Flora Neotropica

Organization for Flora Neotropica
Dicranaceae: Campylopodioideae, Paraleucobryoideae

Author(s): Jan-Peter Frahm
Source: Flora Neotropica, Vol. 54, Dicranaceae: Campylopodioideae, Paraleucobryoideae (Feb. 21,
1991), pp. 1-237
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
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FLORA
NEOTROPIC
MONOGRAPH 54
DICRANACEAE:
CAMPYLOPODIOIDEAE, PARALEUCOBRYOIDEAE
by
Jan-Peter Frahm
Department of Botany
University of Duisburg
Germany
c^s
CANCER
[
Of
\TYROPIOC
FLORAO
NEOTROPICA.
Publishedfor
Organizationfor Flora Neotropica
by
The New York BotanicalGarden
New York
Issued 21 February1991
^^ KîN.,
*9
Copyright ? 1991
The New York Botanical Garden
Published by
The New York Botanical Garden
Bronx, New York 10458
International Standard Serial Number 0071-5794
Library of Congress Cataloging in Publication Data

Flora neotropica. - Monograph no. 1 - New York: Published
for Organization for Flora Neotropica by the New York
Botanical Garden, 1968v.: ill.; 26 cm.
Irregular.
Each issue has distinctive title.
Separately cataloged and classified in LC before monograph no. 40.
ISSN 0071-5794 = Flora neotropica.
works. 2. Botany1. Botany-Latin America-Classification-Collected
works. 3. Botany-Classification-ColTropics-Classification-Collected
lected works. I. Organization for Flora Neotropica. II. New York Botanical
Garden.
QK205.F58
581.98'012-dcl9
85-647083
AACR 2
Library of Congress
ISBN 0-89327-363-5
MARC-S
[8508]
DICRANACEAE:
CAMPYLOPODIOIDEAE,
PARALEUCOBRYOIDEAE
FRAHM'
JAN-PETER
TABLE OF CONTENTS
Introduction.................................................................................
HistoricalAccount.............
...............................................................
Anatomy ....
. . . . ....................................................................
Gametophyte ............................................................................
Leaves .....
......................................................................
Costa . ..... .. ..... ... ......................
.................
...... .................
.
Rhizoids ...........................................................................
Stems ..........................................................
..................
Calyptrae...........................................................................
.........................................
.
Sporophyte .................................
Setae ..........
....... ........
.. ....... .........
.................................
... .
Capsules.............................................................................
Stomata ...........................................................................
Annulus .............................................................................
Peristome...........................................................................
Spores...............................................................................
Cytology ................................
..........................................
Chemistry ...................................................................................
........................
Geography....................................................
Origin and Evolution .........................................................................
Ecology .. ...................
....... ...................
...........................
Substrate ........................................
.....................................
StructuralAdaptations ..........
..........................................................
Water Storage........................................................................
Resistanceto Water Loss ..............................................................
...........
Uptake of Water and Nutrients ................................................
SexualReproduction......................................................................
VegetativeReproduction.................................................................
Systematic Treatment .........................................................................
Campylopodioideae. ......................................................................
1. Atractylocarpus....................................................................
2. Bryohumbertia ....................................................................
3. Campylopodium...................................................................
4. Campylopus .............................
.........................................
5. Dicranodontium...................................................................
6. Microcampylopus ..................................................................
7. Pilopogon.........................................................................203
8. Sphaerothecium...................................................................
Paraleucobryoideae.......................................................................
1. Campylopodiella.............
.................................................
2. Brothera..........................................................................
3. Paraleucobryum...................................................................
Acknowledgments .......................................
...................................
LiteratureCited ..............................................................................
Index to ScientificNames .............
.....................................
...............
2
4
5
5
5
6
9
10
10
10
10
12
12
12
12
14
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22
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24
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25
31
36
37
196
200
216
217
220
224
225
229
229
232
' Departmentof Botany, University of Duisburg,Fachbereich6, Postfach 101629, 4100 Duisburg,Federal

Republicof Germany.
1
~~~~~~~~~~~~2
~~~Flora
Neotropica
ABSTRACT
Frahm,J.-P. (Dept. of Botany,Universityof Duisburg,4100 Duisburg,FederalRepublic
FloraNeotropica54:
of Germany).Dicranaceae:Campylopodioideae,Paraleucobryoideae.
1-238. 1991. The Campylopodioideaeand Paraleucobryoideaeare closely relatedsubfamilies of the Dicranaceae(Musci).Withinthe Dicranaceae,both are characterizedby a broad
costa with a high variationin anatomicalstructures.The Campylopodioideaein the NeoWilliamswith three species,Bryohumbertia
tropicsconsist of eight genera:Atractylocarpus
Portierde la Varde& Theriotwith one, Campylopodium(C. Miiller)Bescherellewith one,
CampylopusBridel with 65, DicranodontiumBruch,Schimper& Giimbel with four, Microcampylopus
(C.Miiller)Fleischerwithone, PilopogonBridelwith six, andSphaerothecium
in the Neotropicsconsist of threegenera:Brothera
with one species.The Paraleucobryoideae
C. Miillerwith one species, CampylopodiellaCardotwith two, and Paraleucobryum(Limpricht)Loeske with two species. All 87 species, as well as infraspecifictaxa are described
and illustrated.
RESUMEN
son subfamiliasestrechamenteafinesa las DiCampylopodioideaey Paraleucobryoideae
cranaceae(Musci). Dentro de las Dicranaceaeambas subfamiliasse caracterizanpor una
costa ancha con una granvariaci6nde estructurasanat6micas.Las Campylopodioideaeen
los Neotr6picosconsistende 8 g6neros:Atractylocarpus
Williamscon 3 especies,BryohumbertiaPortierde la Varde& Theriotcon 1 especie, Campylopodium(C. Miiller)Bescherelle
con 1 especie, CampylopusBridel con 65 especies, DicranodontiumBruch, Schimper&
Giimbel con 4 especies, Microcampylopus(C. Miller) Fleischercon 1 especie, Pilopogon
Bridelcon 6 especies y Sphaerotheciumcon 1 especie. La Paraleucobryoideaeen los Neotr6picos consiste de 3 g6neros:BrotheraC. Miillercon I especie, CampylopodiellaCardot
con 2 especies y Paraleucobryum(Limpricht)Loeskecon 2 especies. Todas las 87 especies
asi como los taxones infraespecificosestan descritose ilustrados.
INTRODUCTION
The Campylopodioideaeand Paraleucobryoideae are subfamilies of the Dicranaceae. This
family of mosses is characterizedby narrowly
lanceolateleaves which aresometimes secundor
falcate with (usually smooth) elongate to quadrate laminal cells which are often differentiated
into basal and upper laminal cells, a single percurrentor excurrentcosta, and the frequentoccurrenceof differentiatedalar cells. The sporophytesareusuallyterminalwith mostly cucullate
calyptraeand cylindricalto ovoid, erector curved
capsules.The plants are erectand can be robust,
formingdense mats, or minute.The Dicranaceae
belong to an evolutionaryline of the superorder
Haplolepideaewith a peristomeconsistingof 16
teeth. The peristome teeth of the Dicranaceae
are entire or divided into 2 (rarely3) prongsand
usuallyare verticallystriateon the outer surface,
an ornamentationwhich is frequentlyfound in
the Dicranales and thereforecalled the "dicranoid" type.
Unfortunately,there are transitions between
the two subfamilies.Certainspecies of Campylopuscannot be distinguishedvegetativelyfrom
species of Paraleucobryum,since they all have a
"leucobryoid"structureof the leaves with ventral and dorsal rows of hyalocysts.Whetherthis
is due to phylogeneticdescentor the consequence
of an independentevolution of this leaf anatomy
is not known.
The Campylopodioideaewere established by
Brotherus(1924). Brotherusincluded 10 genera
in this subfamily:Metzlerella,Microdus,Dicranella, Microcampylopus, Campylopodium,
Campylopodiella, Pilopogon, Campylopus,
Thysanomitrionand Dicranodontium.Of these
genera,Thysanomitrionis now includedin Campylopusas a subgenus,Microdusand Dicranella
Introduction
are placed in a separate subfamily, Dicranelloideae, and Campylopodiellais placed in the
Paraleucobryoideae(Miiller& Frahm, 1987). In
additionthe generaBryohumbertiaand Sphaerothecium are treated here (with species formerly
included in Campylopus).The circumscription
of the Campylopodioideaeused here is similar
to that of Walther(1983). Walther,however,segregated Campylopodiumand Microcampylopus
into different subfamilies (Dicranelloideaeand
Campylopodioideae),although both genera are
very closely relatedand can be regardedas subgenera of one genus (Giese & Frahm, 1985a),
and included Mitrobryumand Maireola in the
Campylopodioideae.The placement of Mitrobryumis not clear since this genus differsfrom
all other Dicranaceaeby its mitrate calyptrae.
Maireolahas been synonymizedwith Ditrichum
(Frahm& Seppelt, 1987). The genus Bryotestua
is not takeninto accounthere becauseit is based
on sterile plants. Another genus, Bartleya, has
been placedinto the synonymyofDicranellacerviculata(Crum& Anderson, 1981).
The systematics of the Campylopodioideae
have been discussed by Frahm (1986) but there
is still no satisfactoryanalysisof this subfamily.
The circumscriptiongiven by Brotherus(1924)
was ill-definedand basedon five characters,three
of which (dioicous sexuality, differentiatedalar
cells, and lack of stomata)were restrictedby the
word "mostly,"a fourthcharacter(differentiated
perichaetialleaves) separatedgenerasuch as DicranellaandAnisotheciuminto differentsubfamilies, generawhich are today usuallyacceptedas
one genus. A fifth character(leaves gradually
thinnertowardsthe margins)is not applicableto
most genera.This confusingassemblageof genera lackedrevisions and monographsto provide
the basic information for generic concepts. A
conspicuouscharactermet in most of these genera is the sinuose seta. Thereforeit is commonly
regardedas characteristicof the subfamily.Only
Pilopogonand Atractylocarpushave straightsetae, but these generaare virtuallygametophytically identical to Campylopusand Dicranodontium. Sinuose setae, however, also occur in
Cynodontium(Dicranaceae), Grimmia (Grimmiaceae) and Campylostelium (Ptychomitriaceae) and have apparently evolved independently several times. This may be the case for
Campylopodiumand Microcampylopus,which
usuallyareplacedin the Campylopodioideaebe-
cause of their sinuose setae and thus their "campylopodioid" appearance,but fit better in the
Dicranelloideaein all other respects.Therefore,
as treatedhere, the circumscriptionof the Campylopodioideaeis probablyartificial.
Whereasmost genera of Campylopodioideae
comprise between 2 and 12 species worldwide,
the genusCampylopushas about 180 speciesand
is thus one of the largestgeneraof mosses.
With the exception of Campylopusand Dicranodontium,all generaof Campylopodioideae
have been revised: Campylopodium(Giese &
Frahm, 1985a), Microcampylopus (Giese &
Frahm, 1985b), Atractylocarpus (Padberg &
Frahm, 1985), Pilopogon(Frahm, 1983a),Bryohumbertia(Frahm, 1982a),and Sphaerothecium
(Frahm, 1986b). Only two subgeneraof Campylopushave been treatedworldwide(Thysanomitrion, Frahm, 1984a, and Campylopidulum,
Frahm, 1986c). For the rest of the genus only
localtreatmentsexist for SouthAmerica(Frahm,
1975a, 1975b, 1977, 1978a,1979a, 1979b, 1979c,
1980a, 1981a, 1981b, 1982c, 1984c, 1986d;
Frahm & Hegewald, 1979; Frahm & Sipman,
1978), Africa (Frahm, 1985a), and Australasia
(Bartlett& Frahm, 1983; Catcheside& Frahm,
1983; Frahm, 1984b, 1987a; Frahm & Mohamed, 1987; Frahmet al., 1985) as well as taxonomic treatmentsof single taxa (Frahm, 1974)
mainly publishedin 14 continuationsof the series "TaxonomischeNotizen zur GattungCampylopus"(Frahm, 1975c, 1976a, 1976b, 1976c,
1978b, 1978c, 1979d, 1980b, 1981c, 1981d,
1982d, 1985b, 1985c).
In these publicationsall characterstatesneeded for classificationhave been criticallyevaluated, the range of expression of these character
statesoverviewed,the delimitationof generadefined and corrected, and many species placed
into synonymy.Worldwide,the numberof species in Campylopushas been reducedfromabout
720 to 180, in Microcampylopusfrom 12 to 2,
in Pilopogonfrom 14 to 8.
The Paraleucobryoideaewere also established
by Brotherus(1924). Brotherusincludedthe genera Paraleucobryumand Brotheraand based the
subfamilyon the broadcosta with a median row
of chlorocysts,differentiatedalarcells, a straight
setaand symmetrical,smooth capsules.Recently
Walther(1983) transferredCampylopodiellato
this subfamily, based on the close relationship
of this genus to Brothera.The entire subfamily
Flora Neotropica
has been monographedby Miller and Frahm to the rankof a genusin 1900, and againreduced
to the rank of a subgenus in 1933. Of the 27
(1987).
species included originally,three were retained
by Giese and Frahm (1985b).
HISTORICALACCOUNT
Pilopogon was introduced in 1826. In 1901
Campylopuswas describedby Bridelin 1819. Brotherusplaced all species of the genus ThysaThe name was chosen to characterizethe curved nomitrionin a subgenusof Pilopogonbecauseof
setae (derived from Greek campylos = curved similarities of the peristome. This caused conand pous = foot). Bridelincludedseveralspecies fusion, because Thysanomitrionwas usually rewith curvedsetae in this genuswhich have noth- gardedas a subgenusof Campylopus.A monoing in common except this kind of seta, for graph of the genus (Frahm 1983a) maintained
example, species laterplaced in Grimmiaor Ra- eight species worldwide,seven of them neotropcomitrium.Species placed presentlyin Campy- ic.
The species of Bryohumbertiawere originally
lopus were mostly described as species of Dicranum, and many authors (e.g., Carl Miiller) included in Campylopus.The genus was based
continuedto use the name Dicranumto describe on an Africanspecies, describedin 1939, which
hundredsof species applicable to Campylopus. was regardedas more or less closely related or
The exploration of the tropics increased the even synonymouswith Campylopus.A re-evalnumberof speciesin this genusdramatically.The uation of its characterstates and additional ulIndex Muscorum(Wijk et al., 1959) lists more trastructuraldifferencesled to the establishment
than 1000 species, about 720 of which were le- of this genus with three species, one each in the
gitimate.
neotropics,tropicalAfricaand SE Asia (Frahm,
The high number of species described (e.g., 1982a).
320 legitimate species cited in the Index MusSphaerotheciumwas introducedin 1865 for a
corumfor LatinAmerica,and a total of 260 spe- species from Colombia. In 1873 anotherspecies
cies describedfrom Africa) prohibitedan over- was addedfrom Sri Lanka.For more thana hunview of this genus.Differenttaxonomicconcepts dred years these two species were the only repused in the last centuryled to the descriptionof resentatives of this genus until a third species,
growthforms as separatespecies, and the lack of from Africa and formerly a member of Camphytogeographicknowledgeand pre-Darwinian pylopus, proved to be a member of this genus
theories on the origin of species resulted in the (Frahm, 1986b).
independent description of species from every
Atractylocarpuswas described by Mitten in
countryand island. The lack of knowledgeof the 1869. Nineteen legitimatespecies were listed in
variability of species and especially the use of the Index Muscorum. Of these nine were acvariable characterstates, such as the transverse cepted in a recentrevision of the genus (Padberg
section of the costa, made this genus a difficult & Frahm, 1985).
one. It enormouslyraisedthe numberof species
The subfamily Paraleucobryoideaewas dedescribed;species which were neitherillustrated fined by Brotherus(1924) to include the genera
nor sufficientlydescribed.
Paraleucobryumand Brothera.Walther (1983)
Duringthe last centurymany new generawere addedCampylopodiella,formerlyincludedin the
establishedor raised from subgenericstatus.
Campylopodioideae,becauseof its structuralafDicranodontiumwas introduced in 1847 for finities to Brothera.
species formerly placed in Dicranum or CamCampylopodiellawas describedfrom Sikkim.
Lateran Africanand a Himalayanspecies were
pylopus.
Campylopodium,originallydescribedas a sec- added. A revision showed the Africanspecies to
tion of Aongstroemiawas raisedto the rankof a be a species of Campylopus,the Himalayanspegenus in 1873 on the basis of its curved setae. cies to be identicalwith the type species, and an
At one time 27 differentspecies were included; andine speciesformerlyregardedas Campylopus
these have been reducedto two (Giese & Frahm, to be a member of Campylopodiella(Frahm,
1985a).
1984c).In furtherstudies(Miiller& Frahm,1987)
Microcampylopuswas originallyestablishedas a third species, again from the Andes, was ina subgenusof Campylopusin 1899. It was raised cluded in Campylopodiella.Recently a fourth
Anatomy
species was added from Papua New Guinea
(Frahmet al., 1988).
Brotherawas originallybased on two nomina
nuda from Asia. Brotherus(1901) reducedthese
names to synonyms of B. leana from North
America,formerlyincludedin Campylopus,and
added a second species. This second species is a
species of Campylopodiella(Frahm, 1984c),and
so the genus is still monotypic.
Paraleucobryumwas firstintroducedby Lindberg (1886) as a subgenusof Dicranum.Loeske
(1907) raisedit to the rankof genus,and included
three holarcticspecies. Subsequentlyother species were transferredto this genus from Dicranum or described as new. All these new combinationsand new species have been reducedby
Miiller and Frahm (1987).
ANATOMY
conversely, display extreme differentiationof

gametangia. Conspicuously, the differentiation
of bud-likeperichaetiais confinedto specieswith
high fertility and it thus can be assumed that
antherozoiddispersalis effective. Specialdifferentiations are found in a group of species with
piliferousleaves suchas Campylopusintroflexus,
C. pilifer, C. julaceus and C. aemulans. Other
species are mostly found fertile and thus give a
comose appearanceas in C. occultus,C. pauper
or C. zygodonticarpus.The highestdevelopment
of such perichaetiais found in Campylopussubg.
Thysanomitrion.Here the perichaetiaare developed on appressedfoliate stalks, especially the
species occurringin SE Asia. This characteris
not developed in the only subantarcticrepresentative of this subgenus(C. clavatus)which is regarded as most primitive, and is hardly developed in the only neotropic representative(C.
richardii).
GAMETOPHYTE
Most generaof Campylopodioideaeand Paraleucobryoideaeare dioicous. Only Atractylocarpus is autoicous and Campylopodiellais paroicous. Conspicuously, these latter genera have
species with small ranges.Dioicism is probably
responsiblefor the broadmorphologicalrangeof
characterstates, especially in the genus Campylopus,that are the resultsof numerousgenetic
variations resulting from out-breeding.Dioicy
also has disadvantages,for example in long distance dispersal,when only one sex is presentin
one habitat and effective methods of vegetative
propagationare requiredto fill the gap between
heterosexualpopulationsand to allow fertilization.
The sex of sterileplantscannotbe determined.
Fertile plants especially of the genera Campylopus and Pilopogon show, in part, a morphological differentiationdue to heterodioicism.In
C.fragilis dwarfmales have been observednesting in female tufts. Fertileplants of a numberof
species producebud-like stem apices consisting
of broaderand blunterleaves. As in many other
mosses this concernsmostly male plants, which
produceflower-likegametangiathat functionfor
dispersalof the spermatozoidsby a splash cup
mechanism.More rarely,femaleplantsalso produce terminal buds which contain several perichaetia. There are also species that lack differentiation, or which display either hardlyany or,
Leaves
The leaves are lanceolateto narrowlylanceolate in shape, graduallynarrowingto a subulate
apex. The marginsare usually entire or serrate
or denticulateonly in the apex. Due to the broad,
often excurrentcosta, the lamina is narrowand
rarelyreachesto the leaf tip, but often vanishes
near midleaf. The lamina is always unistratose.
Laminalcells can be differentiatedinto alarcells,
basal and upperlaminal cells. Alar cells may be
differentiatedor not. They are not developed in
Microcampylopusand Campylopodiumbut differentiatedin all other genera,either weakly or
strongly. Although alar cells can be very conspicuous in some species and hardly developed
in others, this is apparentlynot a specificcharacter.In many speciesthis charactervariesmuch,
obviously controlled by the habitat. Alar cells
function in water uptake from capillary water
along the stem or the tomentum to the leaf.
Therefore,alarcells are usuallylackingin plants
of wet habitats.In Campylopuspiliferspecimens
from rainforestsshow no alar cells since they
receive atmosphericwater taken up by the leaf
surfaces.Specimensfrom dry habitatshave differentiatedalarcells. Alarcells areespeciallywell
differentiatedin plantsgrowingon wet rocksexposed to the sun, which have a high evaporation
rate. In contrast to the older literature,which
used this characterfor differentiationof species,
Flora Neotropica
the presence of alar cells is regardedtoday as

controlledby modificationsand thus usuallyneglected as a useful characterfor separation of
species.
Basal laminal cells differ by their larger size
anddifferentshapefromupperlaminalcells.They
can be either hyaline and thin walled or incrassate. Specieswith hyalinebasallaminalcells usually show a distinct differentiationfrom the incrassateupperlaminalcells.The basalcellsextend
higher up along the margins and are thus
V-shaped in mass. Hyaline basal laminal cells
sheathingthe stem functionfor wateruptakeand
are found especially in species of wet or mesic
habitats. Incrassatelaminal cells can be smooth
or pitted. Basal laminal cells are differentiated
into smaller outer and larger inner ones. The
outer can be narrowand hyaline and forming a
border 2-12 cells wide, or smaller and shorter.
The upperlaminalcells can be quadrate,oblique,
oval or shortlyrectangular.They arelong-rectangularonly in Dicranodontium.
Perichaetialleaves are widened at base and
suddenly contracted to an elongate point. The
broad sheathingbase suggestsa narrowercosta,
which is, however, not narrowerbut as broadas
in normalleaves. The perichaetiathemselvesare
surroundedby anothertype of broad involucral
leaves which are blunt in male plants but show
longly excurrentcostas in female plants.
Special perichaetialleaves are found in Pilopogon, sheathingnearlythe whole seta. This can
be interpretedas another strategyto protect the
young sporophyte.In generaof Campylopodioideae with sinuose setaethe capsulesdevelop protectedbetweenthe comal leaves. This is not possible with straight setae but in this case the
perichaetialleaves take over this function. This
may also cause the longer shape of the capsules
in genera with straight setae in contrast to the
stout capsules of genera with sinuose capsules
which stick between the perichaetialleaves.
Costa
The costa is percurrentor excurrentin all genera treated here. In Campylopusthe excurrent
costae may be concolorous or hyaline, forming
whitish hairtips. These hairpoints can be erect
(e.g., C. pilifer), reflexed (C. introflexus)or recurved(C. griseus)and functionin the reduction
of evaporation. The length of the hairpoint is
determined by modification, since in all these

speciesepiloseplantscan be foundin shadyhabitats.
The costa is the most prominentcharacterof
both subfamilies.In contrastto most othermosses, the costa fills between 1/3 and % of the leaf
width. In this way the usuallyunistratoselamina
with a narrowcosta of most mosses is replaced
here, more or less totally, by a highly differentiated anatomicalstructureprovidingwaterstorage, photosynthesis and mechanical protection
against shrinking.
The costa is composed in most generaof Dicranaceae,except numerous species of Campylopus and all species of Paraleucobryoideae,by
a common dicranoidtype. It consists of (Fig. 1):
1. A ventral chlorophylloseepidermis.
2. A multilayeredband of long, narrowcells,
which are called stereids. Since they resemble
similar cells in the central strand of the stem
which function for water conduction, they are
sometimes also called hydroids.
3. A median layer of enlarged cells, usually
named deuters.The term deuter,of Germanorigin, is merelydescriptiveand meanspointercell;
it refersto conspicuouslylargecells in the median
of a transversesection of a costa. This term has
been used for a median band of enlargedcells in
the Polytrichaceae.In that familythese cells conduct water, and therefore they are also called
hydroids. In the Campylopodioideae,however,
these cells arechlorophyllose("chlorocysts")and
are thus functionallydifferent.Some authorsreplacethe termdeuterby eurycysts(whichmeans,
however,the same and does not regardthe function of these cells), others by "duces"(Latin for
guide cells).
4. A dorsal band of stereids.
5. A dorsal epidermis.
This structureprimarilyprovidesa firmstructure and possible watersupplyby the two bands
of stereidsand assimilation by the ventral, median and dorsal cell layers.
In Campylopusan enormousvariationof this
type of costa is found, which is presumablythe
reason for the rich speciation in this genus. It
allows the species to adapt to a broad spectrum
of ecological niches (Frahm, 1987b). The variation (Fig. 2) concerns:
a. the change, by gradual enlargementfrom
ventral stereids to substereids,small hyalocysts
withfirmwallsandlargehyalocystswithlax walls.
Anatomy
if
AigSfetea^
B^ssillSy
Al
FIG. 1. Transverse sections of leaves in different genera of Campylopodioideae and Paraleucobryoideae. A.

Campylopodiella stenocarpa (Maslin s.n., NY). B. Paraleucobryum longifolium (Crum 1851, DUKE). C. Pilopogon macrocarpus (Allioni 8056, H-BR). D. Atractylocarpus longisetus (Pringle 10603, NY). E. Campylopodium
medium (Steere 6831, MO). F. Microcampylopus curvisetus (Sartorius 60, BM).
b. the change from dorsal groups of stereids

to substereidsor single large cells. The stereids
are sometimes named stenocysts, the large cells
replacinga group of stereids,socii, comites, Begleiter or leptoids. These terms are taken from
the structureof the costa in the Polytrichaceae
in which these cells accompanythe deuter cells
("socii, comites, Begleiter")and function for the

transportof nutrients ("leptoids"),and formtogetherwith the deuter cells-a primitive conduction tissue, with phloem (leptom) and xylem
(hadrom)elements.This is, however,not the case
in Dicranaceaeand thereforean application of
these termsshould be avoided. All possible tran-
Flora Neotropica
1!.
FIG. 3. Transversesection of the costa of Campylopuspilifer (Griffinet al. 92, FLAS). A. The upper
third of the leaf. B. The lower third of the leaf.
FIG. 2. Transversesectionsof differenttypesof

A. Costawithlarge in the upper part and ridged on the basal part
costaein the genusCampylopus.
ventralhyalocystsin Campylopuspittieri(Cleef8806, (Fig. 3) or ridged in the upper part and smooth
in Campylopus in the basal part. Ventral stereids in the
U). B. Costawithventralsubstereids
upper
surinamensis(Mohr1868, NY). C. Costa with ventral
part of the leaf can change to substereidsin the
stereidsin Campylopusuleanus(Ule 148, H-BR).
basal part or substereidsin the upper part can
changeto hyalocystsin the basal part. In humid
sitions are possible from a group of stereidsto a habitats,dorsalstereidscan get largerand change
single largercell by omission of one or more cell to substereids,and dorsallamellaecan get shorter.
divisions (Frahm, 1982e).
In the Paraleucobryoideaethe costa consists
c. the change from smooth dorsal leaves to
ribbed or lamellose surfacesby division of the of a median band of chlorocysts(perhapscomparable to the deuter cells of the Campylopochlorophyllosedorsal epidermalcells.
As shown by the confusion of differentterms dioideae)and ventraland dorsalhyalocysts.The
used for the structureof the costa, there is no medianchlorocystsaresinglein Paraleucobryum
consistent usage. It is proposed to use the de- and Brotherabut divided into 2 to 4 in Camscriptive terms hyalocysts,eurycysts(for deuter pylopodiella.In Paraleucobryuma dorsal band
cells), stenocystsand chlorocysts,althoughin the of chlorocystscan be present.This situationlinks
systematic part the term stenocysts is replaced Paraleucobryumwith certain species of Camby the more common and betterunderstandable pylopus (Frahm, 1982b). Consideringthe basic
term stereids.
sporophyticdifferencesbetweenboth generathis
There is variationin the structureof the costa can be interpretedratheras independentdevelin one leaf and in plantsof differenthabitats.The opment than as phylogenetic linkage. On the othlength of dorsal lamellae decreases to the leaf er hand, Ligrone (1985) has derived the structure
base and the width of ventralhyalocystsincreas- of the leucobryaceousleaf from a dicranoidanes to the leaf base. Thus leaves may be lamellose cestor, which is also hypotheticallypossible and
Anatomy
phylogeneticallyeven more likely, since Leucobryum has the same sporophyte as species of
Dicranum.
The presence of dorsal and ventral bands of
stereids in most other Dicranaceaecan be regardedas the basic structurefromwhich the tendencyto develop hyalocystsseems to be derived.
The presence of ventral and dorsal bands of
stereidsin the highlyadvancedsubgenusThysanomitrion need not be a contradiction.These
species, all except for one, occur in the tropics,
wherethe conservativestructureof the costawith
stereidbandshas advantagesas protectionagainst
shrinkingand for water support.
The ecological significance of the costa of
Campylopushas been discussedby comparisons
of speciespairsdifferingonly in a singlecharacter
in the costal anatomy and a different habitat
(Frahm, 1987b). The differentiationof the costa
concernsseveral characters.
1. Lamelloseand non-lamellosecostae.There
are all possible transitionsbetween costae with
smooth dorsal sides and dorsal sides with lamellae up to six cells high in differentspecies of
this genus. Generally,species with high lamellae
occurin habitatswhichcan easily dryup. Species
with long lamellae can store water and thus extend the photosyntheticallyactive period in dry
habitats. Examples for this adaptation are C.
trachyblepharon(on dry sand in coastal areasof
SE Brazil), C. pilifer on exposed rocks, and C.
lamellinervisin dry caatingaforests. Conspicuously, C. pilifer var. lamellatushas lamellae six
cells high, but it occurs not in drierhabitatsbut
in rainforests.It canbe assumedthatthe lamellae
also allow a better gas exchange in rain forest
habitats with higher humidity and higher temperatures.
2. Ventralhyalocystsand ventral stereids.In
more than half of the species of Campylopusthe
ventral stereids in the transversesection of the
costa of most Dicranaceaeare replacedby ventral hyalocysts.As shown by often narroweradaxial cell walls, these hyalocystsfunctionfor water storage. The size of the hyalocysts varies
between14and %of the leaf width. Specieswith
lax hyalocystsoccur in wet habitats(as in many
paramospecies, e.g., C.jamesonii, C. cavifolius,
and C. nivalis). Species of mesic habitats have
smaller hyalocysts with firm cell walls. In dry
habitats species show ventral stereidsas protection againstshrinking.This is best demonstrated
by the ventral stereids of C. pilifer ssp. galapagensis, which occurs on dry lava flows, and C.
piliferssp. piliferwhich, with ventralhyalocysts,
occurs in less exposed habitats(Frahm, 1987b).
3. Dorsal stereids and dorsal substereids.
Dorsal stereidsoccurin bundlesof 2-4 cells and
functionpresumablyfor mechanicalfixationand
perhaps also water transport.In some species
these stereids are replacedby one nonstereidal
cell (which has a small lumen and thereforeis
usually called substereidal).As in species with
largeventralhyalocystsinstead of ventralstereids, these specieswithout dorsalstereidsare also
characteristicof wet habitats, such as dripping
cliffs or swamps.
Hyalocysts have usually been interpretedas
water storagecells. This is somewhatin contradiction of the fact that species with hyalocysts
often occur in hygric habitats, where this function is not needed. Another possibility (in analogy to Sphagnum) is that they function in the
uptakeof nutrientsin swampyhabitats.Recently
Robinson (1985) added anothertheory. Robinson observed air bubbles in the hyalocysts of
Leucobryaceaeand Calymperaceaefromtropical
lowlandforestsand supposedthat these air bubbles can also function to allow a better gas exchangefor the chlorocystssituatedin the median
of the costa, for chlorocystssurroundedby cells
filled with waterwould have a restrictedgas exchange.The same may be the case for species of
withventralanddorsalbands
Paraleucobryoideae
of hyalocystsand also for some species of Campylopus.However, in the Paraleucobryoideaeas
well as in Campylopus,pores in the surface of
the leaves have not yet been observedas in Leucobryaceaeand Calymperaceae,indicating not
only a structuralbut also a functionaldifference.
Rhizoids
Rhizoids can originatefrom the basal partsof
the stems and the lower dorsal part of the costa.
are
Only in DicranodontiumandAtractylocarpus
rhizoids also borne on the ventral part of the
costa (Crundwell, 1979), showing the close relationship between the two genera. Only stemborne rhizoids are found in Campylopodium
(Crundwell,1979) and Microcampylopus(as in
Dicranella and Microdus)showing in this (and
other respects)connectionof these generato the
Dicranelloideaeratherthan to the Campylopo-
Flora Neotropica
10
dioideae.Thus leaf-bornerhizoidsmay be a good
characterof the Campylopodioideaein a revised
sense. They can be lacking, scatteredor form a
dense tomentum. Since rhizoids function for an
externaltransportof water, low species (such as
species of Sphaerothecium, Campylopodium,
Microcampylopusor Brothera)have usuallyfew
or no rhizoids. Even in largerspecies this characterseems to be controlledby the environment.
It seems thereforeto be as unstableas the presence of alar cells, which cooperatewith the rhizoids in the transportand uptake of water. A
tomentum is found especially in those species
forming dense cushions such as many paramo
species. Sometimes the color of the tomentum
has been used for identification.However, rhizoids are generallyreddish in normal light and
orangebrown in transmittedlight in older parts
by incorporationof phenolic compounds in the
cell walls and whitish in normal light or hyaline
in transmittedlightin younger,outerparts.Thus
a dense tomentummay look whitishas seen from
the outsidebut is reddishinside. Onlythe hyaline
tips of the rhizoids seem to resorbwater.
The structureof the surface of rhizoids has
been successfullyused for differentiationof species in some mosses but has been studied only
in detail for Campylopus(Frahm, 1983b). Rhizoids are usually smooth but a few species with
papillose rhizoid surfaceshave been found.
Recently rhizoid gemmae have been found in
a speciesof CampylopusfromEurope(Arts,1987)
and in one species from the neotropics. They
may, however,have been overlooked,since such
gemmae were not expected in this genus.
pend on such early stages of development. The

calyptraemay be ciliate or entire at base. This
characterseems to be fixed geneticallyand has
been used for differentiationof species in Campylopus but must be used with some caution,
since in Campylopusciliate and non ciliate calyptrashave been found in the same species. It
can be supposedthat in humid habitatsthe tissue
of the calyptramay growon afterseparationfrom
the vaginulaand thus form cilia. In dry habitats
the calyptramay not develop cilia or the cilia are
broken off. Cilia are especially long and conspicuous in Campylopussubg. Thyanomitrion
and subg. Campylopidulum.
SPOROPHYTE
Sporophytesusuallyarise terminallyfrom the
gametophyte. Pseudolateralinsertion has been
found only in a few species of Campylopus,such
as C. shawii and C. controversus.
Setae
The setae are comparableshort. In Sphaerothecium they are only 3-4 mm long and immersed in the perichaetialleaves. Notably, all
three species of Sphaerothecium,found in Colombia, South Africa and Sri Lanka,are known
only from small ranges covering a few square
kilometers.In Microcampylopus
the setaearealso
only 2-6 mm longbut not immersedin the leaves.
In Campylopusthe setae are usually 8, rarely 12
mm long and in Bryohumbertiaand Atractylocarpusthe setae reach a maximum length of 15
mm. The short length of the setae indicates an
origin of the generain open habitats,where the
Stems
capsules are exposed to the wind, and not in
The stems show an internaldifferentiationinto forests or similar habitats, where a short seta
an outer cortical layer with firm, thickened cell would be a disadvantage.
The setae show an interesting twist mechawalls, a median parenchymatictissue and a central stand of small, narrow, elongate cells (Fig. nism. They aretwistedin the drystateand uncoil
when they are wetted even by water vapor. Al4).
thoughthis effecthas been known for more than
a hundredyears,the mechanismfor this torsion
Calyptrae
was not known until recently.Noticeable in all
The calyptraeare cucullatein all genera.This species with coiling setae, the outercortex of the
concerns more or less ripe capsules. In young setae has strongasymmetricthickenings.Recent
sporophytesthe capsule is fully covered by the SEM and TEM studies (Frey & Frahm, 1987)
calyptrawhich is symmetricat this stageand not have revealedthattherearegroupsof smallpores
split. Indicationsof mitratecalyptraefor genera 80 A in diameterin the primaryand secondary
such as Brotherain the literatureprobably de- cell walls of the epidermislayer.Wateror water
Anatomy
11
''
-.
?:
'.'.
FIG. 4. Transversesections of stems in differentgeneraof Campylopodioideaeand Paraleucobryoideae.

A.
Pilopogonmacrocarpus(Allioni8056, H-BR). B. Atractylocarpus
longisetus(Pringle10603, NY). C. Microcampylopuscurvisetus(Sartorius60, BM). D. Campylopusrichardii(Griffinet al. 1035, FLAS).E. Campylopodiella
stenocarpa(Maslins.n., NY). F. Campylopodiummedium(Steere6831, MO).
vapor can pass through them and be absorbed

by pectine in the microfibrilsof the incrassate
tertiarycell walls. These microfibrils,coiled in
the drystate,enlarge,causingan uncoilingmovement of the setae. The setae are straightin some
genera (Atractylocarpus,Pilopogon, Paraleucobryum)but sinuosein all othergenera.This shape
originateswhen the seta firstgrowsupwardsand
thencurvesdownwards.In this mannerthe young

sporophytebecomes situated between the perichaetial leaves, where the capsule develops protected against desiccation. When the capsule is
ripe the seta grows upward again, producinga
second curve and thus a sigmoid shape to the
seta. In this way the calyptra is frequentlyleft
between the perichaetial leaves. The sigmoid
12
Flora Neotropica
shape of the seta causes wide movements of the

Annulus
capsule when the seta is uncoiling.In these genAn annulus is present in the Paraleucobryoiera the setae are twisteddextrorselyin the upper
in Brotheraand in Campylopodiellastenodeae
but
in
the
lower
which
part
sinistrorsely
part
prevents the setae from being torn off by the carpa. It is apparentlylacking in the Campylopodioideae, although in Campylopusthere are
uncoiling movements.
sometimes indications of an annulus found in
the literature.Thisseems, however,to referto an
Capsules
annulus which is visible microscopicallyas difThe capsules are 1.5-2 mm long in the Cam- ferentiatedcells but not dehiscent.
pylopodioideae. They are globose (Sphaerothecium, Campylopussubg.Campylopidulum),long
Peristome
or ovoid
cylindrical(Pilopogon,Atractylocarpus),
The peristome consists of 16 teeth which are
to short-cylindrical,
especiallywhen empty. They
can be straight and symmetric or curved and usually split to half (or more) their length. Only
asymmetric,the latteroften being strumose(Fig. in the two speciesofPilopogonsubg.Thysanomi5). The color varies between yellowish in young triopsisand in the monotypicgenusBrotheraare
capsules to dark brown on old capsules. When the peristome teeth not divided. There are two
emptied, the capsulesarefurrowedand contract- types of peristometeeth: the so called dicranoid
ed below the mouth. In the Paraleucobryoideae type consistingof elongatetriangularteeth,which
capsules are cylindrical.Seta length and capsule is common in most genera, and another type
orientation are important for spore dispersal. consistingof narrow,filiformteeth (Fig. 6). The
Conspicuouslycylindriccapsules are correlated abaxialsurfaceof the dicranoidperistomeshows
with long, erect, nonsinuose setae, as in Atrac- longitudinalstriae,the adaxialsurfacehas transtylocarpusand Pilopogon. Here the spores are verse ribs, usually densely covered by papillae
released by vibrations of the setae by wind. In (Fig. 7, except for Bryohumbertia).In the dry
generawhich have the sophisticatedtwist mech- state the peristome is closed because the inner
anism combined with sinuose setae, the spores transverseribs,formingU-shapedstructures,are
are shed by spiral movements of the setae. In driedup and bent inwards.When moistened(faCampylopusboth symmetric,uprightand asym- cilitatedby the papillae)the tensionbetweenthese
metric, curved capsules are found. Species with ribs is lowered,the distancesbetween these ribs
conspicuouslyuprightcapsules usually occur in are enlargedby absorption of water vapor and
open habitats, whereas those with curved cap- the peristometeethmove outwards.By thismechanism spore dispersalis possible in moist consules grow in forests.
In the Paraleucobryoideaethe capsulesare al- ditions. At the same time the setae uncoil, reways erect, shortly to longly cylindric, and 1.5 sultingin despiralizingmovements or, in genera
with sinuose setae, resulting in wide circular
mm to 2.5 mm long.
The operculumis about half as long as the urn movements. Due to the moist condition when
and obliquelyrostratein most generaand longer the sporesare releasedthey are probablynot disin Bryohumbertia.It is usually darker colored persed far. This has some importancefor these
than the urn.
generathataredioicousbecausefemaleand male
spores must fall nearby to allow fertilizationin
the mature plants. Filiform peristome teeth ocStomata
cur in Campylopussubg. Thysanomitrionand in
Stomata at the base of the capsules are found Pilopogon.Both generashow strongresemblance
in Campylopodium(Campylopodioideae)and also in gametophyticcharacters,but are distinParaleucobryum(Paraleucobryoideae,although guished by sinuose setae in Thysanomitrionbut
there are no stomata indicatedin the literature), straightsetaein Pilopogon(whichshows that this
conspicuously only in these representativesof characteris perhapsovervalued).Here the peridifferent subgenera,but not in closely related stome teeth are longer,ending in slender,round
genera. They are cryptoporousin Campylopo- apices which are papillose. The broader basal
dium but phaneroporousin Paraleucobryum.
partof the teeth (lower1/4)has the same structure
13
Anatomy
<
A/
fI"//
ft
B(B
fl
"
;
Wi
V i
if-
FIG. 5. Shape of capsules in different stages of development in the genus Campylopus sect. Homalocarpus
(A-D) and sect. Campylopus (E-H). A. C. nivalis (Sharp 3043, TENN). B. C. areodictyon (Griffin 356, FLAS).
C. C. argyrocaulon (Hegewald 9171, hb. Frahm). D. C. occultus (Puiggari 331, H-BR). E. C. arctocarpus (Vital
4264, SP). F. C. pauper (Lindig s.n., H-BR). G. C. chrysodictyon (=pauper) (Lindig s.n., NY). H. C. concolor
(Lindig s.n., NY).
as in the common dicranoidtype. In addition to

the openingmechanismdescribedabove, the long
apices of the peristome teeth are twisted when
in the dry state but uncoil when moistened (as
in the pottiaceous type of peristome). In both

groups only the one representativeof Pilopogon
in Africaand the only representativeof Thysanomitrion in the subantarctic(which are regarded
14
Flora Neotropica
11'
"
or
o?
VAaz~~~~~~~~~~isi
'It,
n.y
FIG. 6. Types of peristomesin Campylopodioideae.A. Entireperistometeeth in Pilopogonlaevis (Lindig
s.n., H-BR). B. Bifid peristometeeth in Microcampylopuscurvisetus(Sartorius60, BM).
as the most primitive species) have the normal

dicranoid type of peristome. It can thus be assumed that the species of Pilopogon in South
Americadevelopedafterseparationof the Gondwana continent, caused by changing ecological
conditions stemming from the uplift of the Andes, while Pilopogon africanus remained unchanged in Africa. In this regardthe neotropic
type of peristome would be the most advanced.
The same can be assumed for Thysanomitrion,
in which species evolved from the present subantarctic ancestor and extended to the tropics
with a special speciation in SE Asia.
Spores
The spores are ca. 10-19 um in diameter in
most genera and are thereforebest adapted for
long distancedispersal.Only in Campylopodium
and Microcampylopusare they 18-24 Atm,ca. 21
Lmin Sphaerotheciumand 19-34 umin Paraleucobryum.In the numerous species of Campylopus and species of Bryohumbertiaand Pilopogon
spore size is remarkablyuniform at ca. 13 Am.
The spore ornamentation is not yet known by

SEM studies for all genera. To the extent that the
genera have been studied, it varies between nearly smooth to finely or coarsely papillose in Campylopus to warty in Microcampylopus and Campylopodium (Figs. 8, 9). This does permit
separation of subgenera and sections in Campylopus (insofar as this can be generalized from
the comparatively few species which have been
studied) and even species in Microcampylopus
and Campylopodium.
CYTOLOGY
Chromosome numbers are known for only 18%
of the mosses, mainly for species of Japan, Europe, North America, India, Australia or Antarctica. Very little is known about tropical mosses. According to Fritsch (1982) there are no counts
for Atractylocarpus, Pilopogon, Sphaerothecium
or Bryohumbertia. For Campylopodium n = 13
+ m and 35 + 1 are given, but the count of the
first species belongs to Microcampylopus after a
Cytology
15
. ./
!?
i>
,~~~~~
>
,:.. '~./,~...
.%/,
'.?._,
/.
tz,,~7
I~
"'
~ ; ;~,
,;t
c
__Xs~~
III'
'
tL'^'.
'
-[~~~~~~
?Il~~~nii,,
S_;;li'g,
'
?r ;
'
i".
'
/'.
(?.
z~
_;*,~~j~
;~~~
_ (Ge :
occultus
tM
f' ~
ocutu Ghr
FIG. 7.
~.
/
'if:
?f
*e
'~
'
i'X
-1
1. t hy+"~~. 1F
< il
~~~~~~~~~~~~~~~~~~~~~~~~~
l~~"?;0?
NY).'
Ba
Bb'je r _
logseu
S{
Si
5, J)D.Arcyoaps
a~IB
w#
--
?D
jt.
_
A.
^~~~~~~~~~~~~~~~~~~~~~~~- ~ < ~ - ~ ~ ~~
tt D. Atractylocarpus
t~~~~
2, JE).
10603,
longisetus (Pringle
hs
'i
_,,_.st1
;r
'
t,
_I?s, _
j
! t
~~~~~~~~~Ir
_=~~
fii
?-?/
__
A-
, ~;r?
'i~J ,' 'tf.'/X
_?
(Pinl
1063
n,
Y)
SEM pictures oflpenstomes. A, B. Bryvohumbertiafilifolia (Frahm 1555, hb. Fralm). C. Campylopus
revision of the genera (Giese & Frahm, 1985a,

1985b) and the second count concernsa species
of Dicranella. This shows that chromosome
countsare highlydubiousfor generawhich have
not been revised or are from species which have
been erroneously identified, which again concerns mostly tropicalspecies. In the one species
of Dicranodontiumstudiedcytologically,n = 11-
12 arefound in differentpartsof the range.Chromosomenumbersin Campylopusareknownfrom

14 species (=approx. 7% of the actual number
of species, Frahm, 1983b), and one of them is
now transferredto Bryohumbertia.None of these
counts is from neotropicalmaterial.This is partly due to methodological difficulties.Only two
of these counts were taken from mitosis, all oth-
Flora Neotropica
16
'.
FIGs
s(
l1
,N
. B.
FIG. 8. Spores ofCampylopodioideae. A Atractylocarpusl

1ongisetus(Pringel 10603, NY). B. Campylopodium
medium (Steere 6831, MO). C. Campylopus caroinae(Yital 1714, SP). D. Microcampylopus curvisetus (Sartorius
60, BM).
ers from meiosis, which are more easily obtainable. However, only a few of the species of Campylopusproducesporophytesconsistentlyand the
sporophytes of numerous species are not even
known. The basic numbers are 10, 11, 12, 13
and 18. Chromosome numbers of 10-14 are
commonly regardedas basically diploid (Newton, 1984) and thus all species of Campylopus
can be regardedas diploid, two of them with n
= 18 also triploid. Although most counts are
taken from one population only and not from
differentpopulations, especially in species with
transcontinentalranges,the resultspresentedare
too similar to expect surprisesin additional results. This is not differentfrom most Dicranales
in which n = 13, 14 are found, probablypolyploids of n = 7. In the Paraleucobryoideaen =
12 is given for Brothera leana. For the genus
Campylopodiellan = 14 + m is reportedbased
on one count in one species and n = 12 or 14
for Paraleucobryumlongifolium.As in the Campylopodioideaeall countsfallinto the same range
of the Dicranales.
CHEMISTRY
Flavonoid patternsof several species of Campylopus (Frahm, 1983b), Pilopogon, Campylopodium and Microcampylopus(Frahm,unpubl.)
have been studied. In Campylopusdifferentflavonoid patterns allow separation of subgenera
and sections. Surprisingly, in two species of
Campylopus(C. albidovirens,C. pittierifrom the
neotropics)no flavonoidshave been found. Both
species vegetatively much resemble the genus
Paraleucobryum,which also has no flavonoids
(Muller & Frahm, 1987) and seems to indicate
a phylogenetic connection between these two
genera of differentsubfamilies. However, Leucobryum, a genus with a similar anatomical
structureof the costa, also has no flavonoids(Huneck, 1983), although the genera may not be
closely related (Loeske, 1907). Other chemical
constituentsof the Campylopodioideaeand Paraleucobryoideae have not been studied very
much. In Campylopusintroflexussteroids such
as campesterol have been found as well as the
17
Geography
_(* W**
' "'a
'~
I?
.fiC.
i~Ia
~~~P
....
'~~~~~~~~~~LI
r--3C_L~~~~~~~~~~~~~~~~~~~
'74~ ~~f "~~~~~~4~~~~,"~~
$;
4L
I,
L ~'l,.
FI.
Fram)
cutu
Crnylpu
B
Atatyoaru lni
9
iexou (Ght32
ts(ri s 100,N)
n
(H E
E.C
eir
ho k
rohmetaflfla(rhr
s.n.
orpsopoefCmylpdoda.A.Cmyou
hb.
55 a.Fam.D
A. Campylopusflexuosus (Hakeliers.n., hb.

of sporesof Campylopodioideae.
FIG.9. SEMphotographs
B. Campylopusoccultus (Gehrt 352, JE). C. Bryohumbertiafilifnolia(Frahm 1555, hb. Frahm). D.
Frahm).
(Pringle10603, NY).
longnisetus
Atractylocarpus
triterpenoidhop-22(29)-ene(Huneck, 1983), but

these few resultshave no systematicsignificance.
In many species of Campylopus,Dicranodontium, Bryohumbertiaand Pilopogon, lipids can
be observedin the cells of the lamina and costa.
This effect is visible only in species with firm
basal laminal cell walls and not in those with
hyaline, thin cell walls (and thus not in species
of Paraleucobryoideae).Tropical species have
firmcell wallsin the lowerpartof the leaf, whereas subantarcticand andine species have hyaline
basal laminal cells. Since lipids function as reserve substancesit may be supposed that these
lipids function for balancingthe energy loss by
respirationwhich is especially critical for ecological conditions of the tropics.
GEOGRAPHY
Rangeextent differsconsiderablyin the genera
of the Campylopodioideaeand Paraleucobryoideae. It is smallestin the genus Sphaerothecium,
in which all three species are known only from
a few recordswithin a few squarekilometers.In
contrast, the range of the genus Campylopus
comprises nearlythe whole world between 65?S
and 70?N latitude and an altitudinal range between sea level and 4800 m. This extreme dif-
18
Flora Neotropica
ferenceis mainly due to differentstructuraland

ecological adaptations,which are treatedin detail in the chapterson anatomy and ecology.
Pilopogonis representedin tropicalAfricawith
one species, in Centraland South America with
seven. Only one of these species ranges widely
throughCentralAmerica, the Caribbeanand to
Brazil;all other species are confinedto largeror
smallerparts of the Andes (which can therefore
be regardedas a centreof radiationof this genus),
eitherby geographicalisolation (as in P. schilleri
from Chile) or habitat diversity (as in P. tiquipayae or P. macrocarpus).
Dicranodontiumis a mainly holarctic genus.
Only a few species are found in the tropical
mountainsof SouthAmerica,Africaand SEAsia,
probablyderived from holarcticancestors,since
these species are mostly confinedto the northern
partsof the tropicsand rarelycross the Equator.
Bryohumbertiais representedwith one species
each in the neotropics, tropical Africa and SE
Asia. All threespeciesareapparentlyvery closely
related, the taxa in Africa and SE Asia being
more closely relatedthan either of these is with
the neotropic species.
Microcampylopusis also representedin the
tropics with one species each in the neotropics,
Africa and SE Asia. The African species occurs
also in SE Asia.
Campylopodiumcomprises two species, one
with a wide circumpacificdistributionfrom Japan to New Zealandand Chile,the otherendemic to New Zealandand Tasmania.
Atractylocarpuscomprisesnine closely related
species worldwide. All species are confined to
alpine habitats. There is one species each in all
mountainmassifsof the worldand thus no overlapping ranges. The ranges vary much in size,
betweenEurasianin A. alpinusand endemic species, like A. madagascariensis.
Paraleucobryumis again a holarctic genus
comprisingthreespeciesof which one (P. enerve)
goes down to CentralAmerica. Only one taxon
(P. longifolium ssp. brasiliense) occurs in the
tropics and is endemic to SE Brazil.
Brotherais monotypicand has a disjunctrange
betweenEastAsia and SE of North Americaand
Mexico, probablyas a relict of a formeramphioceanic rangein the Tertiary.
Campylopodiellashows a similar disjunction,
but at the generic level, with one species in the
Himalayasand two in CentralandnorthernSouth
America. The neotropicalspecies are separated

by a differentaltitudinalrangeand have probably
evolved from the same ancestorin the course of
the Andean folding.
Most speciesare found in subtropical,tropical
montane or tropical alpine regions and do not
occurin the lowlandsof equatoriallatitudes.Only
a few speciesof the betteradaptedand (according
to the numberof species)most successfulgenus,
Campylopus,can survive here. These species (C.
surinamensis, C. savannarum)are confined to
open, lighthabitatssuch as sandyshoresof rivers
or sandy soil in heath forests.This effectis probably caused by physiological problems. In experiments,tropicalmontanespeciesdid not reach
a sufficient net photosynthesis under lowland
conditions with high temperaturesand low light
intensity (Frahm, 1987c, 1987d). Only taxa that
are physiologicallyspecially adapted seem able
to grow in the understoryof the equatorialrainforest. A higher light intensity in open habitats
allows the species mentioned above to compensate for the high rates of respirationin part. It
remains an open question why those species of
Campylopusfoundin the Amazon lowlandoccur
only on sandy and not on lateriticsoil.
In Campylopusall types of distribution,from
tropical disjunct species to endemic taxa, are
found. There are only a few species occurring
throughout the tropics. If ranges cover South
America, Africa and Asia (as in C. pilifer or C.
savannarum),the distributionin Asia is confined
to Indiaand Sri Lanka.Such speciesarereplaced
in SE Asia by vicariant species, which demonstrates that isolation of SE Asia by continental
drifthappenedearlierthan the isolation of India
from Africa or Africa and South America.
There are several phytogeographicalrelations
to Africa. Several species are disjunct between
Africaand South America.Inasmuchas they are
either lowland species (Campylopussavannarum), montane (C. flexuosus, C. fragilis), or alpine species (C. nivalis, C. incacorralis,Frahm,
1982b) and also species which produce spores
either frequentlyor extremely rarely, seems to
indicate that both long distance dispersal (especially of fertile alpine species) as well as separation of populations by continental drift (of
sterilelowlandspecies)can be consideredas reasons for these disjunctions.It highlightsthe fact
that at least the lowland species may be older
than the separationof SouthAmericaand Africa
19
Geography
at the end of the Mesozoic. Comparingthe numberof speciesin both continents,Africahas, with
50 species of Campylopus,fewer species than
South America, with 65 species. This is partly
due to a richer speciation of this genus in the
Andes. Although the isolated mountain massifs
in Africaseem to supportendemismby isolation,
this is not the case in Campylopus.In contrast,
this situationin Africa seemed to interruptpossible migrationroutes and caused small ranges
of species. Air currentsin the tropics generally
go fromeastto west (vanZanten1983)and therefore the origin of the alpine species occurringin
Africa and South America should be African.
This is difficult to imagine, since these alpine
species have apparentlyevolved from subantarcticancestorsand had a betterpathwayto the
tropical mountains in South America through
the continuous Andes ratherthan, as in Africa,
by hopping from one mountain to the other.
The phytogeographicalconnection of species
of Campylopusin South America and Africa is
also expressed by subspecies and species pairs
(Frahm, 1988). For example, C. julaceus and C.
are both representedin SE Bratrachyblepharon
zil and in SE Africaby subspeciesdifferingonly
in the height of the dorsal lamellae of the costa
or the presenceof ventral hyalocystsor stereids
in the transverse section of the costa (Frahm,
1985a). Both subspecies occur not only at the
same latitudeon the east coasts of the respective
continents, but also in the same habitatsof nutrient poor sand near sea level. The same disjunctionbetweenSEBraziland Malagasyis found
in the hepaticgenus Bryopterisand the moss genus Phyllogonium.A similareffectis that of species pairs occurringin both continentsas Campylopus sehnemii and C. controversusin South
America and C. cataractilisand C. stenopelma
in South Africa(althoughthere is principallyno
differencesince the differentiationbetween subspeciesand speciespairsis surelyproblematical).
Both species pairs are so closely related that a
common originof both speciesis most probable.
It can be supposedthat both species are derived
from subantarcticancestors,which are still extant (C. incrassatusfor C. sehnemii/cataractilis
and C. purpureocaulisfor C. controversus/stenopelma). After continental separation,both species may have migratednorthwardsto the subtropicaland tropicalregionsand in this way have
developed their own characters.
Disjunctions between Central America and

Asia are illustrated by Campylopusjaponicus
(Mexico-Japan)and Brotheraleana (EastAsiaSE North and CentralAmerica). This is a type
of disjunction found more frequentlyin other
bryophytesas a resultof a previouscircumpacific
range in the Tertiary. The genus Campylopodiella is also disjunctbetweenAsia and the neotropics with C. himalayana and C. crenulatain
the Himalayas and New Guinea and C. stenocarpaand C.flagellacea in CentralAmericaand
the Andes.
A circum-Tethyanrangeis found in Campylopus oerstedianus,occurringin Costa Rica, Jamaica, Georgiaand again in scatteredlocalities
in southern Europe. This indicates a Mesozoic
age of this species, the scatteredpresent distribution being a result of the lack of sporophytes
and probably also unfavorableecological conditions.
A disjunctionbetweenNorthandSouthAmerica is found in Campylopuscarolinae which is
found in the Cerradoregions of Braziland also
in the alluvial coastal plains of Florida,Georgia
and the Carolinas.In both regionsabsolutelythe
same habitat is occupied (white sand in often
burned vegetation in which the minute plants
areburied).Campylopussurinamensisalso grows
on white sand and shows a similar distribution
patternbut occursalso in betweenin the Amazon
lowland. Campylopusangustiretisis disjunctbetween SE Braziland the Caribbeanand is found
in the same vegetationtypes as the previousspecies, but in swampyplaces. All three speciescan
be regardedas relictsof a formercontinuousrange
of a dry vegetation type linking Brazil, the Caribbeanand the SE of North America. In con(which
trast,the occurrenceof C. trachyblepharon
grows in coastal sand and is mainly distributed
in SE Brazil) in N Brazil and the Bermudasis
interpretedratheras a resultof dispersalby birds.
Only few species show pan-neotropicranges
and are found from CentralAmerica to Brazil
(but usually with the exception of the Amazon
lowland).This concernsMicrocampylopuscurvisetus,Pilopogongracilis,Bryohumbertiafilifolia,
Campylopuslamellinervisand C. subcuspidatus
(except for the Andes), C. richardii,and C. arctocarpusand in addition the lowland and montane species with an African-South American
disjunctionsuch as Campylopusfragilis, C.flexuosus,C. savannarum,and C. pilifer.Also, Bryo-
20
Flora Neotropica
humbertiafilifolia and Campylopusarctocarpus

have close relationsto Africa,as indicatedby the
fact that B. filifolia is replaced in Africa by a
closely relatedspecies,B. metzlerelloides,and C.
arctocarpusis replacedby a vicariant race, ssp.
madecassus.
WithinSouthAmericadisjunctionscan be observed between the Andes and SE Brazil.There
are several categories of species: species which
show no differences between the populations
(Atractylocarpuslongisetus, Campylopuscuspidatus, C. heterostachys,C. jamesonii, C. densicoma, C. reflexisetus);populations, as in Pilopogon gracilis,in which differencescan be stated
numerically,but which, however, overlap and
do not allow separationof these populationstaxonomically (Frahm, 1983a); subspecies, as in
Campylopusfragilis subsp.fragilis in the Andes
and subsp. fragiliformis in SE Brazil; or geographicalvicariantspecies such as Atractylocarpus brasiliensisand A. longisetus or A. nanus.
These disjunctions can be explained either by
long distancedispersalor as relicts of a formerly
closed rangethat includedthe Andes and the SE
Brazilianmountains in a presumablycooler climatic period.
Extensionsof boreal rangesto CentralAmerica, rarelyto South America, are found only in
Paraleucobryum enerve and Dicranodontium
denudatum.
DisjunctionsbetweenCentralAmericaand SE
North America are demonstratedby Brothera
leana and Campylopustallulensis.
Campylopusshawii is disjunct in the Caribbean, the Azores and the British Isles.
Several species are endemic to SE Brazil, for
example Campylopus cryptopodioides,C. dichrostis, C. gemmatus, C. julicaulis, C. uleanus
and C. viridatus.Some Campylopusspecies (C.
aemulans,C.julaceus, C. griseusand C. occultus)
are also found on or proceedto the slopes of the
Andes in N Argentinaand S Bolivia.
Campylopusgardneri,C. gastro-alarisand C.
widgreniiare endemic to the arid parts of S and
NE Brazil.
There are numerous species which are confined to the Andes and which have probably
evolved in consequence of the uplift of these
mountainsat the end of Tertiary.This makesthe
Andes the region with the highest number of
species in the world for Campylopusand Pilopogon. Accordingto the humiditygradientin the
Andes from the Equator north and south the

ranges of these species are very different.The
widest rangesfrom Mexico to Bolivia (rarelyto
northern Argentina)are found in Campylopus
albidovirens,C. anderssonii,C. concolor,C. oblongus, C. pittieri, C. sharpii and C. zygodonticarpus. Species confined to the region between
Costa Rica and Peru or northern Bolivia are
Campylopusasperifolius,C. cavifolius,C. huallagensis, C. trivialisand Pilopogon laevis. Only
foundbetweenVenezuelaand Bolivia (andoften
confinedto even smallerrangessuch as Colombia to Peruor Colombiaand Venezuela)and not
in Central America are Atractylocarpusnanus,
Campylopusamboroensis,C. areodictyon,C. argyrocaulon,C. bryotropii,C. capitulatus,C. cleefii, C. edithae, C. incertus,C. jugorum, C. longicellularis,C. luteus,C.perexilis,C. subjugorum,
C. trichophylloides,Pilopogonperuvianusand P.
macrocarpus.These are predominantlyalpine
species. Some species have even smallerranges,
e.g., confined to Bolivia (P. tiquipayae)or Colombia (Sphaerothecium
phascoideum).The only
species confinedto CentralAmerica seems to be
Dicranodontiummeridionale,but this genus has
not yet been monographed and therefore any
phytogeographicalinterpretationsare doubtful.
A phytogeographicaldiscussionfor the species
of Campylopusand Bryohumbertiafromthe Roraima massif is given by Frahm and Gradstein
(1987).
Campylopuscygneusand C. cubensisare confined to the Caribbean,the latter is also found
in the surroundingpartsof the continentin Central and northernSouth America, perhapsas a
result of secondaryspreading.
ORIGIN AND EVOLUTION

Because of the lack of any fossils from either
subfamily, only speculations can be given derived from interpretationof the present ranges
of species and genera.
Atractylocarpusis represented in nearly all
mountainmassifsof the world,each with a single
species. Since these mountains are predominantlyof TertiaryAge, speciationwithin this genus has happened presumablyonly during the
last 50 million years. Atractylocarpusis mainly
tropical and alpine in distribution.It is closely
relatedto Dicranodontium,which is boreal and
montane. The anatomical differences concern
Originand Evolution21
mainlythe sinuoseor erectsetae.A circumboreal
distribution can go back to a Laurasianorigin
and thus it can be assumedthat Dicranodontium
is older than Atractylocarpusand that the latter
has evolved from Dicranodontiumby occupying
new niches in alpine regions and has differentiated into several species on isolated mountain
massifs.
In Campylopusit is notable that there are 15
(revised from 60 described)species in the subantarctic but only one in the subarctic.Moreover, (except for one) only species with hyaline
thin walled basal laminal cells occur in the subantarctic,and the most primitive representative
of the subgenusThysanomitrion
also occursthere.
So it can be assumed that this genus evolved in
the southern part of the Gondwana continent.
Presumably,a rich speciation took place (1) by
isolation caused by continental drift, (2) by
spreading northwardsin the continents under
more favorableclimatic conditions, which were
xeric in the precedingMesozoic, (3) by adaptation to new habitatsin savannasand rainforests
and (4) by speciationin new mountain systems.
For several groups of species, pathwayscan be
constructed illustrating these mechanisms
(Frahm, 1988).The existenceof a circumtethyan
relict (a conspicuouslydry-adaptedspecies with
hairpoints)indicates that representativesof this
genus were also present at the northernpart of
the Gondwanacontinent.
As shownby bud-likeperichaetiaon appressed
foliate stalks and a differenttype of peristome,
found also in the youngest species of Pilopogon
and in certainspeciesof the subg. Thysanomitrion, (especiallythose occurringin SE Asia), the
subg. Thysanomitrionmay representthe youngest branchof evolution in this genus.
The close relationship between Campylopus
and Dicranodontiumleads to the suspicion that
both genera (one in Gondwana, the other in
Laurasia)have a common ancestor. Nearly all
species of Dicranodontiumcan be distinguished
fromcertainspeciesof Campylopusonly by elongate upper laminal cells. In contrastto Campylopus,Dicranodontiumhas never developed the
largevariety of differentstructuresof the costa.
It thereforehas remainedconservativeand less
flexible.Not able to adaptto differentnew habitats, the genus was not able to participatein an
enormousspeciationas was Campylopus.Therefore it remainedconfinedto its formerrangeand
did not spreadsubstantiallyinto the tropics. In

a little knownbryofloristicpaperon the Austrian
Alps, Loeske (1910) discussed relationshipsbetween generaof Dicranaceae.Basedon the similaritiesin the structureof the costa and the lamina with Ditrichum,Loeske derived Campylopus
and Dicranodontiumfrom Ditrichaceae.Paraleucobryumbelongs, accordingto Loeske,to the
same evolutionaryline. For that reason Loeske
proposedto separateCampylopus,Dicranodontium and perhapsalso Paraleucobryumfrom the
Dicranaceaeand to establisha new family,Campylopodaceae.
Pilopogon also resembles certain species of
Campylopusand can be distinguishedonly by
the straightseta and the long perichaetialleaves.
Six of the seven species occurin the Neotropics,
five of them exclusively in the Andes. As indicated by the dicranoidtype of the peristome in
the only Africanspecies and the advanced type
of peristomein the neotropicalspecies,the genus
originatedin the Mesozoic beforethe split of the
Gondwana continent but furtherevolved later
in the Andes.
Bryohumbertiais againcloselyrelatedto Campylopus,differingby longersetae, a longeroperculum and smooth peristome teeth. Three species are distinguishedworldwide,one in each of
the main tropical regions. The differencesbetweenthese speciesareso smallthatcertainsmall
specimens can hardly be referredto one or the
otherof these species,whereassome largegrowth
forms are confinedto the neotropics.This close
relationship indicates a common origin and a
recent differentiationafter the split of the continents which may have not yet reacheda separationinto fully separatespecies.
The three species of Spaherotheciumfound in
Colombia,SouthAfricaand Sri Lankaalso show
only smalldifferences.The nearlyworldwidedistributionindicatesan olderageforthe genusthan
Bryohumbertia,with a late or post Mesozoicspeciation caused by isolation. The small rangesof
all these speciesindicatethat the presentecological conditionsarenot best for these species.Since
these species grow on bare soil and not in sheltered habitats, it may be speculatedthat these
species (two of which are known only from the
type collections made 130 years ago, a third
knownfromonly halfa dozen collections)cannot
survive in such small populations and may be
extinct in the future,if not now.
Flora Neotropica
22
Microcampylopusand Campylopodiumdiffer
mainly in the ornamentationof the spores and
the presence or absence of stomata in the neck
of the capsules.Consequently,both generahave
a relation similar to that of Dicranella(stomata
absent) and Anisothecium (stomata present),
which are often united. A common origin for
both genera can be assumed, perhaps derived
fromAnisotheciumby developmentof the twisted and sinuose seta of the Campylopus-typeor
by sharingthe same ancestorwith Dicranellaand
Anisothecium.Microcampylopusis distributed
pantropically,with one species each in South
America, Africa and SE Asia. These species
probablyevolved afterthe separationof the continents and, because of their close relationship
(the differencesconcernonly length/widthof the
capsule and the spore ornamentation),are descendants of a Mesozoic ancestor. Campylopodiumis SEAsianin distribution,with two species
(the occurrenceof C. medium in Chile is interpreted as a Pacific extension of this range),and
thereforemighthave developedfromMicrocampylopusin this region, or is older, with formerly
vicariantspecies in other partsof the tropicsextinct. A third possibility might be that the Chilean and New Zealand occurrencesare relicts of
a gondwanalandicrange and the occurrenceof
C. mediumin SE Asia is a secondaryextension.
ECOLOGY
SUBSTRATE
Most speciesof Campylopodioideaeand Paraleucobryoideae grow on soil, rocks, decaying
wood or tree bases. This is true for all genera
except Campylopusand Pilopogon. Only a few
species of the lattergenera,occurringin the Andes and the mountains of SE Brazil, are epiphytes. This supportsthe hypothesis of a Gonwanalandicorigin and migrationto the tropics,
where only a few taxa adaptedto this new habitat. Within the neotropics, only one Pilopogon
species, P. longirostratus,is found as epiphyte.
In Campylopusonly a few species such as C.
asperifoliusoccuron treetrunksor bamboonodes.
(Speciesgrowingalso on stems of tree fernscannot be taken into accounthere since they are not
true epiphytes.) Only one species, Campylopus
huallagensis var. weberbauerifrom the Andes,
has become a branchepiphyte.
For an unknownreasonall species of Paraleu-
cobryoideae and Campylopodioideae(as most

Dicranaceae)are strictlyacidophyticand found
only on substrateswith a pH <6. Thereforethese
species are preferablyfound on acidic sand such
as podsoles (andnot on latosoles),on humic soil,
humic soil covering rocks, peat, decomposed
plants such as tufts of Cyperaceae,rotten wood
and acidic rocks.In limestoneareasthese species
growonly on acidichumic soil coveringthe limestone or wood.
All species have a preferencefor open habitats
and are usually not shade tolerant, which may
again be valued as an argumentfor the origin of
at least Campylopusand related genera in subantarctic heathlands. The preference for open
habitatsmakes many species suitableas pioneer
species on open habitats such as soil along road
cuttings or lava flows. Some species are even
aggressive colonists, which is shown by the
spreadingof the circumsubantarcticCampylopus
introflexusin westernEuropeand westernNorth
America, from 0-1000 m altitude.
STRUCTURALADAPTATIONS
WaterStorage
Sub(ant)arcticto temperate,tropicalmontane
or alpine species all have appressedfoliate stems
providing an external capillaryconductingsystem. Only tropical forest species show patent
leaves, as in Campylopuslamellinervis,or verticillate foliate leaves, as in Bryohumbertiafiliand C. hualfolia, Campylopustrachyblepharon
lagensis. In C. savannarum the modification
growingin cerradohabitatshas appressedfoliate
stems while the modification growing in rainforests has patent leaves. These growth forms
seem to be an adaptation for a better gas exchange, which is the major physiologicalproblem for rainforestspecies, especiallyin lowland
rainforestswith high temperaturesand high humidity and probablyalso an effectivemethod to
grow over fallingleaves. Anothercharacterstate
differentiatingtropicaland nontropicalspeciesis
the thicknessof the basal laminal cells. Hyaline,
thin walled basal laminal cells are adapted to
constantly humid conditions and function for
water uptake from the capillarywater of the tomentum and for water storage so long as the
evaporationis not too strong,whichwould result
in a shrinkingof the thin cell walls.Speciesadapted to tropicalenvironmentshave thick, firm,of-
Ecology23
ten pitted basal cell walls, which function for
water storage(Biebl, 1964).
Resistance to Water Loss
Growth form plays an importantrole for resistanceto waterloss. Speciesof exposedhabitats
show dense tufts or cushions (as in high andine
species such as Paraleucobryumenerve,Atractylocarpusssp., Campylopusnivalis, C. cavifolius, C. areodictyon,C. edithae,etc. The same concerns subantarcticspecies.
As in many other acrocarpousmosses a number of speciesof Campylopushave excurrentcostae forming hyaline hairpoints, functioning as
protectionagainst strongradiationand desiccation. In some species these hairpointsvary depending on the habitat, as in Campylopussavannarumwith a concolorousexcurrentcosta in
rainforesthabitats but a distinct hyaline excurrent costa in the "bartletti"'expression of dry
cerradohabitats.
UPTAKE OF WATER AND NUTRIENTS
The anatomies of Paraleucobryoideaeand

Campylopodioideaeshow adaptionsto a less developed internalandwell developedexternalwater conduction. A structuresupportingendohydric water supply is the presence of a central
stand in the stem. Ectohydricfeaturesare easily
wetted surfacesof leaves which become turgescent rapidly,no waterrepellentleaf surfaces,frequent presenceof a rhizoid tomentum and hyaline leaf bases. However, there are no leaf
papillae developed to spread water easily over
the leaf surface;all species have totally smooth
leaves.
SEXUAL REPRODUCTION
Sexual reproductionplays a differentrole in

the genera treated here. Species of Microcampylopus and Campylopodiumare nearly always
foundwith sporophytes.In this respectthesegeneraresembleDicranella,to whichthey ultimately
may be found more closely related than to the
the rate
Campylopodioideae.ForSphaerothecium
of fertilitycannot be estimatedsince the species
are known from only a few collections.It can be
assumedthatspecimenshave been collectedonly
with sporophytes,becausethe speciescan almost
not be distinguishedfrom other generawithout
capsules.In Brotherathereare numerouscollections of fertile material from its Asian range.

However,in North and CentralAmericathe species is found alwayssterile.In the neotropicspecies of Campylopodiella,C.flagellacea is nearly
always sterile and has been found with sporophytes only once, but in contrast,C. stenocarpa
is found usually fertile. Only a small number of
species of Campylopusproducesporophytesfrequently. In many species sporophytesare found
only rarely,and again in many species no sporophytes are known. This is remarkable,inasmuch as Campylopushas developed this special
turnand twist mechanismof the sporophytesfor
spore dispersal, which is, however, apparently
not necessarilyneeded. In contrast to all other
generatreatedhere,some speciesof Campylopus
produce several sporophytes in one perichaetium, usually 3-7. This is especially characteristic for colonists such as C. introflexus,C. richardii or C. occultus.But even closely related
or geographicallyvicariant species can differ in
spore production, like C. introflexus,met frequentlywith sporophytes,versusC. pilifer,which
is rarely found with sporophytes.Dry-adapted
species produce sporophytes more rarely than
species of wet habitats,which apparentlyreflects
betterconditionsfor fertilizationin wet habitats
andbettervegetativepropagationin dryhabitats.
Generally,speciesproducingsporophyteshave
largerranges. This is especially true of species
with large disjunctions,such as between South
America and Africa. Species in which sporophytes are not known have either small ranges
or are knownfrom few localitieswhich also may
be very scattered.An example is Campylopus
oerstedianus,known from less than a dozen records in Costa Rica, Georgiaand SW Europe.
The numberof sporespercapsuleis not known
in even a single genus. However, the spore size
is aboutthe same in all genera,with the exception
of Microcampylopus
and Campylopodium,
which
have sporesof 18-24 ,umdiameterand Sphaerotheciumwith spores 21 ,m in diameter.Therefore, the size and number of capsules may give
an impressionof the fertilityof the genera.
The durationfor the sporophytedevelopment
is not known for tropical species. In temperate
regions all genera show two growth periods a
year, each with one sporophytegeneration.The
sporophytedevelopment thus takes less than 6
months. The fertilityin generalseems to be higher in temperateand subtropicalregions than in
Flora Neotropica
24
the innertropics.This may depend on the photoperiodin the sense that bryophytesusuallyrequire long days for the development of sex organs.
VEGETATIVEREPRODUCTION
The total lack or rareness of sporophytesin

many species shows that there exist effective
methods for vegetative propagation.
The productionof organsfor vegetative propagation is usually seasonal and happens during
a dry season, allowing a dispersalby wind, and
a regrowthin the next humid season. Some populations producepropagulesalways, others never. Plants without propagules show optimal
growth, the others not. Therefore, vegetative
propagationmay indicate ecological stress.
Although there are several differentmethods
of vegetativepropagation,a given methodis usually but not entirelyconfinedto certaintaxa. For
example, brood leaves are characteristicfor C.
fragilis and microphyllousbranchesfor C. flexuosus.However,therehave also been found (but
extremely rarely)microphyllousbranchesin C.
fragilis and brood leaves in C. flexuosus. This
indicatesthatwhileall speciesmay have the same
faculties for all methods of vegetative propagation, they are, however, used differently.
Severalspecies can switch between vegetative
and generativepropagation,which is a most successful response to balance alterationsof habitats.
SYSTEMATIC TREATMENT
CAMPYLOPODIOIDEAE
almost blackish,in loose tufts.Stemserect,rarely

branched,often tomentose in the lower part,comose at tips or not so, rarelyverticillatefoliate.
Dioicous. Perichaetiaterminal, rarely pseudolateral,surroundedby comose leaves; perichaetial leaves with broad sheathingbase, abruptly
contracted to subulae. Stem leaves lanceolate,
erect spreadingto appressed,sometimes homomallous or straight,ending in a fine denticulate,
serrateor smooth, sometimeshyalinetip. Costae
broad, filling 1/3 to 3/ of the leaf width, excurrent
or nearlyso, in transversesection with a median
rowof deutercells,dorsallayersof a (sub)stereidal
band and an epidermallayerand ventrallya single or multilayerredstereidband or a single row
ofhyalocysts, the abaxialsurfacesmooth, ribbed
or lamellose;alar cells lacking, more or less developed or conspicuous, inflated or auriculate,
reddish or hyaline; basal laminal cells hyaline
and thin-walled, rectangular, or incrassate,
subquadrateto rectangular,sometimes pitted, at
margins narrower, shorter or elongate; upper
laminal cells quadrate to rectangular,oval to
elongate, sometimes pitted.
Setae 3-15 mm long, erect or cygneous, yellowish to brownish;capsuleserect,cylindricalto
ovoid or globose, symmetric or asymmetric,
sometimes strumose,smooth or furrowed;opercula usuallylong-rostrate,rarelyconical;annulus
present, dehiscent or not, rarely not differentiated; peristometeeth 16, divided to the middle
or to the base into two prongs,endingin hyaline
papillose points, the outer side striate,the inner
side papilloseor rarelysmooth. Spores 13-15 or
20-24 uimin diam., finely or coarselypapillose
to almost smooth. Calyptraecucullate,entire or
fringedat base.
Plants small, from a few mm to 15 cm high,

yellowish green, light to dark green or olive to
Key to the Generaof Campylopodioideae

1 Stem leaves with broad, sheathingbases abruptedlycontractedinto the leaf apex; spores 20-24 Mmin
diam.
2 Capsulewith stomata, sporespapillose ........................................
3. Campylopodium.
2 Capsulewithout stomata;sporeswarty. .......................................
6. Microcampylopus.
1 Stem leaves (exceptfor perichaetialleaves!)graduallynarrowed;spores 12-15 or 21 mm.
3 Capsuleon 3-4 mm long seta insertedin the perichaetialleaves, spores 21 um in diam. ..........
..... .......................................................................
.
8. Sphaerothecium
3 Capsuleexsertedon longer seta; spores 12-15 mm.
4 Seta erect, more than 1 cm long.
5 Perichaetialleaves sheathing,involving the seta more than halfway up. ............7.
Pilopogon.
5 Perichaetialleaves not involving the seta.
25
Systematic Treatment
1. Atractylocarpus.
6 Capsulestraight;plants equallyfoliate. ..................................
2. Bryohumbertia.
6 Capsulecurved;plants interruptedlyfoliate. ..............................
4 Seta cygneouswhen wet, less than 1 cm long.
5. Dicranodontium.
7 Upper laminal cells elongate,more than 6 times longerthan broad. ..........
7 Upper laminal cells shorter,less than 6 times longerthan broad, quadrateto short rectangular
or oval. ....................................................4.
Campylopus.
Generic delimitation is based mostly on sporophyticcharacters.Successfulidentificationrequiresfertilespecimens,which are, unfortunately, not always present (and in some species of
Campylopus not even known). The gametophytes are very similar anatomicallyand morphologicallyand in some cases the identification
of sterilematerialis problematicalor impossible
without experience.It is generallynot possible

to differentiatesterileplants of Campylopodium
from
andAtractylocarpus
fromMicrocampylopus
Dicranodontiumand not possible for beginners
to distinguishsterile materialof certain species
of Campylopusfrom Sphaerotheciumor Pilopogon. For that reason, an artificialkey for the
identificationof sterile materialis given.
ArtificialKey for Sterile Campylopodioideae

2. Bryohumbertia;Campylopustrachyblepharon,
C. huallagensis.
1 Plants verticillatefoliate. ..............
1 Plantsequallyfoliate or comose at tips.
2 Stem leaves sheathing,from ovate base contractedinto subuli. .................................
...........
3. Campylopodium;6. Microcampylopusand also species of Dicranellasubg.Anisothecium.
2 Stem leaves graduallynarrowed.
3 Upper laminal cells elongate .............................
1. Atractylocarpus;
5. Dicranodontium.
3 Upper laminal cells shorter.
4 Transversesection of the costa with ventralstereids .....................................
...........................................
4. Campylopus;7. Pilopogon;8. Sphaerothecium.
4 Transversesection of the costa with ventralhyalocysts. .........................
4. Campylopus.
1. AtractylocarpusWilliams,Bryologist31:109.
1928, nom. cons. prop.
Mitten,J. Linn.Soc.Bot.12:13.1869,
Atractylocarpus
nom.rej.prop.
exMilde,Bryol.siles.75. 1869.
W.Schimper
Metzleria
Laubm.Deutschl.1:411.1887.
Metzleriella
Limpricht,
exHagen,Kongel.NorskeVidensk.
Metzlera
Schimper
Selsk.Skr.1910(3):15. 1910.
Metzlerella
Hagen,Kongel.NorskeVidensk.Selsk.Skr.
1914(1):62. 1915.
ric, cylindric,without stomata.Annulusnot differentiated.Operculumlongly rostrate,as long

as the capsule.Peristometeeth 16, divided into
two or three prongs,papilloseat tips. Spores 1418 ,tm in diameter,finelypapillose.Calyptracucullate, entire at base.
Type species: A. alpinus (W. Schimper ex
Milde) Lindeberg,typ. cons. prop.
A worldwidemonographhas been published
by Padbergand Frahm (1985). Recent studies
reveal that the type species of Atractylocarpus,
A. mexicanus,must be placed into a genus that
was later describedas CampylopodiellaCardot.
To avoid confusion (Campylopodiellawould be
the next
and forAtractylocarpus
Atractylocarpus,
older name Metzleria would apply) the use of
Atractylocarpushas been proposed for conservation by Frahm and Isoviita, Taxon 37: 967969, 1988.
Autoicousor dioicous.Plants slenderin compact tufts, golden-green,glossy, rarelybrownish

green. Stems usually not branched,erect, 5-35
mm high, reddish tomentose. Leaves erect patent, rarely homomallous, from a lanceolate,
sheathing base contracted into a long subula.
Costa broad, fillingone half of the leaf base, excurrent,in transversesection with a medianrow
of deuter cells and several layers of ventral and
dorsalstereids.Alarcells weaklydeveloped.BasKey to the Neotropic Species of
al laminal cells narrow,rectangular,slightly inAtractylocarpus
crassate and sometimes pitted. Upper laminal
cells elongate,incrassate.
1 Costaridgedat back;capsuleshort,2-3:1.
Seta long, 10-25 mm, yellowish to brownish,
2 Exothecialcells oval, 2-3:1; Peristometeeth
1. A. brasiliensis.
....................
erect, twisted when dry. Capsuleerect, symmetsplit 2/3
26
Flora Neotropica
2 Exothecialcellsrectangular,
5-6:1;Peristome
3. A. nanus.
teethsplitto thebase. ..........
1 Costasmoothat back;capsulelong,3-5:1. ....
2. A. longisetus.
..............................
As shown by the few and less trivial distinguishing characters,all neotropical species are very
closelyrelated.The so-calledAtractylocarpusflagellaceus (C. Muller) Williams (A. costaricensis
(C. Muller) Williams, A. stenocarpus(Wilson)
Zander,A. mexicanusMitten), differsfrom true
Atractylocarpusby its transversesection of the
costa, which has ventral and dorsal hyalocysts,
and the presence of an annulus. It actually belongs to Campylopodiella(Muller & Frahm,
1987).
Specimens examined. BRAZIL. Rio DE JANEIRO:

Serra dos Orgaos, Morro Assu, Liitzelburg 7118 (B,
JE).
The Liitzelburg specimen is sterile and can be

referred to this species only with some doubt,
since the species of Atractylocarpus can be differentiated only by sporophytic characters.
Atractylocarpus brasiliensis is closely related
to A. nanus from the Andes, from which is differentiated only by the less ridged dorsal side of
the costa. These species can be regarded as vicariants that have evolved by the physical separation of populations in the Andes and SE Brazil.
Atractylocarpus nanus and A. brasiliensis differ
from A. longisetus mainly in their shorter capsules and shorter exothecial cells. Both species
occur within the geographical range of A. longisetus. It may be that both represent extreme
(depauperate ?) forms of A. longisetus. This cannot, however, be determined merely by an examination of herbarium material.
1. Atractylocarpusbrasiliensis(C. Miller) Williams, Bryologist 31: 110. 1928. Dicranum

brasilienseC. Muller, Bull. Herb. Boissier 6:
32. 1898. Type. Brazil.Serrado Itatiaia,2300
m, Lagoa,in palude, Ule, Bryothecabras. 109
(holotype,destroyedat B; lectotypenov., BM;
isolectotypes, H, JE, NY, PC, S). Metzleria 2. Atractylocarpus longisetus (Hooker) Bartram,
brasiliensis (C. Muller) Paris, Index Bryol.
Bryologist 49: 110. 1946. Dicranum longiseSuppl. 244. 1900. Metzlerellabrasiliensis(C.
tur Hooker, P1. Crypt. 3a. 1816. DicranodonMiiller) Brotherus in Engler & Prantl, Nat.
tium longisetum (Hooker) Williams, N. Amer.
Pflanzenfam.ed. 2, 10: 191. 1924.
Fl. 15:152. 1913. Metzleria longiseta (Hooker)
Figs. 10, 11.
56: 5. 1920. MetzPlants erect, in dense tufts, to 20 mm high,
yellowish. Stems reddish tomentose. Leaves 46 mm long, from a lanceolatebase contractedto
a subulateapex. Leaf tips dentate. Costae filling
2/3of the leaf base, excurrent,in transversesection
with median deutercells, a multilayeredband of
dorsal stereidsand only a few ventralstereidsin
the middle of the leaf, crenulateat back. Basal
laminal cells incrassate,rectangular,46-64 x 712 um, narrowerat margins forming a distinct
border.Upper laminalcells narrowlyrectangular
to linear, incrassate,45-100 x 4-8 Asm.
Setae 15-20 mm long, golden-yellow, erect,
slightly twisted above. Capsule 1.5 mm long,
about two times longerthan wide, brownto dark
brown.Exothecialcells oval, ca. 2-3:1. Opercula
rostrate,as long as the capsule. Peristometeeth
split up to 2/3into 2-3 prongs.Spores 14-16 tim.
Calyptraebrownish,coveringthe capsule to the
base.
Distribution(Fig. 11). On swampy soil above
forest line. Endemic to SE Brazil.
Brotherus, Bot. Jahrb. Syst.

lerella longiseta (Hooker) Brotherus, Nat.
Pflanzenfam. ed. 2, 10: 191. 1924. Type. Colombia. Fusagasuga, Humboldt & Bonplands.n.
(holotype, BM).
Figs. 1D, 4B, 7D, 8A, 9D, 12, 13.
Atractylocarpusstrictulus(C. Miiller)J.-P. Frahm, J.

Hattori Bot. Lab. 44: 485. 1978. Dicranumstrictulum C. Miiller,Linnaea38: 587. 1874. Campylopus
strictulus(C.Miiller)H. Robinson,Bryologist70: 20.
1967. Type. Colombia.Antioquia:Paramode Sons6n, Walliss.n. (n.v.).
CampylopuslaxiretisHerzog,Beih. Bot. Centralbl.26:
54. 1910. Type. Bolivia. "auf faulem Laubholzim
Bergwaldbei IncacorralProv.Cochabamba,ca. 2200
m," Herzog s.n. (holotype,JE).
Dicranodontiumpusillum Theriot, Hedwigia74: 101.
1934. Type. Colombia.ParamoEl Boquer6n,Troll
2249 (isotypes,B, JE).
Dicranodontiumsetosum Williams, Bull. Torrey Bot.
Club34: 570. 1908.Type.Colombia.Cauca:Paramo
de BuenaVista, Pittier2060 (holotype,NY; isotype,
PC).
DicranummacrodonHampe, Ann. Sci. Nat. Bot. ser.
5, 3: 366. 1865, hom. illeg. (n.v.).
DicranumpittieriRenauld& Cardot,Bull. Soc. R. Bot.
27
'5mm
1.0cm
FIG. 10. Atractylocarpus

brasiliensis(Ue 109, NY). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of
leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule(s).Unlabeled scale bar for microscopicdetails
= 50 Mm.
Belg.31:146. 1893.Type. CostaRica. Foretdu Rancho Flores,Pittier9907 (holotype, PC).

Metzleriaspiripes(C. Miiller) Brotherusin Engler&
Prantl,Nat. Pflanzenfam.1(3):337. 1901. Dicranum
spiripesC. Miller, Linnaea 38: 585. 1874. Type.
Colombia.Antioquia:Piramo de Sonson, Walliss.n.
(holotype,destroyedat B; isotypes, BM, H).
Metzleriastrigulosa(C. Miiller)Brotherusin Engler&
Prantl, Nat. Pflanzenfam.1(3): 337. 1901. Dicranum strigulosumC. Miiller in Renauld & Cardot,
Bull.Soc. Roy. Bot. Belgique31(1):147. 1893. Type.
Costa Rica. Forets du Barba, Tonduz 5510 (holotype, PC).

Metzleriasublongiseta(C. Miiller)Brotherusin Engler
& Prantl, Nat. Pflanzenfam.1(3): 337. 1901. DicranumsublongisetumC. Miiller,Bull. Herb. Boissier 5: 185. 1897 (fide Index Muscorum;n.v.).
Plants to 15 mm high, erect, golden-yellowish
to brown. Stems reddish tomentose. Leaves 4-7
mm long, long-lanceolate, from broad base gradually narrowed into a long acumen. Costa filling
Flora Neotropica
28
60
70
80
^^y^^S^
<
<?L-^'
^^*Mo~
y?
r /
VT
\\
^^^
ss-*:^
--f
!\)~~~~~~~~~
^J
FIG.X1.
Disributon
FIG. 11.
-^
00
^^^^-^
~ots-^'C^:::?^-^-^^
..::.'
]t
400
200
.k.',100
^^S!
~~0
/'
L0
50
200
600
400
by Hendrik
.-J^Prepared
ofAtractlocarus
,,,
300
brsiliesis
(0
and
. nans
800 1000km
500
600
i
__.o
miles
R. Rypkema
(*)0
Distribution of Atractylocarpus brasiliensis (O) and A. nanus (0).
one half of the leaf base, excurrent,in transverse

section with a median band of deuter cells and
multilayeredbands of ventral and dorsal stereids, smooth at back. Basal laminal cells rectangular, slightly incrassate, 57-69 x 18-24 um,
narrowerat margins forming a distinct border.
Upper laminal cells rectangular,incrassate,3678 x 9-14 um. Perichaetialleaves with broad
sheathingbase, suddenly contractedinto a long
fine point formed by the excurrentcosta.
Setae 17-22 mm long, erect, yellow to brown,
twisted when dry in the upper part. Capsules2
mm long, cylindric, ca. 3-5:1, brown to dark
brown.Exothecialcells rectangular,5-6:1. Opercula long rostrate,1.5 mm long. Annuluslacking.
Peristome teeth divided up to 2/3into two prongs.

Spores 14-16 um. Calyptrae golden-yellow with
darker tip, cucullate, covering the whole capsule.
Distribution (Fig. 13). On humid or peaty soil,
on rotten logs, at base of trees at and above timberline, in Central America also in lower altitudes on logs down to 1500 m in Mexico, Costa
Rica, Guatemala, Honduras, Panama, Guyana,
Venezuela, Colombia, Ecuador, Peru, Bolivia and
Brazil.
Cerro San
Specimens examined. MEXICO. OAXACA:
Felipe, Hale 20754 (B). PUEBLA: Amecameca, Sacro
Monte, Pringle 10603 (NY).

GUATEMALA.Chanal,Bernoullis.n. (H).
lower slopes of Cerrode
HONDURAS. MoRAzAN:
29
E
6mm
A
FIG. 12. Atractylocarpuslongisetus(Humboldt& Bonplands.n., BM). A. Plant. B. Leaf. C. Leaf-tip. D.
Transversesection of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic
details = 50 tAm.
Uyuca, Standley & Molina 4307 (F), 4944 (BM). EL

PAAIso: Manzaragua road SW of Guinope, Crum
28515 (F). COMAYAGUA:
Siguatepeque, Standley 56399
(BM, F, NY). INnBUCA:La Esperanza, Standley 25411
(F).
COSTA RICA. LAJUEA: Volcan Poas, Crosby 6281
(MO). SAN Jost: Santa Maria de Dota, 1500-1800 m,
Standley 43169 (B, BM); Cerro de la Muerte, Crosby
3902 (H); Cerro de las Vueltas, Standley & Valerio
43864 (H-BR, PC); El Generil, Skutch 3051 (NY);
foret des Rancho Flores, Tonduz 9910, 9907 (BM,
on Interamerican Hwy. 97 km
H-BR, PC). CARTAGO:
S of Cartago,Bowers837 (FLAS);Sierrade Salamanca

along road from Cerro de la Muerte to San Isidro,
VolcanBarba,arriba
Richards6162 (FLAS).HEREDIA:
de Sacramento,Griffinet al. D 199 (FLAS).
PANAMA. CHIIQui: Boquete, Bajo Chorro, Davidson156 (F).
COLOMBIA.ANTIOQU:Sons6n, Wallissn. (BM).
ARAUCA:
SierraNevada del Cocuy, Boca del Monte,
Paramode Chita, carretera
Cleef10158 (U). BOYACA:
Socha-Sacama,v. d. Hammen & Jaramillo 1643 (U);
SierraNevada de Cocuy, Grubb& GuymerB260 etc.
(BM); Paramo de la Rusia NW de Duitama Fondo,
30
Flora Neotropica
1C
90
0eo
,0
60
70
SC
30
-'.
.....--------^-
200
400
oo
soo
OO0
,. ,
'
--.--
Prepared by Hendrk R. Rypkema
FIG. 13.
Distribution of Atractylocarpus longisetus.
Cleef 7277 (U); Pefia de Arical N de Vado Hondo, Manzanos, Lindig s.n. (BM, NY). HUILA:La Plata,
Cleef 9461 (U); Paramode Chita (carreteraSocha-Sa- vereda Agua Bonita, Diaz et al. 701 (FLAS). MAGcama), v. d. Hammen & Jaramillo 1643 (FLAS,S, U). DALENA:
Parque Nacional de la Sierra Nevada de Santa
CUNDINAMARCA:
CarreteraBogotA-Choachi, Pefia Azul,
Marta,Rangel et al. 341 (FLAS). META:Paramo de
v. d. Hammen & Jaramillo 1644 (U); Paramode Pa- Sumapaz,Cleef4040 (U); CerroNevado de Sumapaz,
Camino Paramo de Guanlacio, Cleef&Jaramillo4052 (U); Lagunasde Buitrago, Cleef7794 (U). SANTANDER:
Cleef 4104 (U); Paramode Sumapaz,San Juan, Cleef tiva-Susa, v. d. Hammen & Jaramillo 1651 (F, FLAS,
8338 (U); San Cayetano, Cleef 6579 (U); W slope of U); vic. LasVegas,Killip&Smith 15888 (NY);Camino
Paramode Guasca,Killip 34075 (F, NICH, S); Suba- Paramode Guantiva-Susa,v. d. Hammen &Jaramillo
choque, Cuchilla El Tablazo, Linares & Bulla 680 1651 (U); carreteraSta. Rosita-Onzaga, Cleef 9855
(COLO);zwischenBarbacoasund Pasto,Lehmanns.n. (U).
Mt. Roraima,McConnell
VENEZUELA.BOLVAR:
(S); inter Tipaquiraet Pacho, Weir205 (BM, FLAS,
Entre El MorH-BR, NY, S); inter Bogota et Fusagasuga,Weir235 & Quelch 344 (BM, H-BR). LIBERTADOR:
(BM, H-BR, NY); Morro del Sabano, Pasto, Andre ro y Aricagua,paramo de Don Pedro, Ruiz-Terdn&
1821 (F, NY); Sabanode Bogota,Villapinzon,Schultes L6pez-Figueiras 8777 (FLAS). MERIDA:Trail from La
12260 (FLAS);Monte del Morro,Lindigs.n. (BM);El Escalera to Puente de Escalera,Luteyn et al. 7846
Boquer6nbei Bogota, Troll2249 (B, PC);Tequenda- (FLAS,NY); propeMerida,Moritz183 (BM, S);prope
ma, Lindig s.n. (BM); Guadalupe, Lindig s.n. (BM); Tovar, Fendler24 (S, BM);Sierrade Santo Domingo,
31
paramo de Mucubaji,Griffinet al. 17506 (B, FLAS,
NAM, NY, PC, S); SierraNevada de Merida,between
Aguadaand La Montana,Griffinet al. 414 etc. (FLAS);
entre Boca de Monte y el Paramo de El Buitre (Aricagua), Lopez-Figueiras12949 (FLAS). SUCRE:Between Cerrode Diablo and Cerrode Neveri, Steyermark 62726 (F). TACHIRA:
Trail leadingto summit of
Paramode Tama, Luteynet al. 7927 (NY);paramode
Tama above Villa Paez, Griffinet al. 127 etc. (FLAS);
paramo El Rosal between La Grita and San Jose de
Bolivar, Griffinet al. 656 (FLAS);paramode Los Colorados 12 km al S de El Zumbador,Ruiz-Teranet al.
8114 (FLAS).
ECUADOR. Quito, Jameson 140 (BM). Chimborazo,Bonplands.n.(BM).Pichincha,Jamesons.n.(BM).
AndesQuitenses,in MonteTunguragua,
Spruce62 (BM,
NY, PC, S). Indanza,Allioni 139 (BM, PC). inter Granadillo et Rosario, Allioni 8114 (H-BR). Anden von
Trail Puno to ChinLotta, Krauses.n. (NY). CARCHI:
Road
gual, Mexia 7616a (B, BM, F, NY, S). ZAMORA:
Zamora-Loja,Holm-Nielsenet al. 3593 (S).
PERU. Tatanara,Lechler 2561 (BM, NY, PC, S).
Tambo de Vaca,Bryan 575a (F, NY). Sandia, Weberbauer2311 (H-BR).AMAZONAS:
Las Palmaszwischen
Balsas und Leimebamba,Hegewald 6974 (F, FLAS,
4462
H, NAM, NICH);E of Chachapoyas,Weberbauer
(H-BR).Cuzco: MacchuPicchu,MenzelP-287 (B);La
Convenci6n, Bues 614 (NY). Huanuco: Huamatios,
3729 (H-BR).LORETO:
N of Moyobamba,
Weberbauer
Weberbauer 4725 (H-BR). SAN MARTIN:Strasse
Frahmet al. 2358 etc. (B)
Chachapoyas-Moyobamba,
BRAZIL.Rio DEJANEIRO:
Sierrados Orgaos,Gardner21 (BM);ParqueNacionalItatiaia,AgulhasNegras,
Frahm 1892 (hb. Frahm).
BOLIVIA.Paradiso,SanJos6-Apolotrail, Williams
1755 (BM, F, H-BR, NY). Unduari, Buchtien s.n.
(H-BR, NY). Between Sorata and Mapiri, Cardenas
1074 (NY). Sillitincara,Yungasvon La Paz, Troll122
(PC). Waldgrenzeueber Tablas, Herzog 2853 etc. (B,
H, JE, PC, S).
as the capsule.Peristometeethsplitto base.Spores

14-16 tm. Calyptraegolden-yellowwith darker
tip, cucullate,covering the whole capsule.
Distribution(Fig. 11). Known only from the
type locality in Peru. The habitat is not known.
This species superficiallyresemblesdepauperate form of A. longisetusbut is treatedhere as a
speciesbecauseof severaldistinctcharacters.The
capsules are shorter than in A. longisetus, the
leaves are shorterand not distinctlyborderedat
the marginswith narrow,elongatecells, the peristome teeth are split to the base (not to 2/3 as in
A. longisetus)and the transversesection of the
costa shows fewer median deutercells and only
a few stereidson the dorsal side and the costa is
ridgedat back. It may be that all these characters
do not representgenotypicaldifferences.
2. BryohumbertiaPotier de la Varde& Theriot,
Bull. Soc. Bot. France86: 422. 1939.
Bridelsect.Filifolii(C.Miller)Brotherus
Campylopus
in Engler& Prantl,Nat. Pflanzenfam.1(3):333. 1901.
DicranumCampylopodes
FilifoliiC. Miller,Genera
musc. frond. 269. 1900.
Type species:B. metzlerelloidesP. Varde& Theriot

A genus with three species, one each in the
neotropics,tropicalAfricaand tropicalSE-Asia.
All species are morphologicallyclosely related
and may appearto be geographicalvicariantsof
the same species.A monographicstudywill likely result in a better understandingof the interrelationshipsof these taxa.
The species of Bryohumbertiawere originally
3. Atractylocarpus nanus Williams, Bryologist
described in Campylopus.They differ in their
31: 109. 1910. Type. Peru. La Convencion,
long,relativelystraightsetae,operculalongerthan
Limasbamba 2500 m, Bues 610 (holotype,
the urn, the presenceof an annulus,smooth cell
Figs. 11, 14. walls on the inner surfaceof the peristometeeth
NY).
Plants in low tufts, to 6 mm high, yellowish- and in their laminal cells, that lack a sharp difgreen. Leaves ca. 4 mm long, lanceolate, gradu- ferentiationbetweenbasalandupperlaminalcells
ally narrowed into a subulate apex. Costa filling (Frahm, 1982a).
half of the leaf base, in transverse section with a
median row of 8-9 deuter cells, dorsal stereids
and ventrally only a few stereid cells, ridged at
back. Basal and upper laminal cells rectangular,
7-14 x 100-1 15 ,um, not conspicously narrowed
at margins.
Setae 8-12 mm long, golden-yellow, erect,
twisted in the upper part. Capsules 1 mm long,
brownish, ovate to short cylindric, 2-3:1. Exothecial cells short rectangular. Opercula as long
1. Bryohumbertiafilifolia (Hornschuch) J.-P.

Frahm, Cryptogamie Bryol. Lich6nol. 3:
366.1982.
Key to the Varieties of B. filifolia
1 Stems not interruptedlyfoliate, only a few mm
lb. var. humilis.
high. ........................
1 Stem interruptedlyfoliate, longer.
32
Flora Neotropica
(1
(S
C
E
4mmZ
FIG. 14. Atractylocarpus

nanus(Bues 610, NY). A. Plant.B. Leaf.C. Leaf-tip.D. Transversesection of leaf.
E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 um.
2 Leavesover 20 mm long; inner basal cells porose. .....................
c. var. longifolia.
2 Leavesless than 20 mm long;inner basal cells
smooth walled ..............
la. var.flifolia.
la. Bryohumbertia filifolia (Hornschuch) J.-P.
Frahm var. filifolia. Dicranum filifolium
Hornschuch, Fl. bras. 1: 12. 1840. Campylopusfilifolius (Hornschuch) Mitten, J. Linn. Soc.,
Bot. 12: 76. 1869. Thysanomitrion filifolium
(Hornschuch) Hampe, Linnaea 25: 361. 1853.
Type. Brazil. "prope Novo-Friburgum," Beyrich s.n. (lectotypus nov., JE).

Figs. 7A, B, 9C, 15, 16.
Campylopusalto-filifolius(C. Mller) Paris,Indexbryol.
ed. 2,1: 297. 1904. Dicranumalto-filifoliumC. Miiller, Hedwigia39: 253. 1900. Type. Brazil.SerraGeral, Ule 835 (holotype, destroyed at B; lectotypus
nov., H-BR).
Thysanomitrionmacrophyllum(C. Miiller)Wilson in
Mitten, Kew J. Bot. 3: 53. 1851. DicranummacrophyllumC. Miller, Syn. musc. frond. I: 402. 1848.
33
c
?
5Br
LE
FIG. 15. Bryohumbertiafilifoliavar.filifolia (Vital 5285, SP). A. Plant. B. Leaf. C. Leaf-tip.D. Transverse
section of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details =
50 Am.
rentz in Geheeb & Hampe, Flora 64: 345. 1881.

Campylopusmacrophyllus(C. Miiller) Mitten, J.
Linn. Soc., Bot. 12: 80. 1869. Type. Peru. Cuchero,
Type. Brazil.Puiggari 176 (n.v.).
Poeppig213 (holotype, destroyed at B; lectotypus Campylopusporphyreodictyon(C. Miller) Mitten, J.
Linn. Soc., Bot. 12: 75. 1869. Dicranumporphyreonov., NY: isolectotype,KIEL).
Campylopusnano-filifolius
(C. Miller) Paris,Ind.bryol.
dictyon C. Miiller, Syn. musc frond. I: 395. 1848.
Suppl.94. 1900. Dicranumnano-filifoliumC. Miil(n.v.).
ler, Hedwigia39: 254. 1900, horn. illeg. Dicranum Campylopuspseudofilifolius(C. Miller) Paris, Ind.
nano-filifoliumC. Miiller, Nuovo Giorn. Bot. Ital.
bryol. 258. 1894. Dicranum pseudofilifoliumC.
N.S. 4:35. 1897. Type. Brazil.Sta.Catarina,Ule439
Miller, Linnaea 42: 471. 1879. Type. Venezuela.
Fendler 40a (holotype, destroyed at B; lectotypus
(lectotypusnov., H-BR).
CampylopusmuelleriLorentz, Verh. Zool.-Bot. Ges.
nov., H-BR; isolectotype,JE).
Wien17:678. 1867.Dicranummuelleri(Lorentz)Lo- CampylopustenuissimusSullivant,Proc. Amer.Acad.
34
Flora Neotropica
100
90
s0
70
60
50
40
0~~~~~~~~~~~~~~~.
~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~
20
FIG.0
1.Dtitoo-
O 2;0 300 40
-Ye,jI
-kmriJlfisla()adCploim
060 6
ei(O
..I..
Prepared by Heodrik RILRypkeffa
FIG. 16. Distributionof Bryohumbertiafilifolias. lat. (@)and Campylopodiummedium(0).

Arts5: 279. 1861. Type. Cuba. Wrights.n. (isotype, ventral and dorsal stereid bands, ridged at back;
GOET).
alar cells differentiated, inflated, reddish, occaCampylopusverticillatus(Hampe) Kindberg, Enum. sionally auriculate and hyaline; basal laminal cells
Hampe,
Bryin.exot. 89. 1889.Dicranumverticillatum
Vidensk. Meddel. Dansk Naturhist. Foren. Kjoe- rectangular, incrassate, 10-16 x 48-93 ,m, narrower and shorter at margins; upper laminal cells
benhavenser. 3, 4: 44. 1872. (n.v.).
short rectangular, ca. 1:2-4, incrassate, 5-6 x
20-30 tm, extending nearly to the apex.
Setae to 15 mm long, erect, straight, rarely
Plants in loose, interwoven tufts, variable in
flexuose or twisted, cygneous only in young sposize, from 1-4 to 8 cm high; stems not tomentose,
rophytes; capsules curved and strumose, coninterruptedly foliate with 1-8 tufts of verticillate
leaves; leaves 5-7(-10) mm long, gradually nar- tracted below the mouth and strongly furrowed
when emptied, brownish to dark brown in age;
rowed into a long, serrate acumen; perichaetial
leaves sheathing the seta; costa up to /3 of the opercula long rostrate, longer than the urn; perileaf width, excurrent, in transverse section with stome teeth reddish with a hyaline tip, the outer
35
SystematicTreatment
surfacestriate,the inner surfacesmooth; spores
ca. 13 ,m in diam. Calyptraecucullate, ciliate
or entire at base.
Distribution (Fig. 16). On humus in forest,
rarelyon rottenwood, basesof trees,on treeferns
or soil-coveredrocks;also in thicketsof secondarygrowthalongroadsor in clearings,sometimes
interwovenin grasses.Usually in rain forests,in
Mexico also in wet oak-pine forest, in southern
Brazil also in Araucariaforests. From sea level
to high montane forests; Mexico, Honduras,
Guatemala, El Salvador, Costa Rica, Panama,
Dominican Republic,Haiti, Cuba, PuertoRico,
Jamaica,Trinidad,Surinam,Venezuela,Colombia, Ecuador,Peru, Bolivia and Brazil.
DOSUL:Fortaleza,
Mar,DusOns.n. (NY). Rio GRANDE
Sehnem 15689 (ASSL);Serrado Faxinal, Sao Francisco de Paulo,Sehnem5322, 5311, 5300 (ASSL).SAo
PAULO:Alto da Serra, Wacket& Decker737 (JE);Jabaguara,Hoehne388 (JE);Serrado BocainaNE Campos da Cunha, Frahm 1558 (hb. Frahm);zwischen
Camposda Cunhaund San Jose dos Bareiros,Frahm
1560 (hb. Frahm);Ilha do Cardoso,Cananeia,Frahm
1556 (hb. Frahm);3.5 km W of Porto Cubatao, Vital
5359 (SP);RibeiraoPires, Vital5285 (SP);km 323 an
der Strasse Sao Paulo-Curitiba, Frahm 1533 (hb.
Frahm);SWJuquiaan derStrasseSP 116,Frahm1557
(hb.Frahm);NE Miracatfian derStrasseSP 116,Frahm
1532 (hb. Frahm);Via Anchietakm 43 oberhalbSantos, Frahm 1142a (hb. Frahm);alongroadSao Miguel
Arcanjo-SeteBarras,Vital7719 (SP);Sao Paulo city,
Parque Estadualdas Fontes do Ipiranga, Yano 196
(SP).
Bryohumbertiafilifolia has been reported from
This species is recognized by its verticillate
Africa(Frahm, 1982b).The Africanplantsmay, foliate leaves, which is otherwise met, in the

however, be conspecific with B. flavicoma (C. Campylopodioideae,
only in Campylopus
Miiller ex Brotherus)J.-P. Frahm.
trachyblepharon and C. huallagensis. This charSpecimensexamined.MEXICO.CHIAPAS:
Lagunas
de Montebello65 km E of ComitAn,Eggers& Frahm
s.n. (hb.Frahm).OAXACA:
SierraJuarez,12 milesabove
Valle Nacional, Sharp et al. 3130 (TENN); 58 km N
of Ixtlan, MickelB 4245 (NY, S). PutBLA:Below Necaxa, Sharpet al. 3117 (NY).
COSTA RICA. SANJosE: 10 km SSE of San Jose,
Crosby9834 (MO);La PalmaareabeyondSan Jeronimo, Crosby 6260 (MO); Cerro Daser area, 9050'N,
84007'W,Crosby6511 (MO).
20 km NE de Medellin,
COLOMBIA.ANTIOQUIA:
Churchill& Sastre-DeJesus 13181a (NY). CUNDINAMARCA:Sasaima, Cleef& v. d. Hammen 4965 (U).
VENEZUELA. Maracuay,Rancho Grande, Mdgdefrau337 (hb. Frahm).
GUYANA. Upper MazaruniDistr., Gradstein5718
(U).
ECUADOR.Zamora,4?5'S,79?W,Ortega601 (MO).
PERU. Rio Huallaga,Ule 1903 (H-BR).
BRAZIL.BAHIA:Mun. de Morrodo Chapeu, Vital
8033 (SP). MINASGERAIS:Km 392 an der StrasseBR
135,Frahm,Camp.Bras.Exsicc.2 (ALTA,B, C, DUIS,
F, FLAS,GOET,GZU, H, HBG, INPA, LBL,JE,KR,
M, MEXU, MO, NAM, NFLD, NY, NICH, PC, SP,
S, TENN, U); Vicosa, Mexia 4916 (FH); Serra da
GrammaDistr. Carangola,Mexia 4302 (FH);Serrado
Caraca,Mun. de SantaBarbara,Vital7703 (SP);Passo
Quatro, Zikan 130 (JE). Rio DEJANEIRO: Serrados
Orgaos,Petropolis,Frahm 1554 (hb. Frahm);Morro
Quamado, Glaziou 7172 (PC); Corcovado, Glaziou
7208 (PC);Itatiaia,Glaziou5622 (PC);Tijuca,Glaziou
5190 (PC). PARANA:Km 51 an der StrasseCuritibaCATARINA:
Joinville,Frahm 1536 (hb. Frahm).SANTA
Serrado Espigao,Frahm 1528 (hb. Frahm);km 181
an der Strasse Curitiba-Lages,Frahm, Camp. Bras.
Exsicc. 1 (ALTA, B, C, DUIS, FLAS, GOET, GZU,
H, HBG, INPA, LBL,JE,KR, M, MEXU, MO,NAM,
NFLD, NY, NICH, PC, SP, S, TENN, U); Serrado
acter may represent an adaptation for better gas

exchange under unfavorable rain forest conditions. It is very variable in plant size, leaf length
and numbers of proliferations, perhaps reflecting
its broad ecological amplitude and broad vertical
geographical range.
lb. Bryohumbertia filifolia (Hornschuch) J.-P.
Frahm var. humilis (Montagne) J.-P. Frahm,
Cryptogamie Bryol. Lichenol. 3: 366. 1982.
Campylopus humilis Montagne, Ann. Sci. Nat.
Bot. ser. 3, 4: 110. 1845. Campylopusfilifolius
(Hornsch.) Mitten var. humilis (Montagne)
J.-P. Frahm, Nova Hedwigia 29: 249. 1978.
Type. Brazil. Blanchet 103 (holotype, PC; isotype, NY).
Figs. 7A, B, 9C, 17.
Campylopusaracuarieti(C. Miiller)Paris, Ind. bryol.
Suppl. 88. 1900. Dicranum araucarietiC. Miiller,
Hedwigia39: 254. 1900. Type. Brazil.SerraGeral,
Ule 661 (lectotype,H-BR).
CampylopushaitensisTheriot, Rev. Bryol. Lichenol.
14: 8. 1944. Type. Haiti. Mt. Formon,Ekman 7561
(isotypes,F, NY, S).
Dicranum rabenii Lorentz, Moosstudien 158. 1864.
Type. Brazil.Raben s.n. (holotype,B).
CampylopusrubricaulisLindbergin C. Miiller, Gen.
musc. frond. 169. 1900, nom. nud. Material.Brazil.
Caldas,Lindbergs.n. (B).
Campylopusstrictisetus(C.Miiller)Brotherusin Engler
& Prantl,Nat. Pflanzenfam.1(3): 334. 1901. DicranumstrictisetumC. Miiller,Hedwigia39:253. 1900.
Type. Brazil. Minas Gerais:Caraca,Ule 1357 (holotype, destroyedat B; lectotypusnov., H-BR).
(Hampe)Jaeger,Ber.ThaCampylopussubarctocarpus
36
Flora Neotropica
This taxon anatomicallyresemblesB. filifolia
var.flifolia. Usually such small plantsare interpretedas juveniles, but in this case the plantsare
found with sporophytes. The taxonomic status
of this taxon can not be decided without field
studiesor cultivationexperiments.It is therefore
treatedhere at the varietal level.
Distribution(Fig. 16). The same range as B.
filifolia, usually on earth covered rocks and at
base of trees. More frequently encountered in
(drier ?) areas such as Minas Gerais in Brazil.
This may indicate that this variety is a depauperate form of B. filifolia or that var. filifolia is
a luxuriantrain forest form.

Distr.
Choapin, Santa Maria,Mexia 9261M (F).
BRAZIL. AMAZNAS:Rd. Manaus-Boa Vista km
Mun. de Morro
10, Frahm 1563 (hb. Frahm).BAHIA:
do Chapeu, Vital7703 (SP).MINASGERMS:
N of Barbacena along rd. BR 135, Frahm 1562 (hb. Frahm).
Rio DEJANEIRO:
Itatiaia,Bueno Rangel,Glaziou9078
(PC); Haut de la Picada Zimler, Glaziou 7370 (PC);
Serrados Orgios, Morro Assu, Liitzelburg6677 (JE);
Palmeiras,Glaziou 9079 (PC);Haut des Orgues,Glaziou 6365 (PC). SAo PAULO:
Between Sao Paulo and
Santos, Camino do Mar km 39, Vitt20638 (ALTA);
Serrado BocainaNE Camposda Cunha,Frahm 1561
(hb.Frahm);Camposdo Jordao,Hoehne642 (JE);Alto
da Serra,Hoehne661 (JE);Pivajussara,Gehrt601 (JE).
Ic. Bryohumbertiafilifolia (Hornschuch)J.-P.

Frahm var. longifolia (Bartram)J.-P. Frahm
comb. nov.
Campylopusfilifolius (Hornschuch)Mitten var. Iongifolius(Bartram)Bartram,J. WashingtonAcad.Sci.
19: 12. 1929. Campylopusharrisii(C. Miller) Paris
var. longifoliusBartram,Contr.U.S. Natl. Herb.26:
64. 1928. Type. Costa Rica. Cerrode las LejasN of
San Isidro, Standley51638 (holotype,FH; isotypes,
NY, US).
1.5cm
var. humilis(Vital
FIG. 17. Bryohumbertiafilifolia
7703, SP). A. Plant.
tigk. St. Gallischen Naturw. Ges. 1877-1878: 496.

1880. DicranumsubarctocarpumHampe, Vidensk.
Meddel. Naturhist.Foren. Kjoebenhavn,ser. 3, 910: 255. 1878. Type. Brazil. Rio de Janeiro: Rio
Preto, Glaziou 9078 (lectotype, BM; isolectotypes,
H-BR, NY).
This variety differs from the typical variety by
its smaller size, and low rosettes. Stems are only
a few millimeters high, and are not interruptedly
foliate.
This taxon differs from the typical variety in

its larger size, more sharply serrate leaves and
especially its porose basal leaf cells.
Specimens examined. COSTA RICA. ALAuELA:
ReservaBiologicade BosqueNuboso de Monte Verde,
15 km SE of Cartago,
Crosby10020 (MO). CARTAGO:
Crosby6087 (MO).
PANAMA. CHIUQui:CerroColorado,20-28 miles
from San Felix, Croat33321 (MO).
3. Campylopodium(C. Miiller)Bescherelle,Ann.
Sci. Nat. Bot. ser. 5, 18: 189. 1873.
Type specimen: C. euphorocladum (C. Miiller)
Bescherelle
Originallya genus of 17 species. A revision
(Giese & Frahm, 1985a)has reducedthe number
to only two species. One species, C. lineare, is
confined to Tasmania and New Zealand. The
other, C. medium,is widespreadfrom New Zealand to SEAsia and Japan.It has also been found
in Chile and in Puerto Rico, from where it has
been reported as Campylopodium pusillum
(=Microcampylopuscurvisetus).This highlydisjunct ocurrencemightbe interpretedas the result
of recentintroductionof the speciesinto the New
World.
The genus is strikinglysimilar to Microcampylopus. Indeed, it is distinguishedonly by the
presenceof phaneroporestomata in the capsule
neck and finely papillose spores.
37
Distribution (Fig. 16). On soil, soil covered
rocks and open banks, colonizing open ground;
Southeast-Asia(Indonesia, Java, Sumatra, Sulawesi, Malaysia,Philippines,New Guinea, Samoa, Society Islands, New Caledonia,Fiji, Hawaii), Taiwan, Japan,Chile and Puerto Rico.
Specimensexamined.PUERTORICO.Cordillera
Central,Jayuya,Masters6873(B, MO,NY).Trailof
Cerrode la Punta,Steere6242(MO).Hillsof Matuillas,northof Villaalba,Steere6831(MO,S).
Campylopodiumcannotbe distinguishedfrom
either Microcampylopusor Dicranella without
sporophytes. Fortunately, Campylopodiumis
commonly fertile. It differsfrom Dicranella by
its cygneous setae, and from Microcampylopus
by the presenceof capsule stomata.
1. Campylopodiummedium (Duby) Giese &

J.-P. Frahm, Lindbergia11: 126. 1985. Di- 4. CampylopusBrid. Mant. Musc. 71. 1819.
dymodonmediusDuby in Moritzi, Syst. Verz.
Plants small to robust,up to 15 cm high, erect,
ZollingerPfl. 134. 1846. Type. Java. Zollinger
411 Za (holotype, G). Figs. IE, 8B, 16, 18. rarelybranched.Stems sparselyto denselywhite
or red tomentose or not tomentose.Leaves5-15
euphorocladum
Campylopodium
(C. Muller)Besche- mm long, lanceolate, erect patent to appressed
relle,Ann.Sci.Nat.Bot.ser.5, 18:189.1873.Aong- foliate,the upperleaves oftenlongerand comose,
stroemiaeuphoroclada
C. Muller,Syn.muse.frond.
1:429. 1848.Type.Java.Zollinger
411 Z(holotype, straight,curledorhomomallous.Leaftips smooth
destroyedat B; lectotypusnov., BM;isolectotype, or denticulate. Costa filling 1/3-7/8of the leaf width
at base, ending in the leaf tip or more or less
NY).
in transversesection on the venlong-excurrent,
For 10 additionalsynonymswith 17 different
tralside with one row ofhyalocysts or 1(-3) rows
genericcombinationsdescribedfromSEAsia and of
stereids,a median band of chlorocysts(deuter
Chile see Giese and Frahm (1985a).
cells),below a layerofstereid or non-stereidcells
Plants 3-20 mm high, usually 13-15 mm high, and dorsally a layer of chlorocysts, which are
in loose mats,yellowishgreento brownish.Stems smooth,ridgedor forminglamellae1-6 cellshigh.
erect, comose foliate in the upper part. Leaves Alar cells lacking or differentiated,reddish or
2-4 mm long with sheatiningbase suddenlycon- hyaline,inflatedto auriculate.Basallaminalcells
tractedinto a long subula. Costa 4 of leaf base, hyaline and thin-walled or incrassate, with
fillingthe subula,in transversesection with me- smooth or pitted cell walls, subquadrateto rectdian deutersand several rows of stereids. Basal angular,usuallysmalleror narrowerat margins.
laminal cells thin walled, narrowlyrectangular, Upper laminal cells incrassate,quadrateto rect25-50 x 4-5 gm. Upper laminal cells short rec- angular,oblique or oval to elongateoval.
Dioicous.Perichaetiaterminal,often bud-like,
tangularto rhomboid, 19-25 x 4-5 mm.
Setae 5-7 mm long, in immaturesporophytes or (rarely)pseudolateral;perichaetialleaves difstrongly cygneous, in mature sporophytesflex- ferentiatedwith broader, sheathing base, conuose when dry and curved when wet. Capsules tractedto a narrow,long subula.
Setae 5-15 mm long,oftenaggregated,
in young
1-1.5 mm long, ovoid, 2-3 times longer than
in
in
and
mature
with
stomata
the
neck, sporophytescygneous, dry
broad,
spophaneropore
striate when empty with six furrows.Annulus rophytesmore or less erectbut sinuoseand twistpresent. Peristome teeth 16, brownish-orange, ed, cygneouswhenwet. Capsuleserectto curved,
striateat base, hyalineand papilloseat apex,split sometimes strumose, striate or furrowedwhen
halfway down. Opercula2 as longas the capsule, empty.Annuluspresentbut not dehiscent.Operobliquelyrostrate.Spores 18-24,tm, finelypap- cula obliquely rostrate,half as long as the capillose, yellowishbrown.Calyptrasmooth at base. sule. Peristome teeth divided to the middle or
Flora Neotropica
38
C4
h1
3I
UU.5mm
FIG. 18. Campylopodiummedium(Steere6242, MO). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule.Unlabeled scale bar for microscopicdetails
= 50 um.
-Subgenus Thysanomitrion(Schwaeger.)Kindb.
emend. J.-P. Frahm,which has only one species
(C. richardii)in the Neotropics (Frahm, 1984a).
This subgenusis essentiallycharacterizedby sporophyticcharacters;narrow,filiform,deeplysplit
peristome teeth, a symmetric capsule, scabrous
base and very finely papillose spores. Most species produce comose perichaetia on appressed
foliate stems, have an areolationwith incrassate
The genus can be divided into three subgenera. and pitted cells, show side nerves and a transmore deeply into two prongs, reddish and horizontally striate below, hyaline and papillose
above. Spores ca. 13 um in diameter, nearly
smooth to papillose. Calyptraeciliate at base or
not, sometimes both conditionsin the same species.
Lectotype species: Campylopus flexuosus
(Hedw.) Brid.
SystematicTreatment39
- SubgenusCampylopuswhich has two sections,
Campylopusand Homalocarpus(Frahm,1983b).
Section Campylopushas asymmetric,often stru-Subgenus CampylopidulumVital, which has mose
capsules, whereas section Homalocarpus
two species worldwide:Campylopusperpusillus
Mitten has symmetric, upright, nonstrumose
(tropicalAfrica)and C. carolinae(Braziland SE
North America). The subgenus has nearly glo- capsules.
bose capsules on very short (only 2-4 mm long
These infragenericcategories can be distinsetae)thatareimmersedin the perichaetialleaves, guished only by sporophyticcharacters,which
and the calyptraeare long ciliatewith cilia nearly are rarely present. Therefore,for practicalpuras long as the calyptra.The subgenusresembles poses, this treatmentof the genusdoes not utilize
the genus Sphaerothecium,but Sphaerothecium this classification.
has largerspores,about 23 gm in diam., and the
calyptraeare entire.
verse section of the costa with ventraland dorsal

bands of stereids.
Key to the Neotropical Species of Campylopus

1 Basal laminal cells thin-walled,hyaline.
2 Leaves endingin a hyaline hairpoint.
C. griseus.
3 Leaf tips recurved(when reflexed,see note for C. introflexusunder C. pilifer). ........29.
3 Leaf tips straight.
4 Stems appressedfoliate, filiform.
1. C. aemulans.
5 Costa not lamelloseat back. ..............................................
36. C.julaceus.
5 Costa lamellose at back. ..................................................
4 Stems not filiform.
47. C. pilifer.
6 Costa lamelloseat back. ....................................................
6 Costa not lamellose.
7 Upperlaminalcells shortrectangularto oblique,ventralhyalocystsof costa equalor smaller
44. C. oerstedianus.
than the median deutercells. ........................................
7 Upperlaminalcells oval to elongateoval, ventralhyalocystsof costa largerthanthe median
deutercells.
23. C. edithae.
8 Leaf tips hooked. ....................................................
8 Leaf tips not hooked.
3. C. amboroensis.
9 Alar cells lacking ...............................................
10. C. bryotropii.
9 Alar cells conspicuous. ...........................................
2 Leaf tips concolorous.
10 Upper laminal cells quadrateor rectangular.
11 Upper laminal cells quadrate.
12 Plants producingbrood leaves in the comal tufts.
11. C. capitulatus.
13 Lowerpartsof stems appressedfoliate. ...........................
25. C. fragilis.
13 Lowerpartsof stems not appressedfoliate. ...........................
12 Plants without brood leaves.
43. C. occultus.
14 Stems ending in comal tufts. ......................................
14 Stems equallyfoliate.
37. C.julicaulis.
15 Stems appressedfoi;ate. ........................................
15 Stems with patent leaves.
30. C. heterostachys.
16 Leaves broadestat base, spinouslydentateat tips. .........
52. C. sehnemii.
16 Leaves broadestbelow midleaf, serrateat tips .................
11 Upper laminal cells rectangular.
7. C. areodictyon.
17 Plantsproducingmicrophyllousbranches. .............................
17 Plantswithout microphyllousbranches.
33. C.jamesonii.
18 Robust plant up to 10 cm high;leaves more than 1 cm long. ........
18 Plants up to 5 cm, leaves 3-7 mm long.
19 Upper laminal cells 3-4 times longerthan broad.
20 Plants to 1.5 cm high;upperstem leaves longerthan lower ones. 61. C. trivialis.
58. C. tallulensis.
20 Plants taller,upperstem leaves as long as lower ones.........
19 Upper laminal cells 1.5-2 times longerthan broad.
60. C. trichophylloides.
21 Leaves 2.5 mm long, appressedfoliate. ................
21 Leaves 5-6 mm, erect patent.
40
Flora Neotropica
22 Leaf apex short, blunt. ............................

..... 46. C. perexilis.
22 Leaf apex elongate. ...................
41b. C. nivalisvar. multicapsularis.
10 Upper laminal cells oval to elongateoval.
39. C. longicellularis.
23 Costa with ventralstereids. ........................................
23 Costa with ventralhyalocysts.
24 Transversesectionof costae as in Paraleucobryum,
with ventraland dorsalhyalocystsand
median chlorocysts.
C. pittieri.
25 Alar cells lacking. ..................................................48.
.......
2. C. albidovirens.
25 Alar cells present. ....................................
24 Transversesection of costae not as above.
26 Plants to 12 cm high; leaves 7-12 mm long.
27 Costa contractedat base, graduallyintergradingwith laminal cells; leaves > 1 cm
C. cavifolius.
long. ........................................................13.
27 Costa not contractedat base, well separated;leaves < 1 cm long...............
..........................................................
8. C. argyrocaulon.
26 Plants to 5 cm high, leaves to 5 mm long.
28 Upper laminal cells 4:1.
29 Costa short lamellose at back with lamellae 1 cell high. .... 56. C. subjugorum.
29 Costa not lamellose, rather,smooth or ridgedat back.
30 Alar cells indistinct,not protrudinginto the costa. ..........
41. C. nivalis.
30 Alar cells distinct, protrudinginto the costa. ................53.
C. sharpii.
28 Upper laminal cells 6-8:1. .....................................
35. C. jugorum.
1 Basal laminalcells incrassate,colored.
31 Leaves ending in a (sub)hyalinetip.
32 Basal laminal cells short, subquadrate.
. 51. C. savannarum.
33 Outerbasal laminal cells quadrate.......................
....
33 Outerbasal laminal cells long and narrow ...............................
34. C. japonicus.
32 Basal laminal cells long, rectangular.
34 Basal laminal cells smooth, minute plants to 1 cm high.
35 Leaf tips straight;costae with ventralstereids. .........................
12. C. carolinae.
35 Leaf tips homomallous;costae with ventralhyalocysts. ................
26. C. gardneri.
34 Basal laminal cells pitted, largerplants.
36 Upper laminal cells elongate,also pitted.
37 Transversesection of the costa with ventralhyalocysts,yellowishplants..........
..............................................................
19. C. cuspidatus.
37 Transversesection of costa with ventralstereids,blackishplants. .... 50. C. richardii.
36 Upper laminal cells not pitted ......................................
10. C. bryotropii.
31 Leaves not ending in a hyaline tip.
38 Leaf tips cucullate. .....................................
19b. C. cuspidatusvar. dicnemioides.
38 Leaf tips not cucullate.
39 Basal laminal cells short rectangularto subquadrate.
40 Outerlaminal cells quadrate.
41 Laminareachingto leaf tip. .....................................
22. C. dichrostis.
41 Laminavanishingbeforeleaf tip.
42 Plants equallyfoliate. .....................................
51. C. savannarum.
42 Plants comose foliate ..................................
65. C. zygodonticarpus.
40 Outerlaminal cells elongate.
20. C. cygneus.
43 Upper laminal cells quadrate. .....................................
43 Upper laminal cells short rectangularto oblique.
44 Plants comose foliate ........................
45b. C. paupervar. lamprodictyon.
44 Plants equallyfoliate. .......................................
4. C. anderssonii.
39 Basal laminal cells longly rectangularto elongate.
45 Plants verticillatefoliate.
46 Costa lamelloseat back. ..................................
59. C. trachyblepharon.
46 Costa not lamelloseat back.
47 Leaves ca. 10 mm long. ...................................
31. C. huallagensis.
47 Leaves ca. 5 mm long. ..........................................
14. C. cleefii.
45 Plants not verticillatefoliate.
48 Basal laminal cells pitted.
49 Upper laminal cells rectangularor oval.
50 Upper laminal cells oval.
51 Leaves ca. 15 mm long.
52 Costa filling2/3of leaf width. ...................
55. C. subcuspidatus.
SystematicTreatment
41
52 Costa filling /3 of leaf width. .........................

54. C. shawii.
51 Leaves ca. 6 mm long.
53 Transversesection of costa with ventralhyaloycsts.
54 Upper laminal cells pitted. ......................
62. C. uleanus.
54 Upper laminalcells smooth. ....................
64. C. widgrenii.
53 Transversesection of costa with ventralstereids.
55 Upper laminal cells pitted. ..................
27. C. gastro-alaris.
55 Upper laminal cells smooth. ....................
63. C. viridatus.
50 Upper laminal cells rectangular.
56 Costa longly excurrent,sharplydentate.
57 Upper laminal cells 4-6:1. ..........................18.
C. cubensis.
57 Upper laminal cells 2:1. ........................
16. C. controversus.
56 Costa shortlyexcurrent,serrate. .......................
6. C. arctocarpus.
49 Upper laminal cells quadrateto subquadrate.
58 Robustplants,leavesca. 10 mm long,lamelloseat back. .......38. C. lamellinervis.
58 Medium sized plants, leaves 5-7 mm long, not lamellose. ...............
.........................................
24. C. flexuosusvar. incacorralis.
48 Basal laminal cells smooth.
59 Leaves endingin a long, slendertip, which is as long or longerthan the lamina.
60 Leaf tip serrate.
61 Upper laminal cells oblique to oval. ..................
21. C. densicoma.
61 Upper laminalcells subquadrateto short rectangular. .......40.
C. luteus.
60 Leaf tip smooth. ...............
........................49.
C. reflexisetus.
59 Leaf tips shorter.
62 Upper laminal cells subquadrate.
63 Plants equallyfoliate.
64 Alar cells protrudinginto the costa.
65 Upper laminal cells irregular,quadrate, short rectangularor
............
42. C. oblongus.
oblique. .........................
65 Upperlaminalcells subquadrate,in distinctrows. .. 15. C. concolor.
64 Alar cells not protruding.
66 Leaves to 3 mm long, blunt, crisped. ...... 17. C. cryptopodioides.
66 Leaves longer,lanceolate,not crisped. ...........24.
C.flexuosus.
63 Plants endingin a comal tuft ............................
45. C. pauper.
62 Upper laminal cells rectangularor oval.
67 Upper laminal cells rectangular.....................
57. C. surinamensis.
67 Upper laminal cells elongateoval.
68 Leaf tips sharplyserrate.
69 Vegetativepropagationby microphyllousbranches;piramo species. ..........................................
32. C. incertus.
69 Vegetativepropagationby brood leaves;epiphyticspecies....
9. C. asperifolius.
.............................................
68 Leaftips smooth. ................................
28. C. gemmatus.
1. Campylopusaemulans(Hampe) Jaeger,Ber.
C. Muller,Hedwigia39:260.
Dicranumfilicaudatum
1900. Type. Brazil. Minas Geraes: Serrade Ouro
S. Gall. Naturw. Ges. 1870-1871: 444. 1872.
Preto, Ule 1367 (holotype,destroyedat B;lectotypus
Thysanomitriumaemulans Hampe, Vidensk.
nov., H-BR).
Meddel.Naturhist.Foren.Koebenhavnser. 3,
setaceo-rigidus(Hampe)Jaeger,Ber.Tha2: 273. 1870. Pilopogon aemulans (Hampe) Campylopus
tigk.St. GallischenNaturwiss.Ges. 1877-1878:496.
Brotherusin Engler& Prantl, Nat. Pflanzen1880. Dicranumsetaceo-rigidumHampe, Vidensk.
fam. 1(3): 336. 1901. Type. Brazil.Rio de JaMeddel. Naturhist. Foren. Kjoebenhavnser. 3, 910:257. 1878. Type.Brazil."adRio Preto,"Glaziou
neiro, Glaziou3307 (isotypes,H-BR, NY, PC,
9077 (isotypes,NY, PC).
20.
S).
Figs. 19,
stricticaulis
Ind.
Campylopus
(C. Muller)Paris,
bryol.
Suppl. 97. 1900. Dicranum stricticauleC. Miller,
Bull. Herb. Boissier6: 38. 1898. Type. Brazil.Serra
do Itatiaia, Ule 1795 (holotype,destroyedat B; lecDicranum auribrunneumC. Miiller, Hedwigia 39:
260. 1900.Type.Brazil.Goyaz,Serrade Balisa,Ule
totypusnov., H-BR; isolectotype,S).
1533(holotype,
atB;lectotypus
destroyed
nov.,PC). Campylopussubincrassatus(Hampe)Jaeger,Ber.Thain EnCampylopusfilicaudatus
(C.Miller)Brotherus
1880. DicranumsubincrassatumHampe, Vidensk.
gler& Prantl,Nat. Pflanzenfam.
1(3):332. 1901.
auribrunneus
inEnCampylopus
(C.Miiller)Brotherus
gler& Prantl,Nat. Pflanzenfam.
1(3):332. 1901.
42
Flora Neotropica
/ti
w
4.7mm
0,
5.3cm
AM
~es CI~C~ r
FIG. 19. Campylopus aemulans (Vital 5710, SP). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section of
leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details = 50 Am.
Meddel.NaturhistForen.Kjoebenhavnser. 3, 4: 47.
1872. Type. "Brasiliaaustralis,"Glaziou 4507 (isotype, H-BR).
Campylopusbrunneo-bolax(C. Miiller) Brotherusin
Engler& Prantl,Nat. Pflanzenfam.1(3): 333. 1901,
nom. nud.? (accordingto Index Muscorum,basionym ignot.).
Plants filiform with appressed foliate slender
stems, to 8 cm high, erect, yellowish green above
and reddish below, tomentose at base. Fertile
plantswith comal tufted perichaetia.Leaveslanceolate from an ovate base, 4-6 mm long. Costa
of the leaf base, excurrentin a hyfilling 1/3(-1/2)
aline, serrate tip, which may be short or even
lackingin shaded habitats,in transversesection
with ventral hyalocysts and dorsal groups of
stereids, ridged at back. Alar cells reddish, inflatedor less differentiatedto lacking.Basallaminal cells elongate rectangular, hyaline, thin
walled, 13-22 x 26-45 Am,narrowerat margins.
lO
90
43
70
0o
60
50
ir
L.,/
--X1~~~~~~1
200400
0
600
800100,
10 200 3D00
.
--
00 00
go ..
Prepared by Hendrik R. Rypkema
FIG. 20.
Distribution of Campylopus aemulans (0) and C. albidovirens (@).
Upper laminal cells oval to elongate oval, 3-6

x 13-19 ,m, incrassate.
Sporophyte not known.
Distribution (Fig. 20). On open soil, sand and
gravel in sunny habitats along roadsides, on

banks, and other disturbedsites. Found only in
southeasternBrazil from Minas Gerais to Rio
Grandedo Sul and also in Argentina.
(H-BR);Mun. de PadreParaiso,along road 116, Vital

5896 (SP). PARANA:
Mun. de Senges, along highway
PF 11, Vital5841 (SP);Mun de Castro,along Hwy PR
11, Vital 5822 (SP); Ponte Grossa, Parque Nacional
Vila Velha, Frahm 1894 (hb. Frahm);2 km W of Rio
GuaribobaNW of Curitiba, Vitt21464 (ALTA). Rio
DE JANEIRo: Schiffner 759 (H-BR); An Felsen der Ti-
juca, Ule 2048 (H); Serra do Itatiaia, Schiffner630

(H-BR), Dusen 132 (H-BR, NY, S); Itatiaia,zwischen
Pico de Registro und Agulhas Negras, Kalb & Poelt
s.n. (hb. Frahm);Abrigo Reboucas, Griffin& Vital44
(FLAS);Felsen der Serrade Balisa, Ule 1533 (H-BR);
Specimens examined. BRAZIL. ESPhUTOSANTO: an Felsen der Achmer, Ule 2005 (HBG, H-BR); Ad
Serrade CaparaoNat. Park, Pico Bandeira,2880 m, confines Rio de Janeiro-MinasGeraes,Schiffner460,
Schdfer-Verwimp8069 (hb. Frahm). MINASGERAIS: 463 (B). Rio GRANDEDOSUL: Mun. de Vacaria, along
Ule 1366, 1368 (H-BR).Serrade OuroPreto, Ule 1367 BR 116, Vital5710 (SP);Sao Franciscode Paula,Dal
44
Flora Neotropica
CATARINA:
Pont et al. 221 (NY). SANTA
Lajes,Sehnem ually mergingwith the lamina, slightlyribbedat
5434 (SP). SAo PAULO:
Mos~n 19 (H-BR), Schiffner back. Alar cells conspicuous,inflated,reddishor
2676 (B, HBG,H-BR);Mun.de Brotas,16 km W of

Vital
Itapirina,Vital2196(SP);Mun.de Jacupiranga,
2833 (SP);Iguape,Schiffner
2676,2159 (NY).
NEof C6rdoba,v.Hiibschmann
s.n.
ARGENTINA.
(hb.Frahm).
hyaline, occupyingpart of the costa. Basal laminal cells rectangular,hyaline, thin-walled, 1638 x 58-90 um, narrowerat margins. Upper
laminal cells irregularlyrectangular,13-26 x 3This species is identifiedby its filiformstems. 9 jum,vanishingnear midleaf.
Setae 15 mm long, pale brown, curved and
In its stiff, filiform stems it closely resembles
twisted.
Capsules2 mm long, pale brown, cylinPilopogongracilis(Hooker)Bridel,which is also
dric.
Opercula
long rostrate.
found in SE Brazil. Fertile specimens of PiloVegetative
propagation
by means of small mipogon are differentiatedby the long sheathing
branches in the axils of the upper
crophyllous
and
setae.
leaves
the
Sterile
perichaetial
straight
leaves.
specimenscanbe differentiatedonly by the transDistribution(Fig.20). On rocks,soil andwood.
verse section of the costa, which shows ventral
In
Mexico in coniferousforests,south of Mexico
stereids in Pilopogon but ventral hyalocysts in
in
open
placesin montane rainforestsfrom 2500
Campylopusaemulans.
m to the forest-lineand in subalpineforestsand
not
or
aemulans
have
may
Campylpus
may
thicketsup to 4000 m; Mexico, Guatemala,Coshyalinehairtippedleaves.Epilosespecimenshave
been describedas C. setaceo-rigidus,subepilose ta Rica, Hispaniola (Dominican Republic), Colombia, Venezuela, Ecuador,Peru, Bolivia.
forms as C auribrunneus.
In leaf characterssuch as leaf shape, basaland
Mineral
Specimensexamined.MEXICO.HIDALGO:
upper laminal cells and in its hyaline hairpoint, El Chico, Sharp et al. 1805 (TENN). MEXICO
D.F.:
this species resembles C. pilifer, which differs, Lagunasde Zempoala,Frahm, Camp. Exsicc. 6 (B,
however, by the presence of a costal lamella at BING, BUF, C, DUIS, DUKE, EGR, F, FLAS, G,
back. It resembles Campylopusjulaceus in re- GOET, GRO, GZU, H, HBG, JE, KRA, M, MEXU,
MO, NAM, NFLD, NICH, NY, PRC, PRE, POZ,
gards to its appressedfoliate stems, leaf shape RNG, S, U, UPS);Nevado de Toluca,BrotherAmable
and hyaline basal laminal cells. Campylopusju- 1906 (NY). MICHOACAN:
Capacuaro,Rees & Baltazar
laceus, however, has oval, incrassateupperlam- 763 (MEXU); 6 km NE de Zinapecuaro, 19?54'N,
34 miles
100?47'W,Delgadillo4974 (MEXU).OAXACA:
inal cells.
E of Mitla,Sharpet al. 2883b (TENN);northof Oaxaca
on Hwy. 175 at SierraJuarezgap, Sharp et al. 3033b (TENN).
2. Campylopus albidovirens Herzog, Biblioth.

Bot. 87:19.1916. Type. Bolivia. "Quellwiesen
GUATEMALA. QUEZALTENANGO:
Cerro Quemado
an der WaldgrenzefiberTablas,"Herzog2782 above Los Valvos, Standley86137 (F).
VolcanPoas, EggersCR
(holotype, JE; isotypes, B, H-BR, FH, NY, 1.1COSTA RICA.ALAJUELA:
(hb. Frahm).
S).
Figs. 20, 21.
COLOMBIA.BOYACA:
Paramode
Misc.
Theriot,Smithsonian
angusti-alatus
Campylopus
Collect.85(4):2. 1931. Type. Mexico.Rio Frio,
BrotherAmable1724(holotype,PC;isotypes,FH,
H-BR).
49:110.
Bartram,
Bryologist
Campylopus
guatemalensis
1946.Type.Guatemala.Dep. Totonicapan,
Standley86159(holotype,FH;isotypes,F, FH,MICH).
Plants pale yellowish green or whitish green,
in dense mats. Stems to 3 cm high, equally foliate, reddishtomentose below. Leaveserect, ca.
4-5 mm long, from an ovate base graduallynarrowed to an acumen of varying length. Costa
filling 3/4-4/ of the leaf base, excurrent, denticulate
at the extreme apex, in transversesection with

ventralhyalocysts,median chlorocystsand dorsal hyalocystsalternatingwith chlorocysts,grad-
Pisva, Cleef4608
(U); CordilleraCentral,Mt. Purac6,Killip6655 (NY).
CUNDINAMARCA:
CuchillaEl Tablazo,Linares& Bulla
363 (COL);Paramode CruzVerde,CarreteraBogotaCoachi, Cleef 3182 (U); Paramo entre Cogua y San
Cayetano, Cleef 6130 (U); Paramo de Palacio, Cleef
3681 (U); ParamoAlto, Smeets 15N (U).
VENEZUELA.JUNIN:
Paramode Tami abovePaez,
SierraNevada
Griffinet al. 153 (FLAS).LIBERTADOR:
de Merida,Aguada,Griffinet al. 1623 (FLAS).MERIDA: Between Aguada and La Montafia, Griffinet al.
235, 1660 (FLAS);nearLagunade Los Anteojos,Griffin et al. 346 (FLAS).Rangel,Sierrade SantoDomingo,
Paramode Mucubayi,Griffinet al. 932 (FLAS).
ECUADOR.PINCHINCHA:
CordilleraOriental,west
of summitalongQuito-Baezaroad,SteereE 196 (NY).
PERU. ANCASH:
CordilleraBlanca, Laguna Llanganuco,Hegewald7534 (hb. Hegewald);LagunaLlaca,
Frahm, Camp.Peruv.Exsicc. 1 (ALTA,B, BM, BUF,
DUKE, EGR, F, FLAS, G, GRO, GZU, HBG, KR,
45
2mm
3.7mm
5.3cm
FIG. 21. Campylopusalbidovirens(Cleef 6130, U). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of
leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule.Unlabeled scale bar for microscopicdetails =
50 Atm.
of Chuma,Lewis79-1005(F);NevadoJankhoUma
aboveSt. Antonio,Lewis79-1544(F).
37578 (COLO). AYACUCHO:
Along road from Huanta
This species resembles the genus Paraleucoto SanFranciscoabove Tambo,Frahm,Camp.Exsicc.
54 (ALTA, B, BING, BM, BUF, C, DUKE, EGR, F, bryum in its very broad costae, alar cells proFLAS,G, GRO, GZU, H, HBG, KRA, MEXU, NAM, trudinginto the costa and the lamina vanishing
NFLD, NICH, NY, PC, PRC, POZ, RNG, S, U). LA near midleaf. More
importantly,the transverse
LIBERTAD:
Cerro Sango, Hegewald 5949 (hb. Hegesection
of
the
costae
with ventraland dorsal hywald).
BOLIVIA. COCHABAMBA:
Cerro Chua Laguna N of alocysts and median and dorsal chlorocysts
Corani, Lewis 79-2288A (F). LA PAZ:Near Curva NNW
closely resembles the costa of Paraleucobryum
NAM, NICH, NY, MEXU, NFLD, PC. POZ, RNG,
S, U); QuebradaTulparajunear Huaraz,ArmstrongB
46
Flora Neotropica
4cm
v^
3mm
FIG. 22. Campylopusamboroensis(Herzog273, JE). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails = 50 Mm.
longifolium,from which it differsonly by a few

substereidaldorsalcells in the middle of the leaf.
The relationshipof this species to Campylopus
is basedentirelyon its cygneoussetae of the sporophyte,which has been found only once in Colombia.
Campylopusalbidovirensis closely related to
C. pittieriWilliams,which has, however, no alar
cells, a more turgid,leucobryoidappearance,and
the costa narrowed at leaf base with broader,
laxer ventral cells of the costa. Campylopuspittieriis a paramospecies, whereasC. albidovirens
is a species of montane forests. These species can

be regarded as a species pair that has developed
from a common ancestor by means of ecological
differentiation.
This species varies considerably in the length
of the leaf tip, which is longer in shaded, humid
habitats.
At high elevations this species produces microphyllous branches in the axils of the upper
leaves that resemble those of C. areodictyon. Both
species are similar in the leaf structure and can
be easily confused. They are distinguished only
4. Campylopusanderssonii (C. Miller) Jaeger,
Ber. Thitigk. St. Gallischen Naturwiss. Ges.
1870-1871: 436. 1872. Dicranumanderssonii
C. Miller, Bot. Zeitschr.14: 169. 1872. Type.
Ecuador.GalapagosIsl.: CharlesIsl., Anders3. CampylopusamboroensisTheriot,Rev. Bryol.
son
s.n. (holotype, destroyedat B; lectotypus
Lichenol. 11: 54. 1939. Type. Bolivia. Cerro
nov.,
NY; isolectotypes,FH, H-SOL, S).
273
Amboro, Herzog
(holotype, JE; isotypes,
Figs. 24, 25.
23.
S, PC).
Figs. 22,
by the transversesection of the costa, which has

dorsal groups of stereids in C. areodictyonbut
only substereidsin C. albidovirens.
Herzog,Biblioth.Bot.87:21. 1916,
Plants blackishor blackishbelow and brown- Campylopusfulvus
horn.illeg.Type.Bolivia.RioParacti,1800m, Herish above, to 4 cm high, in loose tufts. Stems
zog 4996(holotype,JE;isotypes,B, FH,H-BR,S).
equally foliate with erect patent leaves, the leaf Campylopus
insularisBartram,
Proc.Calif.Acad.Sci.
Isl.:
ser.4, 21:80. 1933.Type.Ecaudor.
apices often slightly recurved. Leaves 4-6 mm
Galapagos
DuncanIsl.,Stewart3323(lectotypus
nov.,FH).
long, narrow lanceolate. Costa filling /3 of the
leaf width, in transversesection with ventralhy- CampylopuslongisubulatusTheriot, Rev. Bryol. Lich6nol.11:49. 1939.Type.Bolivia.Tablas,1800m,
alocysts,ridgedat back,excurrentinto a hyaline,
Herzog4566(holotype,PC;isotypes,B, JE).
serrateawn.Alarcells lacking.Basallaminalcells CampylopussubincacorralisTh6riot,Rev. Bryol. Lithin walled, rectangular,3-10 x 19-58 ,um.Upchenol.11:44. 1939.Type.Bolivia.Tablas,3400m,
Herzog2799(holotype,n.v.;isotype,B).
laminal
cells
oval
to
per
incrassate,
elongateoval,
3-6 x 13-19 um.
Plants yellow-greento olive brown, to 7 cm
Sporophytenot known.
high,
usually2-4 cm high,whitish-tomentosebeDistribution(Fig. 23). On wet rocksand moist
with blackish stem and long, crowded,
soil in alpine vegetationin the Andes from 3500 low,
sometimes slightlycurved comal leaves. Leaves
to 4400 m in Venezuela, Colombia, Peru, and
to 11 mm long, lanceolate,graduallynarrowed
Bolivia.
to a long, slender subula, serratein the upper
third. Costa filling 1/2of the leaf base, smooth at
examined.
COLOMBIA.
CUNDINAMARSpecimens
CA:Paramode Palacio, Cleef 4090 (U). MAGDALENA: back, excurrentas a long awn, in transversesecSierra
NevadadeSantaMarta,Cuatrecasas
24570(NY). tion with large ventral hyalocysts and dorsal
Sierrade Santo Domingo, groups of 2-5 stereids. Alar cells large and inVENEZUELA.MERIDA:
29 (FLAS).
Gonzales-Pereira
flated. Basal laminal cells short, subquadrateto
PERU. ANCASH:
CordilleraBlanca, LagunaLlaca,
Frahm,Camp.Peruv.Exsicc.2 (ALTA,B, BM,EGR, short rectangular(1-2:1), thick-walledand pitF, FLAS,G, GRO,GZU,HBG,KR,MEXU,NAM, ted, at marginsnarrowand elongate (30 x 30NFLD,NICH,NY, POZ,S, TRT,U); LagunaQue- 60 ,Am).Upper laminal cells irregular,subquadrococha,Hegewald7662(hb.Hegewald).
rate, short rectangularor obliquely, 6-9 x 1030 ,m, smaller at margins, extending in a few
CampylopusamboroensisresemblesC. pilifer
rows nearly to the apex.
marginal
in its hyaline tipped leaves, hyaline basal lamiSporophytenot known.
nal, and oval upper laminal cells, but is distinDistribution(Fig. 25). On shaded, earth-covguished by longer, elongate oval upper laminal eredrocksand soil in humid forestsat elevations
cells and especially by the transversesection of between 500 and 3000
m; Mexico, Costa Rica,
the costa without dorsal lamellae.
Colombia, Bolivia and on the GaVenezuela,
For 40 yearsit had been known only from the
lapagos Islands of Duncan, Isabela, Santa Cruz
type locality in Bolivia, but accordingto recent and San Cristobal.
recordsthe species seems to be widespreadbut
of scatteredoccurrencethroughoutthe Andes.
Fincade
MEXICO.CHIAPAS:
examined.
Specimens
Theriot in the type description referredthis Liquidambar,15?42'N,92?45'W,Delgadillo4673
species to subg. Thysanomitrion,despitethe fact (MEXU);CerroTres Picos, Mun. de Villa Corzo,
SierraJuarez
that the sporophytewas unknown.The presence Breedlove25023 (hb. Frahm).OAXACA:
N of Oaxacaon Hwy.175,E. Sharp14b,Smith
Gap
of ventralhyalocystsin transversesection of the et al. 460
(TENN);12 milesaboveValleNacionalon
costa indicatesthat this species should be placed Hwy. 175, Sharp et al. 3126 (TENN); 23-24 km NE
de Llanode las Flores,Delgadillo805 (MEXU).Pass
in subg. Campylopus.
Flora Neotropica
48
80
90
60
70
40
50
30
0 l
- ---------------
"2
.-- -
S
---------20
0.-0
'I
200
02
400
600
800
1000km
mil
FIG. 23. Distributionof Campylopusamboroensis(0) and C. angustiretis(0).

above Llano de las Flores on the road between Ixtlan
COLOMBIA. BoYAcA:Sierra Nevada del Cocuy,
de Juarez and Tuxtepec, Sharp et al. 2381, 2397d Quebrada Bocatoma, Cleef8743 (U). CUNDINAMARCA:
(TENN);E slope of SierraJuarez60 miles W of Tux- Subachoque,CuchillaEl Tablazo,Linares& Bulla 544
tepec, Websteret al. 253 (TENN).
(COL);Paramode Sumapaz,Cleef8430 (U);Cordillera
COSTA RICA. HEREDIA:Laguna Barba, 10?05'N, de La Leonora60 km NNE de Bogota, v. d. Hammen
83?55'W,Crosby10928 (MO). SANJOSE:Macizo de & Jaramillo 3178 (U).
CedralSW of Aserri, Crosby3755 (MO).
VENEZUELA. ARAGUA: Henri Pittier Nat. Park,
49
A~~~~~~~~~
FIG. 24. Campylopusanderssonii(Gradstein& WeberM 36, U). A. Plant.B. Leaf.C. Leaf-tip.D. Transverse
section of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails =
50 'um.
Owens30 (FLAS);between Caracasand Colonia Tovar, Nee & Whalen16879 (NY). MERDA: La Carbonera area near La Azulita, Griffinet al. 2243 (FLAS);
El Portachuelo,carreteraEl Morro-Aricagua,LopezF. 12596 (FLAS).TACHRA:
Pairamode Tama, RuizTerdn8369 (FLAS).TRUJILLO:
Paramode La Canada,
Ruiz-T. 9193b (FLAS).
Neotrop. Exsicc. 3 (U). LOJA:CajanumaE of Loja,

Laegaard53629D (NY).
BOLIVIA.COCHABAMBA:
E face of CerroChuaLaguna, 17?13'S,65?53'W,Lewis 79-2224A (F). TARiJA:
Caste6n,alongroadto Tarija,21?28'S,64?12'W,Lewis
79-478 (F).
This species is distinguished from all other

ECUADOR. GALAPAGOS
ISLANDS:Isla Santa Cruz,
Weber13020, 14151, 14219, 39189, 41243 (COLO), species of the genus with long subulate leaves by
13668 (COLO,NY), Gradstein& WeberM 36 (U), M its subquadrate basal laminal cells and its serrate
19 (NY, U); IslaIsabela,CerroAzul, Gradstein,Bryoph. leaf tip.
50
Flora Neotropica
o100so
._90
.
70
80
60_50
/~
200300
600..
000.00
,-
40
~30
FIG. 25. Distributionof Campylopusanderssonii.
This species has variable basal laminal cells.

namensis C. Muller var. angustiretis (Austin)
J.-P. Frahm, Bryologist 83: 582. 1980. Type.
They can be more or less pitted, with alar cells,
that are very distinct and have collenchymatous
U.S.A. Florida: Jacksonville, Austin s. no. (hothickenings(e.g., the types of CampylopusanFigs. 23, 26.
lotype, NY).
derssoniiand C. subincacorralis)but less develdelicatulusWilliams, N. Amer. Fl. 15:
oped in the types of C. longisubulatusand C. Campylopus
137. 1913. Type.Cuba.Herradura,Britton6523 (hofulvus. In other expressions(e.g., the types of C.
lotype, NY; isotype, PC).
longisubulatusand C. fulvus), the basal laminal CampylopusditrichoidesBrotherus,ActaSoc. Sci. Fenn.
19: 7. 1891. Type. Brazil.MinasGerais: Wainios.n.
cellsaremoreor less subquadrateor shortrectan(holotype,H-BR; isotype, S).
gular,the leaf apex can be variablytoothed and
ditrichoidesvar. robustiorBrotherus,Bih.
the back of the costa can be smooth or toothed Campylopus
Kongl. Vetensk.-SvenskaAkad. Handl. 21 afd. III
in the upperpart of the leaf.
(3): 6. 1895. Type. Brazil.Glaziou 11761 (holotype,
5. Campylopusangustiretis(Austin)Sullivant&
James, Man. Moss. N. Amer. 80. 1884. DicranumangustireteAustin, Bot. Gaz. 4: 150.
1879. Campylopusgracilicaulis Mitten ssp.
angustiretis(Austin) Kindberg,Gen. Eur. N.
Amer. Bryin. 2: 204. 1897. Campylopussuri-
H-BR).
CampylopusleptodictyonBrotherus,Denkschr.Akad.
Wiss. Wien math.-nat.K1.83:259. 1926. Type. Brazil. Glaziou 11744 (lectotype,H-BR).
Plants to 5 cm high, usually smaller, often only
a few mm high, in loose mats. Stems usually not
branched, equally foliate with conspicuous
51
3mm
5?m
FIG. 26. Campylopusangustiretis(Griffin88, FLAS).A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 tm.
spreadingleaves, the upperleaves dense, forming

a comal tuft. Leaves6 mm long, lanceolate,long
decurrent, from an ovate base gradually narrowed into a fine point, denticulate at the extreme apex, keeled. Costae filling only /4 of the
leaf base, excurrent,in transverse section with
ventralhyalocystssmallerthan the median deuter cells, with conspicuouslythinneradaxial cell
walls, dorsallywith a narrowband of substereid
cells, smooth at back. Alar cells large, inflated
and widelyauriculate,long decurrent.Basallam-
inal cells incrassate, elongate rectangular, often

slightly pitted, 6-13 x 32-48 Am, narrower at
margins. Upper laminal cells incrassate, elongate, 6-10(-12) times as long as broad, not sharply delimited from the basal laminal cells, 4-6 x
26-45 tm.
Distribution (Fig. 23). On wet sandy soil in
swampy places, on wet earth covered rocks and
on soil beside streams. In the Caribbean region
in coastal areas, up to 1000 m, in Brazil from
52
FloraNeotropica
800 to 2250 m; Jamaica,Cuba, SE and E Brazil,

reportedalso in Haiti and Puerto Rico.
Key to the Varieties of
Campylopus
arctocarpus
examined.
CUBA.PinardelRio,Vinales,
Specimens
1 Leaveslanceolatewith shortexcurrentcostae,not
Sameks.n. (PRC).
falcateat stem tips. ........
6a. var. arctocarpus.
PARISH:
JAMAICA. CLARENDON
Mason River Savanna, Crosby3120 (MO);MasonRiver Field Station,
1 Leaveslong and narrowwith long excurrentcos-
tae, falcateat stem tips. ...... 6b. var. caldensis.

Griffin87 (FLAS);MasonRiverSavannahabout23/4
milesNW of Kellits,Crosby3119 (DUKE);between
Portland
GapandBlueMtn.Peak,Hegewald8107
(hb. 6a. Campylopus arctocarpus var. arctocarpus
Frahm).
Figs. 5E, 27, 28.
BRAZIL.Goias:20 kmN of AltoParaiso,Irwinet
al. 32696 (NY). MINASGERAIS:

Serrade Caraga,Ule
1351 (H-BR, HBG). PARANA:Curitiba,capao da ImCorbuia,Dombrowski5734 (FLAS).Rio DEJANEIRO:
CampylopussellowianusJaeger,Ber. Thitigk. St. Gallischen Naturwiss.Ges. 1877-1878: 385. 1880. DicranumsellowianumHampe,Vidensk.Meddel.Na7633(H-BR),v.Hoehnels.n.

covado,Marzuichelli
(H);
turhist.Foren.Kjoebenhavnser. 3,4:43.1872, nom.
771
Gandeira
Glaziou
Itatiaia
Gavea,
(H);
11744(NY).
scabrisetumHampe.Campro Thysanomitrion
illeg.
NationalPark,Vital240(SP),vonSachsenCoburg
392
pylopussellowiiC. Miillerex Kindberg,Enum.Bryin.
Km 107 along road 282 E
(H-BR). SANTACATARINA:
Type. Brazil.Sellow s.n. (n.v.).
of PonteSerrada,
Frahms.n.(hb.Frahm).SAoPAULO: exot. 51. 1888.scabrisetum
Thysanomitrion
Hampe,Icon. muse. 23B.
Km 221 entreBrotase Itapirina,Vital1454(SP).
1844. Type. Venezuela.Moritz 144a (n.v.).
roraimaeBrotherus,Trans.Linn.Soc. Bot.
Campylopusdelicatulusvar. carolinae(Grout) Campylopus
ser. 2, 6: 89. 1901. Type. Venezuela.McConnell&
Theriot, Mem. Soc. CubanaHist. Nat. 15: 232,
Quelch527 (holotype,H-BR).
1941 has nothingin common with C. angustiretis Campylopusglaziovii (Hampe) Jaeger,Ber. Thatigk.
St. GallischenNaturwiss.Ges. 1877-1878:385. 1880.
and is treated here as its own species, C. caroDicranumglaziovii Hampe, Vidensk. Meddel. Nalinae. Campylopusangustiretisdiffers from all
turhist. Foren. Kjoebenhavnser. 3, 6: 138. 1875.
otherneotropicspeciesof Campylopusby its narType. Brazil.Rio de Janeiro,Glaziou6368 (isotype,
row costa and elongateupperlaminalcells. Since
NY).
the sporophyteof this species is not known, it is Campylopusrigidiusculus(Hampe) Jaeger,Ber. Thatigk.St. GallischenNaturwiss.Ges. 1870-1871:432.
placed tentatively in Campylopusand may ul1872. Dicranum rigidiusculumHampe, Vidensk.
timately have to be transferredto anothergenus.
Meddel. Naturhist. Foren. Kjoebenhavnser. 3, 2:
As in Campylopyuscarolinae, C. angustiretis
272. 1870.Type.Brazil.Sierrade Piedade,Warming
is disjunctbetween SE North America and Bras.n. (isotype, PC).
zil. Also the habitatof these speciesin both parts Campylopusventri-alaris(C. Miiller)Brotherusin Engler & Prantl, Nat. Pflanzenfam.1(3): 334. 1901.
of their rangeis identical.
Dicranumventri-alareC. Miller, Hedwigia39: 251.
Campylopusangustiretishas been considered
1900. Type. Brazil. Minas Gerais, Ule 1347 (holoa variety of C. surinamensis. It resembles the
type, destroyedat B; lectotypusnov., H-BR).
"donnellii"-expressionof C. surinamensisin cer- Campylopuscrispicoma(C. Miiller)Jaeger,Ber. Thatigk.St. GallischenNaturwiss.Ges. 1877-1878: 383.
tain anatomicalcharacterssuch as the long upper
1880.DicranumcrispicomaC. Muller,Flora34: 529.
laminal cells and its habit that has stems ending
1875. Type. Venezuela. Funck & Schlim 366 (hoin comal tufts. However, C. surinamensishas a
lotype, destroyedat B; lectotypusnov., NY).
broader costa (/3 of the leaf base), shorter and Campylopusdiscriminatus(Hampe)Jaeger,Ber. Thawider basal laminal cells, a longer nerve that is
1880. Dicranum discriminatumHampe, Vidensk.
excurrentinto a coarsely serrate awn, a transMeddel. Naturhist.Foren. Kjoebenhavnser. 3, 9verse section of the costa with ventralhyalocysts
10: 254. 1878. Type. Brazil.Rio de Janeiro,Glaziou
slightlylargerthanthe medianrowofdeutercells,
7147 (isolectotype,H-BR).
and more distinct groupsof dorsal stereids.Fur- CampylopuspelichucensisWilliams, Bull. New York
Bot. Gard. 3: 110. 1903. Type. Bolivia. Pelichuco
thermore, the leaves of C. angustiretisare disWilliams2835 (holotype,NY).
River,
keeled
while
the
of
leaves
C.
surinamensis
tinctly
Campylopusrectisetus(Hampe) Jaeger,Ber. Thatigk.
are concave.
DicranumrectisetumHampe,Vidensk.Meddel.Na6. Campylopusarctocarpus(Hornschuch)Mitturhist. Foren. Kjoebenhavnser. 3, 6: 137. 1875.
Type. Brazil.Rio de Janeiro,Glaziou6364 (isotype,
ten, J. Linn. Soc., Bot. 12: 87. 1869. Dicranum
PC).
arctocarpumHornschuch, Fl. bras. 1(2): 12. Campylopus
microthecaHerzog,Beih. Bot. Centralbl.
1840. Type. Uruguay.Montevideo,Sellows.n.
26(2): 55. 1910. Type. Bolivia. CerroAmboro,Her(holotype, destroyedat B; lectotype, BM).
zog 271 ((holotype,JE;isotype, B).
53
4=Q45
i .oS
A
FIG. 27. Campylopusarctocarpusvar. arctocarpus(Sharp3082a, TENN). A. Plant. B. Leaf. C. Leaf-tip.D.
details = 50 Mm.
CampylopusstrictifoliusBrotherus,Act. Soc. Sci. Fenn. bands, ridgedat back, shortlyexcurrentin a ser19(5): 9. 1891. Type. Brazil. Minas Gerais:Caraca, rate point. Alar cells inflated, reddish brown.
Wainios.n. (holotype, H-BR).
Basallaminalcellsincrassate,rectangular,ca. 1:3For synonyms from Africa see Frahm(1985a).
x
Plants in loose, yellowish to dark green tufts,
1-2(-3) cm, rarely to 4 cm high. Stems equally
foliate and slightly comose at tips, reddish tomentose. Leaves 6-9 mm long, narrow lanceolate, erect patent when wet or crisped when dry,
serrate at tips. Costa up to 1/2of the leaf base, in
transverse section with ventral and dorsal stereid
1:6,with pitted walls, 8-16 60-80 .um,shorter

and narrowerat margins. Upper laminal cells
quadrateto subquadrateor shortlyrhomboid,in
distinct rows, incrassate,6-10 x 8-12 Am.
Vegetativepropagationby means of short microphyllous branches in the axils of the comal
leaves.
Seta 8 mm long, light brown. Capsulecurved,
Flora Neotropica
54
1*O
90
70
80
60
so
0o
30
FIG.
bi
28
Distbution
of
Campylopus
''. 'R4
atoarctocarpus
/'
j2
FIG. 28. Distributionof Campylopusarctocarpuss. lat.
slightly strumose, 1.5 mm long, furrowedwhen

emptied. Calyptraciliate.
Distribution(Fig. 28). On rottenlogs, soil covered rocks, and also soil in montane forests to
3000 m, rarely below 1000 m. Usually in rain
forests,in Mexico also in pine, pine-oakand Liquidambarforests. Mexico, Guatemala, Honduras, Costa Rica, Panama, Cuba, Jamaica, Dominican Republic, Colombia, Venezuela,
Surinam, Ecuador, Peru, Brazil, Bolivia, Uruguay, north to Florida.
Also in tropical Africa as ssp. madegassus
(Bescherelle)J.-P. Frahm in Tanzania, Malawi,
Maskarenes, Mauritius, Reunion, Seychelles,
Comores, Tristan da Cunha and MalagasyRepublic.
55 km NE of Tuxtla, Sharp et al. 4271c (TENN). JALsco: 10 miles W of Autlan, Crum 1325 (MICH).
MICHOACAN:
6 miles E of Pulvilla on Hwy 15, Norris
& Taranto 15681 (TENN); Puerto Garnica, Delgadillo
540 (MEXU, NY). OAXACA:Sierra Juarez, 12 miles
above Valle Nacional, Sharp et al. 4522b, 4524 (TENN);
Cima del Cerro Salom6n, 16?46'N, 94?11'W, Ishiki
1275b (NY). TAMAUuPAS:
Above Gomez Farias, Sharp
3633 (TENN). VERACRUZ:
Los Huerfanos near Yecuatla, Sharp et al. 3082a (TENN); 2 km al sur de Coatepec, Guzman 90a (TENN).
GUATEMALA. JALAPA:Alta Verapaz, Standley div.
no. (FH). MORAZAN: Standley div. no. (FH).
QUEZALTENANGO:Volcan Santa Maria, Standley 83421
(NY).
HONDURAS. MORAZAN:
Cerro de Uyuca, Standley
(FH).
NICARAGUA. JINOTEGA: Along Hwy. 3 1.9 km
NW of Aranjuez, Stevens & Krukoff5649 (MO).
COSTA RICA. ALAJELA: San Rafael de San Ra-
m6n, Alfaro22035d (NY); La Fuente,Alfarola (NY);

Specimens examined. MEXICO. CHIAPAS:Lagos de
Montebello, Delgadillo 3483 (MEXU); Yerba Buena
Santiago de San Ram6n, Brenes 21864 (NY). SANJOS:

Cerro Daser area, 9050'N, 84?07'W, Crosby 9842 (MO).
55
SystematicTreatment
PANAMA. PANAMA:Vic. Cerro Jefe, Lewis & Dress-
ler 7604 (MO);CerroMali,Mori & Gentry4377 (MO).

NW of Los Planesde Hornito,Antonio4119B
CHmRQUi:
(MO).
CUBA. ORIENTE:
Pico Turquino,Acunia333 (NY);
Cupeyal,Samek et al. 22 (MO).
JAMAICA. ST. ANDREW:
Sir John's Peak, Britton
1168, Harris 11135 (NY), Crosby3028 (MO);Above
Newhaven Gap, Bengry314 (NY). ST. THOMAS:
Blue
Mountains,Crosby3404 (MO), Rothrocks.no. (NY),
Brittondiv. no. (NY).
DOMINICANREPUBLIC.LAVEGA:
S of La Vega,
Liogier 5 (NY); La Vega, Elliott 691 (FH); La Manaclita S of La Vega,Liogier15842 (NY); BetweenConstanzaandJarabacoa,Allard16516 (NY). Alto de Casbito, Julia &Jimenez 9211-B (NY);Constanza,Allard
16516 (NY).
COLOMBIA. ANTIOQUIA:
Vic. Medellin, Charetier
The ssp. arctocarpusis replaced by the ssp

madegassus(Bescherelle)J.-P. Frahmin Africa,
which differs in smaller size, less pitted basal
laminal cells and a costa being shortlylamellose
in the upperpart of the leaf.
Specimensof this species can be very variable
with respectto size, lengthof the leaf tips, color,
and size of the alar cells, which seems to reflect
the broad geographicalrangeand broad ecological amplitude.
The plantsare usuallycrispatewhen dry, only
the type of C. strictifoliusdiffersby straight,not
crispateor curledleaves. It has been regardedas
a form (Frahm,1979a),but the taxonomicvalue
of this taxon seems questionable.
153 (NY). META:Cordillerade Macarena,Schultes

11208 (NY). VAUPES:
Rio Apaporis, Schultes 11795
6b. Campylopusarctocarpus(Homschuch)Mit(FLAS).
ten var. caldensis (Aongstroem)J.-P. Frahm,
VENEZUELA.AMAZONAS:
CerroYavi, Phelps70a
stat. nov.
ChimantaMassif,Steyermark& Wur(NY). BOLIVAR:
Figs. 28, 29.
dack 1259 (NY). LARA:Parque Nacional Yacombu,
Griffin& Lopez 348 (NY). MERIDA:La Azulita,Stey- CampylopuscaldensisAongstroem,OfversForh.Kongl.
Svenska Vetensk.-Akad.33(4): 5. 1876. Campyloermark97129 (NY). YARACUAY:
E of Ampara,Steypusarctocarpus
Mittenssp.caldensis
ermark 10620 (NY). ZUIA: San Jose de los Altos,
(Hornschuch)
(Aongstroem)J.-P. Frahm, Rev. Bryol. Lichenol.
Griffin102 (FLAS).
45(2): 137. 1979. Type. Brazil.Withoutlocality,
GUYANA. PakaraimaMts., Mt. Latipu, Maas &
Widgrens.n. (holotype,S).
Westra4219 (U).
Campylopusfalcatulus
SURINAM. Lelygebergte,Florschitz 4844 (U).
Bartram,Contr.U.S. Natl.Herb.
26: 64. 1928.Type.CostaRica.Cerrode las Lajas,
FRENCH GUIANA. Montagnede l'Inini, 3?34'N,
Standley 51507 (holotype, FH; isotypes, H-BR, S,
53032'W,Cremers9081 (hb. Frahm).
NY).
ECUADOR. PIHINCHA: Quito-Baeza Hwy., Steere
Campylopuscrispatus(C. Muller)Brotherusin Paris,
& Balslev25115 (NY).
Ind. bryol. Suppl.91. 1900. DicranumcrispatumC.
PERU. CAJAMARCA:
Tambillo, Raimondi 5355 (B).
Miller, Bull. Herb. Boissier6: 36. 1898. Type. BraBRAZIL.BAHLA:15 km N of Leucois,Boom &Mori
zil. Ule 1799 (holotype,destroyedat B; lectotypus
1078 (NY); Serrado Tombador7 km S of Morrode
nov., H-BR).
Chapeu,Irwinet al. 32311 (NY); 23 km N of Seabra,
Irwin et al. 30942 (NY). MINAS GERAIS:
19 km S of Campylopusperfalcatus(C. Miiller)Brotherusin Engler & Prantl, Nat. Pflanzenfam.1(3): 334. 1901.
Padre Paraiso, Vital 5897 (SP). Rio DEJANEIRO:TereDicranumperfalcatumC. Miiller,Hedwigia39: 250.
sopolis, Glaziou6368 (PC);Serrados Orgaos,Glaziou
1900.Type.Brazil.Sta.Catarina.Ule 1124 (holo17979 (PC);Near Abrigo Massena, Vital 4864 (SP);
type, destroyedat B; lectotypusnov., H-BR).
ItatiayaNat. Park,Rose 20509 (NY), Eiten 7652 (NY);
ParqueNacional Tijuca, Pico de Papageio,Landrum Campylopusscopelliformis(C. Muller) Brotherusin
Engler& Prantl,Nat. Pflanzenfam.1(3): 334. 1901.
2209 (MICH). SANTACATARINA:
Campos dos Padres,
C. Miller, Hedwigia39: 248.
Reitz 2464 (NY). Itajai,Reitz 2993 (NY). Sio PAULO: Dicranumscopelliforme
1900. Type. Sta. Catarina:Serrado Oratorio, Ule
Camposde Jordao,north of Capivari,Landrum2810
706 (holotype,destroyedat B;lectotypusnov., H-BR).
(MICH);Serrado Bocaina,Frahm,Camp.Bras.Exsicc.
26 (ALTA,B, C, DUIS, FLAS,GOET,GZU, H, HBG, CampylopussubcubitusWilliams,Bull.New YorkBot.
Gard.3(9): 111. 1903. Type. Bolivia. LowerSorata
INPA, JE, KR, M, MO, NFLD, NICH, NY, PC, S,
River, Williams1749 (holotype,NY; isotypes, FH,
SP, TENN, U); Between Sao Paulo and Santos, Vitt
JE, H-BR).
20646 (ALTA); 28 km NE Sao Paulo-ParanaState
borderalong Hwy 230, Vitt20806 (ALTA);Campos CampylopustablasensisTh6riot,Rev. Bryol.Lichenol.
11: 152. 1939. Type. Bolivia. Rio Tablas, Herzog
do Jordao, Vital9676 (SP);Alto da Serra,Hoehne686
4534a (holotype,PC; isotype, JE).
(JE).
Campylopus arctocarpus is closely related to
C. flexuosus in habit, areolation, presence of microphyllous branches in the axils of the upper
leaves, and the shape of the leaves, but differs in
the transverse section of the costa with ventral
stereids and in the pitted basal laminal cells.
Differs from ssp. arctocarpusby conspicuous

hamate leaves (cf. C. falcatulus, C. perfalcatus),
which are longerand narrowerwith long excurrentcosta. The laminaextendsnearlyto the apex
in ssp. arctocarpus,but vanishesin ssp. caldensis
after 2/3 of the leaf length.
56
Flora Neotropica
1406 (hb. Frahm);ParqueNat. Itatiaia,Griffin& Vital
22.150 (FLAS);Serrado MarbetweenParatiand CunCATAMNA:Serra
ha, Frahm 1405 (hb. Frahm).SANTA
do Espigao30 km NW Dr. Pedrinho,Frahm 1490 (hb.
Alto da Serra, Gehrt499 (JE);
Frahm). SAo PAULO:
Serrado Bocaina, Mun. de Silveira, Vital 7298 (SP);
2 km NE of Pico de Itapeva, Mun. de Pindamonhangaba, Vital 7073 (SP). NE Campos da Cunha,Frahm
1404 (hb. Frahm);Km 277.5 along road from Sio
Paulo to Curitiba,Frahm 1526 (hb. Frahm).
7. Campylopus areodictyon (C. Miiller) Mitten,
J. Linn. Soc. Bot. 12: 77. 1869. Dicranum areodictyon C. Miiller, Syn. musc. frond. 394.
1848. Type. Venezuela. Merida: Sierra Nevada, 9000 ft, Funck& Schlim 1082 (holotype,
destroyed in B; lectotypus nov., NY; isolectotype, H-BR).
Figs. 5B, 30, 31.
CampylopusnitidusWilson in Mitten, Kew J. Bot. 3:
52. 1851. nom. nud. in synon.
Campylopussubconcolor(Hampe)Mitten,J. Linn.Soc.
Bot. 12: 86. 1969. Dicranum subconcolorHampe,
Linnaea32: 138. 1863. Type. Colombia.Andes Bogotenses, La Penna, "in sylvis ad latera Barrancas
2900 m," Lindig 213b (isotype, FH).
CampylopusbreweriBartram,Bol. Soc. Venez.Ci. Nat.
25: 47. 1963. Type. Venezuela.Merida:Sierrade la
Culata,Brewers.n. (holotype,FH; isotype, VEN).
Plants in loose to compact tufts, to 6 cm high,

light yellowish green. Stems erect, whitish tomentose at base, often branched in the upper part
with appressed branches. Vegetative propagation
by means of numerous small microphyllous
branches in the axils of the upper leaves. Leaves
ca. 4 mm long, narrow lanceolate, canaliculate
in the apex. Leaf tip nearly smooth or scarcely
serrate. Costa filling /2--2/3of the leaf width, contracted at base, in transverse section with large
ventral hyalocysts, without dorsal groups of
stereids, slightly ridged at back. Alar cells lacking. Basal laminal cells thin walled, the inner
shortly rectangular, 10-16 x 32-60 ,m, the outer
narrow and elongate, forming a marginal border
of 5-9 cells. Upper laminal cells in the leaf shoulFIG. 29. Campylopusarctocarpusvar. caldensis ders
subquadrate to irregular, incrassate, the up(Frahm 1401, hb. Frahm).Plant.
permost laminal cells short rectangular to oval,
6-13 x 16-32 ,m, ca. 3-4:1.
Seta 5-6 mm long, light brown to dark brown
Distribution (Fig. 28). Costa Rica, Bolivia,
in age, sinuose. Capsule 2 mm long, light brown,
Brazil. At the same habitats as ssp. arctocarpus furrowed when empty. Lid obliquely rostrate, 0.5
in elevations between 1000 and 2000 m.
mm, darker colored. Calyptra ciliate at base.
Distribution (Fig. 31). On humic soil and soil
Km
BRAZIL.
51.5
examined.
PARANA:
Specimens
covered
rocks especially in the Paramo regions,
Frahm
1577
rd.
(hb. Frahm).
along Curitiba-Joinville,
Rio DEJANEIRO:Teres6polis, Serra dos Orgfos, Frahm
also open soil and base of trees and shrubs in the
57
6mm
5cm
FIG.
30.
Campylopus
areodictyon
(Grin
et
47 ,
FAS).
Plant.
B.
Leaf
C.
Leaip.
Transverse
FIG. 30. Campylopusareodictyon(Grifln et al. 477, FLAS). A. Plant. B. Leaf. C. Leaf-tip.D. Transverse
50 ,m.
subalpine belt, from 2600 to 4500 m in Venezuela, Colombia, Ecuador, Peru and Bolivia.
Specimensexamined.COLOMBIA.ARAlCA: Sierra
Nevada del Cocuy, Cleef 8900 (U). BOYACA:
Sierra
Nevada del Cocuy, Alto Valle Lagunillas,Cleef 5708
Subachoque,ParamoEl Tabla(U). CUNDINAMARCA:
zo, Frahm 885244 (COL);Paramode Palacio, 1.5 km
al S de las Lagunasde Buitrago,Cleef4124 (U);Paramo
de CruzVerde ca. 5 km al ENE de LagunaEl Verjon
3365 m, Cleef 3386 (U); Sabanade Bogoti, Schultes
12216 (FLAS);CuchillaEl Tablazo, Linares & Pulla
770 (COLO);Andes Bogotenses, Weir180 (NY); Macizo de Sumapaz,3720 m, Cuatrecasas27059 (NY);

La Vega near Bogota,Apollinaires.n. (NY, PC);Piramo de Sumapaz,Andabobos, Cleef771 (U); Bogota,
Guadeloupe,Onraedt78.6308 (hb. Frahm).
VENEZUELA. MEIRmA:
La Negrita to the Bosqueronof Quebradade las Canas,Luteyn 7838 (NY);
SierraNevada,Aguada,Onraedt78. V.5848,5798, 5799
(hb. Frahm);near Lagunade los Anteojos, Griffinet.
al. 346, 456, 462 (FLAS); between Aguada and La
Montafia,Griffinet. al. 235 (FLAS);bosque de La Vagabunda entre El Morro y Aricagua,Ruiz-Teran &
Flora Neotropica
58
I~nz~
|f
<f<Q
LpzFger
eb
Q
9559
3000m,
andChilifruta,
cayo,betweenHuaytapallana
Hegewald9247 (hb.Frahm).
60
70
80
BOLIVIA. LA PAZ:Prov. Murillo, NE of Milluni,
200 400 600 800 10

,oo00km
4750m,Lewis79-1653,79-1671(F);Wslopeof Cerro
ChekharaN of La Paz, Lewis79-1788A(F); Prov.
NevadoJankoUma,4750m,Lewis79-1558
Larecaja,
(F).
100 200 300 400 500 600 .mil
(F Preparedby HendrikR.Rypkema
/O~~~~~
o
^^'
0"?/
FIG. 31.
^_10
Distribution of Campylopus areodictyon.
Lopez-Figueiras 9559 (FLAS); piramo de Don Pedro

ca. 20 km ESE of El Morro, Ruiz-Terdn 8706 (FLAS).
TACHIRA:Distr. Jauregui, Paramo El Batallon, Grifin
et. al. 574 (FLAS); Distr. Junin, Paramo de Tama above
Villa Paez, Griffin et al. 477 (FLAS).
Base of Volcan SincholaECUADOR. PICHINCHA:
hua, Holm-Nielsen et al. 6621 (GB); Cotopaxi, Quevedo-Latacunga road, 78?56'W, 0?58'S, Holm-Nielsen
et al. 3387 (GB).
PERU. ANCASH:Cordillera Blanca, along road from
Huaraz to Huanuco between Huallanca and Chiquian,
4500 m, Frahm, Camp. Peruv. Exsicc. 3 (ALTA, B,
BM, EGR, F, FLAS, G, GRO, GZU, HBG, KR,
MEXU, NAM, NFLD, NICH, NY, POZ, S, TRT, U);
along road from Catac to Chavin de Huantar E of
Cahuish, Frahm, Camp. Exsicc. 53, 78 (ALTA, B, BM,
BUF, C, DUKE, EGR, F, FLAS, G, GRO, GZU, H,
HBG, KRA, LGHF, MEXU, NAM, NFLD, NICH,
NY, POZ, PC, PRC, RNG, S, TRT, U). JUNiN: Huan-
In anatomical respects, Campylopusareodictyon is relatedto C. nivalis, but is distinguished

by shorter,less apiculateleaves with conspicous
subquadrate(ca. 1.5:1) cells in the basal part of
the upper laminal cells ("leaf shoulder").Macroscopically, it is easily recognized by microphyllousbranchesin the axils of the upperleaves,
which are producedfrequentlyby sterile plants.
Such microphyllousbranchesare found also in
C. albidovirensgrowingin paramohabitats.Both
species are very similarmacroscopicallyand can
be distinguishedonly by the differenttransverse
section of the costa.
Campylopusareodictyonhas often been confusedwith otherspeciesof Campylopus.Material
published from Bolivia (leg. Bridges)and Ecuador (leg. Jameson and Spruce)by Mitten (1869)
as well as many other herbariumspecimens collected in CentralAmerica consist of C. nivalis.
The records from the Juan Fernandez Islands
(Brotherus, 1924) are based on C. fragilis. A
specimen labeled as isotype in KIELconsists of
C. albidovirensand has led to misidentifications.
Fordistinguishingcharacterssee notes underthis
species.
Of the two isotypes, the specimen in H-BR
bears the handwritingof C. Miiller,whereasthe
specimen in NY has been distributed by the
"Leipziger Botanischer Tauschverein." Thereforethe H-BR specimenis proposedas lectotype.
Campylopussubconcolorhas been regardedas
synonymouswith C. leucognodesby Florschutzde Waardand Worrell-Schets(1980), but the latter is treated here as synonymous with C. argyrocaulon,as publishedby Florschiitzand Florschiitz-de Waard(1974).
The morphologicalvariation is influencedby
the humidity of the habitat. In dry habitats the
tufts are very low, producingnumerousflagellae,
and the leaves are rathershort and blunt. In wet
habitats such as dripping cliffs, the plants are
relatively lax and the leaf apices considerably
longer.Apparently,the length of the upperlaminal cells is not correlatedwith the length of the
59
4cm
FIG.32. Campylopusargyrocaulon(Hegewald9171, hb. Frahm).A. Plant.B. Leaf.C. Leaf-tip.D. Transverse

sectionof leaf.E. Basallaminalcells. F. Upperlaminalcells.G(H). Capsule(s).Unlabeledscalebarformicroscopic
details = 50 ,m.
leaf apex and thus not changed in habitats of

differenthumidity.
8. Campylopusargyrocaulon(C. Muller)Brotherusin Engler& Prantl,Nat. Pflanzenfam.1(3):
332. 1901. DicranumargyrocaulonC. Muller,
Linnaea38: 588. 1874. Type. Colombia. Antioquia: Paramo de Sonson, Wallis s.n. (ho-
lotype, destroyed
H-BR).
in B; lectotypus nov.,
Figs. 5C, 32, 33.
CampylopusareodictyonSchimperin Mandon,PI.Boliv. no. 1612, nom. nud.

Campylopusleucognodes(C. Muller)Paris,Ind. bryol.
Suppl. 93. 1900. DicranumleucognodesC. Miller,
Nuovo Giorn. Bot. Ital. n. ser. 4: 32. 1897. Type.
Bolivia. Cochabamba:Germain 1105. (ectotype,
H-BR; isolectotype,NY).
Flora Neotropica
60
BO
70
)\
J
|_
<1
/ Af
200
Seta 6-8 mm long, sinuose, light to dark brown.

Capsule ca. 2 mm long, cylindric, olive-green to
brownish or dark brown in age, furrowed. Annulus darker colored. Lid longly rostrate, 1 mm
long, orange brown. Calyptra ciliate at base.
Distribution (Fig. 33). On wet humic soil in
paramo and wet puna vegetation between 3500
and 4800 m, forming large and dense tussocks,
in wet habitats also on soil covered rocks, rarely
at lower altitudes in subalpine open (Weinmannia, Drimys) forests, also on rocks in wet seepy
cliffs. Venezuela, Colombia, Ecuador, Peru, Bolivia.
60
400
600
800
1000km
O100 200 300 400 500 600 mil
dPreparedby HendrikR.Rypkema
----20
L/
4>4
10
Specimens examined. COLOMBIA. ARAUCA:Sierra

Nevada del Cocuy, Cleef8893 (U). BOYACA:Paramo
de la Rusia NW de Duitama, Cleef7329 (U). CUNDINAMARCA:Represa de Neusa, Cleef4169a (U); Paramo
de Palacio, Cleef 3812 (U); Paramo de Cruz Verde,

Cleef 3146 (U); Piramo de Sumapaz, Chisaca, Cleef
4910 (U); Paramo de Sumapaz, Cuatrecasas27057
(NY). Cordillerade La Leonora60 km NNE de Bogota,
v. d. Hammen & Jaramillo 3211 (U); Paramode Sumapaz,LagunaCurucues,Mdgdefrau1366 (hb.Frahm).
CAUCA:Parque Nacional del Purace, v. d. Hammen &

Jaramillo 3642 (U). META:Paramo de Sumapaz, Cleef
Mun. de Sta. Rosa de Cabal,
1148 (U). RISARALDA:
f the leafwidth, excurrent,in transversesection
valle de la Quebrada,van Reenen & Cleef 896 (U).

CordilleraCentral,Swale, Pennell 12115 (NY).
Paramode Don Pedro 20
VENEZUELA. MERIDA:
km al ESEde El Morro,Ruiz-Teran& Lopez-F. 8706
(FLAS);Distr. Miranda,above Pico del Aguila,Griffin
et al. 1358 (FLAS);Alrededoresde la Lagunade los
Patos,distr.Rangel,Badilllo&Pdez 6827 (MY);Sierra
de Santo Domingo, Paramode Mucubaji,Griffinet al.
1286, 1209, 1039 (FLAS);Sierrade Merida, Laguna
de Los Anteojos, Griffinet al. 278, 300 (FLAS).
ECUADOR. Tunguragua,Spruce 52 (NY). Quito,
Jameson s.n. (NY); PICHINCHA:21 km E of Pito on
roadto Baeza,0?23'S,78?13'W,Churchill& Sastre-De

Jesus 13522 (NY).
PERU. ANCASH:E side of tunnel Cahuish, Hegewald
Las Lagunas, HegePlants large, to 13 cm high, in compact tufts, 7688 (hb. Frahm). CAJAMARCA:
Cuzco: Between
wald
6165
6164,
6191,
Frahm).
(hb.
at
reddish
whitlight green tips, densely
(rarely
Urcos and Juliaca along road from Puno to Cuzco,
ish) tomentose below. Stems erect, unbranched. Frahm, Camp.Peruv.
Exsicc. 4 (ALTA, B, BM, EGR,
Leaves
8-9
mmBasal
narrowlanceolate,
FIG. 33.
Distinct
cells thin-wgyrocaulon.led,
long, laminal
ending F, FLAS, G, GRO, GZU, HBG, KR, MEXU, NAM,
group.
in a long, almost smooth tip. Costa filling half NFLD, NICH, NY, POZ, S, TRT, U). JuNiN:30 km
ofrPlants
the leaflarg,
in0:,
transverse
section
toexcurrent,
13 cm high
in compact
tufts, E of Huancayo,Hegewald9171 (hb. Frahm).
width,
BOLIVIA. Without locality, Bridges s.n. (NY).
with
ventral
and dorsal
light thick-walled
reddish (rarelongay
greenatmargtips,
denselyhyalocysts
whit- COCHABAMBA:
area of Laguna Tarucani, Lewis 79-2465
smooth
atStems
back. erect,
Lamina
tomentosine
below.
formish)
unbranched.
substereids,
vanishing (F); E face of CerroChua LagunaW of Tambo Pata,
below
midleaf.
cells
x a Lewis 79-2219A (F). LAPAZ:1 km S of PuertoAcosta
mmAlar
Leaves
oval8-9
narrow
6ending
long,
lax, lanceolrassate,
hyaline, forming
distinct
Basal
laminal
just N ofLago Titicaca,Lewis 79-749 (F);Head of Rio
in a long,
almost
smooth
Costathin-walled,
tip. cells
half6:
filling
group.
N of La Paz, Lewis79-1812B (F);Nevado Jankrectangular,6-16 x 51-79 /m, ca. 4-10:1, in 7- Zongo
ho Uma above Mina St. Antonio, Lewis 79-1576A,
12 rows at margins thin-walled and elongate, 1586
(F).
forminga distincthyalineborder.Upper laminal
cells oval to elongateoval, incrassate,3-6 x 10Campylopus argyrocaulon has often been confused with other species of Campylopus. As syn19 tm,ca. 4-6:1.
61
FG34C
FIG. 34.
Campylopus asperifolius (Lewis 792260, F).
types of this species (Mandon 1612, 1613) have

been distributed as Campylopus areodictyon
Schimper nom. nud., non (C. Miiller) Mitten,
specimenswere often named C. areodictyonand
are filedin herbariaunderthis species. Robinson
(1967) supposedthis species to be identicalwith
C. pittieri Williams, and Florschiitz-deWaard
and Worrell-Schets(1980) placed C. leucognodes
in C. subconcolor.C. subconcolor,however, is
regardedhere as a synonym of C. areodictyon
and C. leucognodes of C. argyrocaulon,as in
Florschiitzand Florschiitz-deWaard(1974).
The speciesis easily recognizedby its largesize

and long leaves, which end in a long, smooth or
finelyserratetip. Both charactersare in common
with C. cavifolius,which is, however,even larger.
Small forms of C. cavifolius are distinguished
from C. argyrocaulonby the costa contractedat
base, which is ill defined from the lamina, and
by large,lax ventralhyalocystsin transversesection of the costa.
The appearanceof this species varies between
(a) slender plants with appressed foliate stems
growingin dense tufts on wet soil, and (b) laxly
62
Flora Neotropica
foliate plants in loose tufts with more spreading

leaves growingin subalpineforests and also on
moist rocks and dripping cliffs. The latter has
been usually been named C. leucognodes, although there are no anatomicaldifferences.
ReSpecimens examined. COSTA RICA. CARTAGO:
fugio Tapanti,Arocha 1194, 1141 (hb. Frahm).SAN

JOSE:La Hondura, Valerio 261 (NY); Las Nubes,
Standley38754 (NY); Cordillerade Talamanca,Cerro
de la Muerte,Eggers CR 6, 3 (hb. Frahm).
COLOMBIA.BOYACA:
Pefiade ArnicalN de Vado
Hondo, Cleef9495 (U). CUNDINAMARCA:
Subachoque,
9. CampylopusasperifoliusMitten,J. Linn. Soc.

Bot. 12: 79. 1869. Dicranodontiumasperifolium (Mitten) Brotherus, in Engler & Prantl,
Nat. Pflanzenfam.ed. 2, 10: 190. 1924. Type.
Ecuador. Abitagua, 7000 ft, Spruce 54 (lectotype, PC;the originalmaterialin the Mitten
herbarium,NY, is lacking).
Figs. 34, 35.
CampylopustrichophorusHampe ex Herzog,biblioth.
Bot.87: 20. 1916.Type.Ecuador.

Loja:Krauses.n.
(n.v.).
weddeliiBescherelle,
J. Bot. (Morot)5:
Campylopus
147. 1891.Type.Peru.Carabaya:
Weddels.n. (isotype,BM).
Plants to 10 cm high, in loose tufts, yellowish
to brownish,conspicuouslyglossy golden green.
Stems laxly foliate with distant, spreading,flexuose or slightly homomallous leaves, often comose at tips, not tomentose. Leaves 6-7 mm
long, from ovate base graduallylanceolate,ending in a long, slender, smooth tip. Costa filling
one halfofthe leafwidth, excurrent,in transverse
sectionwith largeventralhyalocysts,medianand
dorsal chlorocysts,without dorsal stereids.Alar
cells large,inflated,protrudingconspicuouslyinto
the costa. Basallaminalcells rectangular,slightly
incrassate,elongate oval upper laminal cells, 36 x 26-42 gm.
Sporophytespseudolateral.Seta to 1 cm long,
twisted, light brown to blackish in age. Capsule
erect, symmetric, olive-greenin young capsules
to brownish in mature capsules and blackish in
old capsules, furrowed.Annulus blackish. Lid
obliquelyrostrate,1 mm long, reddishto reddish
brown or blackish,darkercolored than the capsule. Calyptranot known.
Vegetativepropagationby meansof smallpropaguliferousleaves producedin the axils of comose leafs at stemtips.
Distribution (Fig. 35). Epiphytic on bark of
trees, branches and even small branchlets in
cloud-forestsand subalpineforests, on nodes of
Chusqueain bamboo scrubsfrom 1400-4000 m,
usuallybetween2600 and 3300 m in CostaRica,
Venezuela,Colombia, Ecuador,Peru and Bolivia.
ParamoEl Tablazo,Frahm885245 (COL);Cordillera

de la Leonora60 km NNE of Bogota, v. d. Hammen
& Jaramillo3781 (U); Paramode Guasca 50 km NE
of Bogota, 3300 m, Sipman et al. 10612 (COLO,U);
Sabana de Bogota, Piramo de Cruz Verde, Schultes
12209 (FLAS).HUILA:
CordilleraCentral,Paramode
las Papas, Bishop s.n. (FLAS). MAGDALENA:
Sierra Ne-
vada de SantaMarta,Rangel et al. 394, 289 (FLAS).

VENEZUELA.Paramode Bocon6,Griffinet al. 1062
Paramo El Batallon, Griffin& Du(FLAS).MERIDA:
garte PV 1097 (NY); Paramo de Pozo Negro, RuizTerdn 6575 (FLAS). TACHIRA:
Jauregui,Paramo El
Batall6n,Griffin& Ruiz-Terdn466 (FLAS).
ECUADOR.LOJA:CajanumaSE ofLoja, Laegaard
536236 (NY); between Loja and Saraguro,Steere &
Balslev 25898 (NY); MORONA-SANTIAGO:Gualaco-Suc-
E of San Gabriel,
ua road, Steere27775 (NYJ).NAPO:
Steere 9139f(NY); E side of CerroSumaco, 77?39'W,
0?36'S,Lojtnant& Molau 13058 (NY).
PERU. AYACUCHO: 21 km vom Pass Huamina in
RichtungAyna, Hegewald9005 (hb. Frahm).

La Paz, Laguna Huichin-
cani, 67017'W,17054'S,Lewis 87484 (MO);E face of

CerroChuaLagunaW ofTambo Pata, Lewis 79-2260
(F). An Baumastender WaldgrenzeiiberTablas,Herzog 2858 (JE).
This species is morphologicallyand anatomically differentfrom all other species of Campylopus by the distant, lax, flexuose leaves and
by the laminal cells, which are not sharply differentiated between basal and upper laminal cells.
Probably for these differences, Brotherus placed
this species in Dicranodontium. A final decision
concerningthe generic placement was not possible until recently, as no sporophytes were
known. Recently, however, it has been found
once with sporophytes(Lewis 79-2260) in Bolivia. This specimen indicates clearly that the
species has to be placed into Campylopus.
As in othergeneraof the Campylopodioideae,
epiphytic species are rareand thus it can be assumed that these speciesarethe youngestbranch
of evolution.
Campylopus asperifolius was described by
Mitten by two syntypes, one collected by Spruce
in Ecuador, which has been chosen as lectotype,
the other by Lechler (no. 2627) in Tatanara, "An-
des chilenses,"which is probablysituatedtoday

in Peru. The latter specimen was labelledin the
herbariumof C. Muller as "Campylopuspeni-
63
cillatus" and was used by Herzog as a syntype
of Campylopustrichophorus,not knowing that
exactly this specimen had alreadybeen used by
Mitten for the descriptionof C. asperifolius.
Campylopusasperifoliusis characterizedby its
epiphytic occurrence,its metallic, glossy color,
the lax, distantand spreadingleaves and the presence of propaguliferousleaves in the axils of the
uppermost leaves. In anatomical respects it is
conspicuousby the alar cells protrudinginto the
costa and elongate cells throughout the whole
lamina.
90
70
80
-b- ..
----0
_....
60
.. 200
..
____.
--
"
- '~
.
,
~)C>
00
000
00
000m
200 300 400SOO0600,,
Prepared by Hendrik R. Rypkem
Ii
20
10. CampylopusbryotropiiJ.-P. Frahm, Nova

Hedwigia39: 152. 1984. Type. Peru. Ancash:
CordilleraBlanca, Laguna Llanganuco 3850
m, Frahm 825118 (holotype, B; isotypes, C,
EGR, F, FLAS,G, GRO, GZU, H, HBG, KR,
KRA, M, MEXU, NAM, NFLD, NY, PC, S,
U, UPS, USM).
Figs. 36, 37.
Plants to 4 cm high, dark to yellowish green,
often brownishbelow and lighterabove, in loose
tufts. Stems equally foliate, tomentose below;
rhizoids papillose. Leaves 6-7 mm long, erect
patent when wet, more or less appressedwhen
dry, and lanceolate. Costa filling half of the leaf
width, excurrentin a nearly smooth or scarcely
dentate awn, which may be hyaline or (rarely)
concolorous, in transverse section with ventral
hyalocysts and a dorsal layer of stereids. Alar
cells inflated,hyaline or reddish,usuallyreddish
inside and hyaline outside, protrudinginto the
costa. Innerbasal laminalcells elongate,hyaline,
shortrectangularto oval, 6-10 x 19-54 ,m, the
marginalcells in 8-10 rows very narrow.Upper
laminalcells incrassate,oval to shortlyelongate,
with thickened corers, 3-5.5 x 20-35 ,um(410:1).
Distribution(Fig. 37). On wet rocksand boulders, on wet ground in paramos, around ponds
and in mires, in elevations from 3500 to 3730
m in Venezuela,Colombia and Peru. The range
is not yet sufficientlyknown, since this species
has been describedonly relatively recently.It is
supposedto be an andine element and is to be
expectedalso in Ecuadorand Costa Rica.
Specimensexamined.COLOMBIA.CUNDINAMARCA:Cabecerasde la QuebradaChusaentre Paramode
Palacioet Paramode Chingaza,Cleef5336b (U).
VENEZUELA. MERIDA: Distr. Rangel, SierraNe-
FIG. 35. Distributionof Campylopusasperifolius.

vada de Santo Domingo, Paramode Mucubajialong
roadto LagunaNegra,Fransen1246, 1247, 1432, 1451
(GB);SierraNevada de Merida,Poelts.n. (hb. Frahm);
Barclay9586 (MICH);Teleferico,Cleef&Huber4808
(U); Paramode Santo Christo, 20 km al S de Mucuchies,Ruiz-Terdn8489 (FLAS);ParamodeTamaabove
the village of Paez, Griffinet al. 322 (FLAS). Distr.
Libertador,Lagunadel SantoCristo,Ruiz-Terdn8491
(FLAS).
PERU. ANCASH:
Parque Nacional Huascaran,Laguna Llaganuco,Frahm 823903 (hb. Frahm).
Campylopus bryotropii is related to C. cuspidatus by the hyaline tipped leaves and the shape
of the upper laminal cells and therefore specimens were named C. cuspidatus before C. bryotropii was described. The latter is, however,
smaller, has a narrower lamina, hyaline, not pitted basal laminal cells and thin-walled, not collenchymatous alar cells.
11. Campylopus capitulatus Bartram, Bull. Brit.
Mus. (Nat. Hist.) Bot. 2: 52. 1955. Type. Ec-
Flora Neotropica
64
\ VG
.F'
2cm
'
, ij ,)
i ) i
*O
?n0?
jf ?Jj)
if
E
FIG. 36. Campylopus bryotropii (Frahm 823903, hb. Frahm). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
section of leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule. Unlabeled scale bar for microscopic
details = 50rm.
in transversesection with ventralhyalocystsand

dorsal groups of (sub)stereids.Brood leaves 2
mm long, narrowlanceolate,with poorly developed lamina, contracted at base, often hooked
Plants to 5 cm high, yellowish green, in loose or curled.Alarcells hardlydeveloped.Basallamtufts. Stems appressed foliate, abruptly ending inal cells hyaline, thin-walled, rectangular,16in a comose tip consistingof long, flexuoseleaves 26 x 45-80 gm, narrowerat margins. Upper
with numerous small, heteromorphic brood laminalcells sharplydifferentiatedfrom the basleaves. Leavesabout 5 mm long, lanceolate,from al ones, incrassate, 8-13 x 13-19 Am, nearly
ovate base graduallycontractedto a serratetip. quadrate.
Costa filling '/2 to 2/ of the leaf width, excurrent,
uador.Chaupi-Sagcha,Pululagua,on a stump,
Bell 591 (holotype, FH; isotype, BM).
Figs. 38, 39.
65
80
70
60
50
40
fQ~~~~~~~
^
^
FS^
?
O^^
o ?0
<0
.
>
200
0
<
400
600
800 1000km
100 200 300 400 500 600 miles
Prepared by HendrikR.Rypkema
o~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.....
FG37Ditbtino
aplpsbytoi(0an0dC
aoie(*
;~~~-?-?-?~~~~~~~~.-?-?-.-?'
FIG. 37.
FI.37
(L of Campylopubryotropii
--Distribution
YI/~~~~~~ (0) and C. carolinae(@).10
isrbuin
fCmplpu
Distribution (Fig. 39). On rotten wood, logs

and stumps, rarely epiphytic in montane rainforests, cloud forests and subalpine forests
through the Andes from Venezuela, Ecuador and
Peru to Bolivia, so far not known from Colombia, in elevations from 2000 to 3500 m.
Specimens examined. VENEZUELA.
TACHIRA:
ParamoEl Batallonabove the town of La Grita,Griffin

et al. 550 (FLAS).MERIDA: Campo Elias, La Carboneraarea,fincaSanEusebio,Griffinet al. 1588 (FLAS).
ECUADOR.NAPO:Road betweenTena and Baeza,
Steere E 135 (NY). PICHINcHA:Old road from Quito
to Sto. Domingo, 24 km W of Chiriboga,Brako 4737

(NY); 20 km W ofLloa, Frahm 144, 145 (hb. Frahm).
PERU. AMARONAS:
Hornopampa bei Balsas, Hege-
wald 6675 (hb. Hegewald).Cuzco: Pisac, Hegewald

8650 (hb. Hegewald);MachuPicchu, Hegewald8826,
rytopi()
n Ccroiae()
8860 (hb. Hegewald).JuUNv: Huasahuasibei Tarma,
Hegewald8355 (hb. Hegewald);Muna,Bryan489

(MO).
BOLIVIA.COCHABAMBA:
Antahnacana,7 milesSSW
of Locotal,Hermann25228a(hb.Frahm).
Although describedin 1955, this species had

alreadybeen found in Peru in 1923 (Bryan489,
MO), but erroneouslytaken by Williams for C.
gracilicaulis(C. surinamensis).Indeed,it is morphologicallysimilar to C. surinamensiswith appressed foliate stems and apical comal tufted
leaves, but it is distinguishedfrom the latter by
nearly quadrate(not rectangular)upper laminal
cells and the habitat on rotten wood at high elevations, whereas C. surinamensisis a lowland
species growingon sandy soil.
Flora Neotropica
66
if
I
,/C
'' C
n^0 C
F
4mm
4cm
FIG. 38. Campylopus capitulatus (Bell 591, FH). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section of
leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details = 50 Mm.
Campylopuscapitulatusis closely relatedto C.

fragilis with respectto the presenceand shape of
brood leaves, the lax, hyaline basal laminal cells
and the incrassate, subquadrateupper laminal
cells and the structureof the costa, but is distinguishedby the conspicuousappressedstem leaves
and comal tufts. Thus, the species are thoughtto
be phylogeneticallyrelated. Since C. fragilis is
confinedin the Andes to the alpineand subalpine
belt, C. capitulatusmay be interpretedas an an-
dean rainforestraceof C.fragilis. Althoughmorphologically very closely related, both species

were found in a mixed tuft (Frahm 144, 145) in
Ecuadorat 2800 m elevation, which shows that
they are geneticallydistinct.
Another species, occurringin SE Asia, resembles C. capitulatus:C. laxitextus Lac. It is distinguished only by long, subhyaline excurrent
costae and longer, short rectangularupper laminal cells.
67
12. Campylopus carolinae Grout, Moss fl. N.
Amer. 1: 249. 1939. Type. U.S.A. North Carolina: Brunswick Co., near Southport, Anderson
& Evans 6180 (holotype, DUKE). Figs.-----' 8C,
37, 40.
60
70
80
>
200
40
60
60
0800000km
100 200 300 400 500

6 00 mil
Campylopus cerradensis Vital, Cryptogamie Bryol.

Lich6nol.5: 20. 1984. Type. Brazil.Sao Paulo:Mun.
de Itirapina,Vital 1714 (holotype, SP).
Plants dark green to brownish green or blackish, to 1 cm, rarely to 2 cm high, in loose tufts.
Stems with appressed or erect patent leaves.
Leaves lanceolate, 2.5-4 mm long, the margins
entire and convolute in the upper part. Costa 1/3
of the leaf width, excurrent in a hyaline point,
in transverse section with dorsal stereids and a
small group of ventral stereids in the median part
of the leaf. Alar cells lacking. Basal laminal cells
rectangular, incrassate, 13-19 x 35-50 Mm.Upper laminal cells oblique to oval, incrassate, 56 x 19-25 jm.
Capsule globose, ca. 1 mm in diam., scabrous
at base, shortly rostrate. Seta 2-4 mm long,
curved. Calyptra long-ciliate at base.
Distribution (Fig. 37). Very typically buried in
loose sand in Cerrado vegetation in Brazil at altitudes up to 1500 m, also in open grassland,
open pine or pine-oak forest of the coastal plain
in SE North America in Florida and North Carolina.
FIG. 39. Distributionof Campylopuscapitulatus.
rate subgenus,based on sporophyticcharacters,

was not recognizedbefore 1984. Spore dispersal
is presumablynot very successfuldue to the imSpecimens examined. BRAZIL. BAHIA:Lagoa de mersedcapsules,but this species (as also C. perAbaete, Boom et al. 882 (NY). MATOGRosso: Mun.
means of deciduousstem
de Brasilandiaca. 16 km SE ofXavantina, Vital2195 pusillus)propagatesby
some advantage in
have
to
which
seems
tips,
BR
135 from
Km 67 along road
(SP). MINASGERAIS:
Curvelo to Diamantina, Frahm 1441 (hb. Frahm). colonizing xeric pioneer habitats.
PARA:Serrado Cachimbo, km 780-820 on CuiabaSantar6m highway (BR 163), Reese 16335, 16339
Mun. de Brotas,ca. 16 km W of
(LAF). SAo PAULO:
Itapirina,Vital2200 (SP);ca. 8 km E-NE of Itapirina,
Vital 1434, 1508, 1714, 1742, 2065 (SP).
This species is the only representative of the

subgenus Campylopidulum Vital in the Neotropics. A second species, C. perpusillus Mitt.,
occurs in tropical Africa.
Such disjunction between Brazil and southeastern North America is also found in Campylopus angustiretis, which also occurs on sandy
soil. It may be interpreted as relic of a larger,
continuous range in former, drier climatic periods.
Sporophytes are produced extremely rarely and
therefore the necessity for placement in a sepa-
13. Campylopuscavifolius Mitten, J. Linn. Soc.

Bot. 12: 87. 1869. Type. Ecuador."AndesQuitenses," Chimborazo, Jameson s.n. (lectotypus nov., NY).
Figs. 41, 42.
Theriot,Arch.Bot.2(10):187.
apollinairei
Campylopus
s.n. (ho1928.Type.Colombia.Bogota,Apollinaire
lotype,PC;isotype,NY).
One of the largestspeciesof Campylopus,plants
to 10 cm high or more, yellowish green above,
brownish below, in wet habitats darkercolored
to even blackish, in tufts. Stems equally foliate
with erect spreading leaves, tomentose below.
Leaves 10-12 mm long, narrowlanceolate,from
an ovate base graduallycontracted into a very
long, fine, smooth tip, the narrowleaf tip as long
Flora Neotropica
68
C
1cm
[
E
FIG. 40. Campylopuscarolinae (Vital 1714, SP). A. Plant. B. Leaf. C. Leaf-tip. D. Transversesection of
50 Am.
as the basal part of the leaf. Costa at leaf base ill

defined, graduallyintergradinginto the lamina,
excurrent,in transversesectionwith largeventral
hyalocysts,taking half or more of the thickness
of the leaf, withoutdorsalstereids,graduallynarrowed into the lamina, ridgedat back in the upper partof the leaf. Alar cells weaklydeveloped,
hyaline. Basal laminal cells rectangular,thinwalled, 10-13 x 19-45 Am,elongateand narrow
at margins.Upper laminal cells oval to elongate
oval, incrassate,6-10 x 22-45 ,m, ca. 4-6:1.
Sporophytepseudolateral.Seta 15 mm long,
curved, brownish. Capsule 2 mm long, light
brown to dark brown in age, erect, symmetric,
cylindrical, furrowed when empty. Calyptra
fringedat base.
Distribution(Fig. 42). Terrestrialin paramos,
often in largetufts in wet depressionsand around
ponds, often associated with Sphagnum spp.,
rarely on wet rocks, in the drier northern and
southern part of its range in subalpine shrubs
and forests, in Panama, Venezuela, Colombia,
69
i li
FIG. 41. Campylopus cavifolius (Cleef7278, U). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section of leaf.
E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details = 50 Mm.
Ecuador, Peru and Bolivia, in elevations between

3400 and 4300 m.
cizo alto N de Belen, Cleef 9803 (U); SierraNevada

del Cocuy, Barclay 7439, 7448 (MICH); Alto Valle
Lagunillas,Cleefet al. 10386 (U); Alto de la Cueva,
Cleef 9982 (U); Piramo C6ncavo, Cleef 8573, 8678,
8523, 8642 (U);QuebradaBocatoma,Cleef8753, 8760
(U); QuebradaBocatoma,Cleef8751, 8741, 8739 (U);
N-NW de Duitama,Cleef7293, 7296, 7354 (U); Cerro
Nevado del Sumapaz, Cleef 1282 (U); Piramo de la
SarnaentreSogamosay Vado Hondo, Cleef9398b(U).
CAUCA:
Volcan Purack,Lagunade San Rafael,Cleef&
Fernandez 537 (U); valle Rio Vinagre, Cleef & Fer-
Specimensexamined.PANAMA. Bois de Boquete

near David, FrereHelion s.n. (S).
COLOMBIA. ARAUCA: Sierra Nevado del Cocuy,
QuebradaEl Play6n, Cleef 8976, 9150, 10066 (U);
Gradstein,Bryoph.Neotrop.Exsicc. 55 (U). BOYACA:
Paramode la Rusia, Cleef 7278 (U); Paramode Pisva,
Cleef4413, 4506a (U);carreteraSocha-LaPunta,Cleef
4330,4333,4335,4338,4340,4343,4346,4416, 4506b,
4643 (U); al N del Alto de Granados,Cleef 4580 (U); nandez 658 (U). CUNDINAMARCA:
Piramo de SumaParamode la Rusia, Cleef 7516, 7357, 7274 (U); Ma- paz, Chisaca,Cleef3583 (U); LagunaNegra,Cleef190
Flora Neotropica
70
70
80
and can hardlybe distinguishedmacroscopically.
60
Campylopus pittieri is usually paler (leucobry-
...
?-'^^-/
Prepared by Hendrik R.Rypke.ma
FIG. 42. Distributionof Campylopuscavifolius.

(U); Cabecerasdel rio S. Rosa, Cleef5268, 5269, 5284
(U); Media Naranja,Cleef& Jaramillo232 (U); Paramo de CruzVerde, Cleef 3383 (U); Andabobos,Cleef
771, 1684 (U); Paramo entre Cogua y San Cayetano,
Cleef6539 (U); Cabecerasde la QuebradaChuza,Cleef
& Florschitz 3640 (U); Paramo de Palacio, Cleef &
Uribe6714, 6735, Cleef& Jaramillo 4029, Cleef 317,
4032, 5157 (U); Cabecerasde la QuebradaChuza,Cleef
5336, 9631 (U); Lagunasde Buitrago,Cleef278, 4092
(U);Paramode Sumapaz,VallequebradaHonda,Cleef
1560 (U). META:Paramode Sumapaz,CerroNevado
de Sumapaz,Cleef1282, 1283, 1361, 1430, 7701, 8137,
8207 (U); Hoya El Nevado, Cleef 864 (U); Hoya Sitiales,LagunaLa Primavera,Cleef7580 (U);Quebrada
El Buque,Cleef 7793 (U); CerroNevado de Sumapaz,
Cleef1252a (U); LagunaLa Guitarra,Cleef863, 848,
849, 8280 (U).
Paramode La Negra,RuizVENEZUELA.MERtDA:
Terdn8401 (FLAS);Sierrade SantoDomingo, Paramo
de Mucubaji,Griffinet al. 1083 (FLAS).
Volcan Chiles km 34-36 TulECUADOR.CARCHI:
can-Maldonadoroad, Holm-Nielsenet al. 5858 (GB).
oid), especially when dried up, and has shorter

leaf tips. MicroscopicallyC. cavifoliusis distinguishedby smallerventralhyalocystsof the costa
and a differenttransverse section of the costa.
This demonstratesthat species of differentphylogenetic origin have occupied the same ecological niche and have developed homologous adaptations such as growth in cushions, dense
tomentum, largeventral hyalocystsin the costa,
hyalinebasal laminal cells or oval upperlaminal
cells.
14. Campylopuscleefii J.-P. Frahm, Cryptogamie Bryol. Lichenol. 7: 441. 1986. Type. Colombia. Boyaca: Paramo La Rusia, "subparamo cerca del puente del Rio Surba," Cleef
6269 (holotype, U).
Figs. 43, 44.
Plants 0.5 to 5 cm high, yellowishgreenabove,
dark green to blackish below. Stems interruptedly foliate, the leaves erect patentwhen wet but
appressedwhen dry. Leaves 4-5 mm long, lanceolate, subtubulosein the upper part, with entire margins.Costa filling half of the leaf width,
excurrentin a serrate, sometimes at the outermost tip hyaline awn, in transversesection with
ventraland dorsalstereids,smooth at back. Alar
cells hyaline and thin-walledor reddish and incrassatein older leaves. Basal laminal cells rectangular,2-6:1, incrassateand slightlyporose, 610 x 26-42 Atm,at marginsin 2-8 rowselongate.
Upper laminal cells irregular,short rectangular
to oblique or rhomboid,3-10 x 10-19 Am, (2-)
4(-6): 1, incrassate.
Seta 1.5 mm long, curved.Capsule1 mm long,
globose.
Distribution(Fig. 44). Terrestrialin paramos
between 3300 and 3875 m, rarelyup to 4400 m,
found only in Colombia.
Specimens examined: COLOMBIA. BOYACA: Par-
amo de la RusiaNW-N de Duitama,Cleef6797, 6813,

Balslev8661 (NY). NAPO:Road Quito-Baeza,Paramo 6819, 7238, 7251, 7285, 7422 (U); CarreteraSochade Guamani, Ollgaard& Balslev 10062, 10161 (NY). Los Piiios km 41, Cleef9879 (U); ParamoentreCogua
PERU. ANCASH:Cordillera Blanca, Tunnel Cahuish,
y SanCayetano,Lagunaverde,Cleef6355 (U); Paramo
al NW de Belen,cabecerasQuebradaMinas,Cleef1886,
Hegewald7688 (hb. Hegewald).
BOLIVIA. COCHABAMBA:Cerro Chua Laguna W of
1895 (U); Piramo de Guina, Sta. Rosita, Cleef 9754
TamboPata,Lewis79-2276A (F);Rio Sanjana,Herzog (U); SierraNevada del Cocuy, Alto Valle Lagunillas,
3299 (NY).
Cleef 5698 (U); vereda S. Jose de la Montafia, Cleef
Road Ibarra-Mariano Acosta, Ollgaard &
IMBABURA:
Campylopus cavifolius is often associated with
C. pittieri.Both species even growin mixed tufts
Paramo de Sumapaz cerca

1766 (U). CUNDINAMARCA:
de Lagunitasal S. de Juan, Cleef 8410 (U); Piramo

entreCoguay SanCayetano,LagunaVerde,Cleef6355
71
SystematicTreatment
1Il
WE
^-^B
W)'*/
i|M
\
(U). META:Paramode Sumapaz,Hoya de la Quebrada even hybridogencapsuleshave been observedin

Sitiales, Cleef893 (U).
this genus.
Campylopus cleefii resembles C. thysanomitroides in the areolation, the dark color and the
habitat in paramos, but is distinguished by the
comose foliation and ventral stereids in transverse section of the costa.
A capsule has been found only once. It is unusually globose and has an unusually short seta
and is perhaps not representative, as aborted or
The variation observed concernsthe size and

color of the plants,the developmentof alarcells,
the lengthof leaf apices and the lengthoflaminal
cells.
15. Campylopusconcolor(Hooker)Bridel,Bryol.
univ. 1: 476. 1826. DicranumconcolorHooker, Musci exot. 139. 1816. Type. Colombia.
72
Flora Neotropica
80
70
60
50
40
0d
<k
^/^
I\
o~
^s^
?~~
0
0
200
400
600
800
1000km
100 200 300 400 600 600 miles
Prepared by Hendrik R.Rypkema

FIG.
10. Distribution
of
Campylopus
cleefti
(*)
and
C.
controversus
(0).
01~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
oy
I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.d
~~~~~~~~~~~~~~~~~~~~~~~~~~~
........
FIG. 44. Distributionof Campylopuscleefii(W and C. controversus(0).
Alto de Aranda, Humboldt & Bonpland s.n.

Figs. 45, 46.
(isotypes, H-BR, KIEL).
Plants to 9 cm high,in green,loose tufts.Stems
reddishtomentose, equally foliate with erect patent leaves, often comose at tips. Leavesto 9 mm
long, from ovate base graduallynarrowedinto a
long, serrate tip. Costa filling
l/22/3
of the leaf
base, excurrent,the excurrentpart half as long

as the leaf, in transversesection with ventralhy-
alocystsand dorsalgroupsof stereids,smooth or

(in the upper part) slightly ridged at back. Alar
cells well developed, hyaline or reddish, protrudinginto the costa. Basallaminalcells rectangular,incrassate,10-16 x 32-48 gm, narrowand
elongate at margins. Upper laminal cells
subquadrateto short rectangular,small, opaque,
5-6 x 10-15 ,m, in distinct rows.
Sporophytes2-5 aggregatedin the comalleaves.
Seta 8-9 mm long,brownish,twistedand curved.
73
2mm
A
FIG. 45. Campylopusconcolor(Hegewald8762, hb. Hegewald).A. Plant.B. Leaf.C. Leaf-tip.D. Transverse
section of leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule.Unlabeled scale bar for microscopic
details = 50 Am.
Capsule 2 mm long, asymmetric,strumose,furrowed when dry, brownish to blackish brown.

Operculumobliquelyrostrate.Calyptraciliate at
base.
Distribution (Fig. 46). In montane and subalpine forests between 2300 and 3300 m, most
frequentlybetween2700 and 3200 m, rarelyalso
in paramosup to 3700 m elevation throughthe
Andes in Guatemala, Costa Rica, Panama, Co-
lombia, Venezuela, Ecuador, Peru, Bolivia to

northern Argentina.
Specimens examined. GUATEMALA. QUEZALMujulicabetween S. Martin Verde
and Colomba, Standley 85675 (NY). SAN MARCOS:
BarrancoEminenciabetweenSan Marcosand San Rafael, Standley86229, 86300, 86391 (NY).
PANAMA. CmHIIQ: Bois de Boquete,FrereHelion
s.n. (S); Finca Lerida,Salazar et al. 790c (MO, NY);
Volcin de Chiriqui,Davidse& D'Arcy10194B (MO).
TENANGO: Above
74
Flora Neotropica
9?0
t:
80
70
60
o....' o.
0IO
-I~~~~~~~~~~~~~~~~~0
OL?a;'~~~~~~~~
{~20
.............
%
0
0
-"
-
200
400
600
800
:..
-.
100km
100 200 300 400 500 600 miles
FIG .
Preparedby HendrikR.Rypkema 46.?
FIG. 46.
~ fCmyop
~ Ditiuin
~ 'oclr
~ '
.........
Distribution of Campylopus concolor.
COLOMBIA. BOYACA: Chiquinquira,v. d. Ham- Hammen et al. 2267 (U); entre Arcabucoy Villa de
men 2808, 2814 (NY); Alto de Onzaga,v. d. Hammen Leiva, v. d. Hammen et al. 2651, 2804 (U); Carretera
2820 (NY, U); Camino de Soata al Alto de Onzaga,v. Arcabuco-Tunja,v. d. Hammen et al. 2807 (U); Card. Hammen &Jaramillo1667 (U); Paramode la Rusia reteraVilla de Leiva-Chiquinquira,v. d. Hammen et
San CayeNW-N de Duitama,Cleef7092 (U); PefiaBlanca,Cleef al. 2871 (NY), 2814 (U). CUNDINAMARCA:
7343 (U); Arribade la carreteraTunja-Arcabuco,v. d. tano, Hda. Portugal,Cleef6038, 6073, 6044, 6578 (U);
75
CarreteraMosquera-LaMesa, v. d. Hammen et al.

ed. 2,1: 320. 1904.DicranumorthopelmaC. Miiller,
2146 (U); Torca, Cleef 7 (U); Sasaima,Finca S. Jose,
Hedwigia39: 257. 1900. Type. Brazil.Sta. Catarina:
SerraGeral, Ule 1128 (holotype,destroyedat B; lecCleef4963 (U); Sabanade Bogotav. d. Hammen et al.
1999 (U); Carreteraentre Bogota y Puente Comun, v.
totypus nov., H-BR).
d. Hammen et al. 1925 (U); Paramo de Coachi near Campylopusouro-pretensisParis in Brotherus,Nat.
Pflanzenfam.1(3): 334. 1901, nom. nud. cf. Index
Bogota, Killip & Ariste-Joseph11978a (NY); Bogota
Muscorum(n.v.).
Chiquinquira,Lindigs.n. (NY); Paramode CruzVerde, carreteraBogota-Coachi,Cleef 3059 (U). HUILA: Campylopuspleurocarpus
(C.Miller) Paris,Ind. bryol.
Rio Paez Valley, Pittier 1271 (NY). SANTANDER:
CerSuppl.95. 1900. DicranumpleurocarpumC. Muller,
retera Guantiva-Onzaga,v. d. Hammen 1730 (NY),
Bull.Herb.BoissierVI: 35. 1898. Type.Brazil.Serra
do Itatiaia, Ule 1798 (holotype,destroyedat B; lec1727, 1865 (U); al oriente de Santander,Uribe227
(FH).
VENEZUELA. Colonia Tovar, Pittier 439 (MO). CampylopusrectipesC. Muller, Genera muse. frond.
MERIDA: ParqueNacional de la SierraNevada above
270. 1900. nom. nud. cf. Index Muscorum(n.v.).
Tabay, Griffinet al. 1866 (FLAS);SierraNevada de CampylopusstramineolusC. Miiller, Genera muse.
Santo Domingo, Piramo de Los Granates,Lopez-Fifrond. 270. 1900. nom. nud. cf. Index Muscorum
gueiras15426 (FLAS);CampoElias,Roadto LaAzuli(n.v.).
ta and Jaji, Fransen 1170, 1228, 1176, 1178 (GB);
SierraNevada de Merida,between La Aguadaand la
Plants in loose tufts, glossy dark green, to 12
Montaiia,Griffinet al. 1771 (FLAS).TACHIRA:
Junin, cm
high. Stems slender, unbranched, equally foParamode Tami above Paez, Griffinet al. 948 (FLAS).
liate, at tips often homomallous, radiculose.
TRUJILLO:Bocon6, Paramo de Canada, Ruiz-Terdn
Leaves 6-8 mm long, narrow lanceolate, longly
9184 (FLAS).
PERU. AMAZONAS:18 km above Leimebamba,
pointed, serrate at tips. Costa longly excurrent,
Hutchinson& Wright4879 (NY). APURIMAC:
Quebra- ridged and serrate at back, taking 1/3 of the leaf
da Pallca bei Quishuari,Hegewald 9828 (hb. Hegebase, in transverse section with ventral and dorZwischen Contumaza und Cascas,
wald). CAJAMARCA:
Hegewald7357 (hb.Hegewald);Lopezet al. 9060 (NY). sal stereids. Alar cells well developed, hyaline or
Cuzco: Aguascalientesund MachuPicchu,Hegewald reddish. Basal laminal cells rectangular, incras8678, 8762, 8817, 8819, 8859 (hb. Hegewald).
sate and pitted, narrower at margins. Upper lamBOLIVIA. Yamacarhi-Yungas,Jay 1893 (NY).
inal cells short rectangular to rhomboid or oval.
Yungas,Rusby3115 (NY).
Seta 6-9 mm long, brownish, pseudolateral.
2 mm long, brownish, furrowed when
Capsule
16. Campylopus controversus (Hampe) Jaeger,
and
dry
empty, slightly curved. Operculum
Ber. Thitigk. St. Gallischen Naturwiss. Ges.
1877-1878: 385. 1880. Dicranum controver- obliquely rostrate.
Distribution (Fig. 44). On wet rocks, wet banks
sum Hampe, Vidensk. Meddel. Naturhist.
and logs in rainforests and cloudforests in SE
Foren. Kjoebenhavn ser. 3, 4: 42. 1872. Type.
Brazil. Glaziou 5204 (holotype, BM; isotypes, Brazil, from the lowlands to the forestline (2300
m), usually above 1000 m, rarely above the foL, NY).
Figs. 44, 47.
restline up to 2600 m.
Campylopusflaccidus Brotherusin Engler& Prantl,
SerNat. Pflanzenfam.1(3): 334. 1901. DicranumflacSpecimensexamined:BRAZIL.MINASGERAIS:
cidum C. Muller, Hedwigia 39: 251. 1900, nom. ra de Ibitipoca,Pico de Piao, Sucre6902 (RB).Rio DE
Ule178, 1087, 1798, 1897,1921, 2044 (HBG);
invalid. Type. Brazil.Minas Gerais:Serrade Ouro JANEIRO:
Tijuca, Ule 210 (H-BR);Pico de Papageio, Ule 1087
Preto, Ule 1362 (holotype,H-BR).
Campylopushumoricola(C. Miller) Paris,Ind. bryol. (H-BR);Corcovado,Bandeira245, 440 (NY);Itatiaia,
ed. 2, 1:313.1904. DicranumhumoricolaC. Miller, Glaziou5204 (PC);ParqueNacionil de Itatiaia,AgulHedwigia 39: 256. 1900. Type. Brazil. Rio de Ja- has Negras, Vitt21491, 21499 (ALTA);Schiffner319,
neiro:Corcovado, Ule 1650 (holotype,destroyedat 559, 567, 601, 1395 (H-BR);Dusen 486 (H-BR, NY);
Landrum2156 (NY); Frahm, Camp.Bras. Exsicc. 25
B; lectotypusnov., H-BR).
Campylopusmacrogaster(C. Muller)Brotherusin En- (ALTA, B, C, DUIS, FLAS, GOET, GZU, H, HBG,
gler & Prantl, Nat. Pflanzenfam.1(3): 334. 1901. INPA, JE, KR, M, MO, NFLD, NICH, NY, PC, S,
DicranummacrogasterC. Miiller,Hedwigia39: 252. SP, TENN, U); Abrigo Reboucas, Schifer-Verwimp
1900. Type. Brazil. Sta. Catarina:SerraGeral, Ule 7557 (hb. Frahm);Rio Campo Belo, Landrum2190
1123 (holotype, destroyed at B; lectotypus nov., (NY); Teres6polis,ParqueNational Serrados Orgaos,
Frahm 1598 (hb. Frahm).SAo PAULO:
ReservaBiol6H-BR).
Campylopusmosenii Brotherus,Bih. Kongl. Svenska gica de Mogi Guacu, Vital2676 (SP);Ilha do Cardoso,
Vetensk.-Akad.Handl. 21 Afd. 3(3): 6. 1895. Type. Vital 10322 (SP).
Brazil.Sao Paulo:Alto da Serra,Mosen18 (holotype,
This species is closely related to Campylopus
H-BR).
Campylopusorthopelma(C. Miiller)Paris,Ind. bryol. stenopelma from S and SE Africa. The ranges of
Flora Neotropica
76
li
FIG. 47. Campylopuscontroversus(CampylopodesBrasiliaeExsiccatae 25). A. Plant. B. Leaf. C. Leaf-tip.

D. Transversesection of leaf. E. Basallaminal cells. F. Upper laminalcells. G. Capsule.Unlabeledscale bar for
microscopicdetails = 50 mm.
both species are on the same latitude and in the

same geographicalsituationat the East coasts of
the continents. They can thereforeregardedas
vicariant species derived from the same Mesozoic ancestorand could even be regardedas subspecies.
17. CampylopuscryptopodioidesBrotherus,Bih.
Kongl.SvenskaVetensk.Akad. Foerh.26 Afd.
III(7): 9. 1900. Type. Brazil. Mato Grosso:

Lindman 475 (holotype, H-BR; isotype, S).
Figs. 48, 49.
ramuligerBrotherus,Denkschr.Akad.
Campylopus
KI.83:262. 1926.Type.BraWiss.Wienmath.-nat.
zil. SaoPaulo:Apiahy,Schiffner110 (holotype,not
presentin H-BR;isotypes,BM,PC).
Plants in green, loose, soft tufts to 3 cm high.
Stems erect,denselyfoliate,sometimesbranched,
77
? 0
C) o0 0
.o
FIG. 48. Campylopuscryptopodioides

(Vital 9491, SP). A. Plant. B. Leaf.C. Leaf-tip.D. Transversesection
of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Mm.
tomentose,with severalbudsof deciduousbroodleaves. Leaves to 3 mm long, lanceolate, contractedto a subtubulose,serratetip, erect patent
when wet, rugoseand appressedwhen dry. Costa
filling 1/3 of the leaf width, vanishing in the leaf
tip, smooth at back. Alar cells thin-walled, hyaline, isodiametric.Basallaminalcells shortrectangular,incrassate, 10-13 x 16-29 gm, at margins shorter and narrower.Upper laminal cells
small, quadrate,incrassate, 13-16 x 13-22 tm.
Distribution(Fig. 49). On tree trunksand rot-
ten wood, found only in Southern Brazil and

adjacent Argentina between sea level and 1500
m.
Specimens examined. BRAZIL. MINASGERIS: Ser-
ra de Mantiqueira,Camanducaia,Schdfer-Verwimp
6708 (hb. Frahm). PARANA:
Mun. de Quitandinha,
25?53'S, 49?30'W, Vital 9491 (SP). Rio GRANDEDO
SUL:MorroItacolumi,Sehnem 4767 (LBLC);Rio do

Simos, Sehnem s.n. (hb. Frahm). Rio DEJANEIRO:Ser-
ra dos Orgaos,v. Liitzelburg6503, 6718 (B, JE). SAO

PAULO:
Serrado MarnearUbatuba,Schdfer-Verwimp
7866, 8278 (hb. Frahm);ReservaBiologicado Alto da
Serrade Paranapiacaba,Vital 10407 (SP).
Flora Neotropica
78
70
80
.0
50
60
30
~"
-:---~-1- .
.... <^. ....ft<
...........20
B:1~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
O~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
..
I ,?.~4
~'
J
.
202
~~~~~~3_O~~~~~~~~~~~~~~~~~3
0
200
4.
Dstibuio
FIG
'
I
800 1000km
600
400
100 20030
-Prepared
400 500 600miles
R.F!ypkom'a"-
aplpscytpdode
\,
~'-teyT----__
by
Hendr'i
of
'o "
"
'
^^<^^"\
0
ft'
*/! \
Of
uess()
'y
,/
"':~",?'
30T~-
i^^
ofCampylopus
FIG.49. Distribution
(O).
(0)andC.cubensis
cryptopodioides
79
Gr
2mm
C UUuoo ~
11"o~n~n~
Ii
I O'
u001
A
4cm
FIG. 50. Campylopuscubensis(Samek s.n., PRC). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of
50 Am.
ARGENTINA. IguaSu,v. Hiibschmanns.n. (hb. v.

Hiibschmann).
This species is easily recognized by the small
lateral branches consisting of brood-leaves, the
short, relatively blunt, crisped leaves with quadrate laminal cells, which reach down along the
margins to nearly the leaf-base.
18. Campylopus cubensis Sullivant, Proc. Amer.

Acad. Arts Sci. 5: 278. 1861. Type. Cuba,
Wright 39 (holotype, NY).
Figs. 49, 50.
Campylopusharrisii(C. Miiller)Paris,Ind.bryol.Suppl.
92. 1900. DicranumharrisiiC. Miiller, Bull. Herb.
Boissier 5: 553. 1897. Type. Jamaica. Newhaven
Pass, Harris 11008 (holotype,destroyed;lectotypus
nov., NY; isolectotype,S).
Flora Neotropica
80
Plants tall, to 7 cm high, green to yellowish
green. Stems reddish tomentose, sometimes
branched. Leaves erect patent, flexuose, rarely
distant or even recurved,about 7 mm long, lanceolate, ending in a very long-pointed, slender,
coarselyserrateapex, also leaf marginserratein
the upperthird of the leaf. Costa taking '/3of the
leaf base, in transversesection with ventral and
dorsal stereids,ribbedat back in the upperpart.
Alar cells forming largereddish brown auricles.
Basal laminal cells rectangular,incrassate and
pitted, 6-10 x 26-48 um, shorterand narrower
at margins.Upper laminal cells incrassate,4-6
x 10-29 4um,short rectangularto oblique, in
rows.
Vegetative propagationrarely by microphyllous branches(found only in Ekman 3798).
Seta about 1 cm long, yellowish, sinuose or
cygneous, aggregated.Capsule to 2 mm long,
asymmetric, furrowed,yellowish with red, not
dehiscent annulus. Operculumlongly rostrate,
oblique, reddish, darkerthan the capsule. Calyptrafringedat base.
Distribution (Fig. 49). On soil, rotten wood
and base of trees in forests,in Hispaniola,Cuba,
PuertoRico, Jamaicaand Trinidad,also in Central and northernSouth America in Costa Rica,
Panama,Venezuelaand the Guianas.Usually in
rain forests, in Cuba frequentlyin pine forests,
in elevations between 500 and 2000 m.
12965,15734, 15740 (NY);betweenConstanzaand

Jayaco,Norris& Jones11060 (NY);Mt. Diablotin,
Fishlock41a (NY).
PUERTORICO.Maricao500-900 m, Britton2640,
Steere5487,Buck3855,Reese14609(NY);Sierrade
Cayay,Steere6711 (NY).
TRINIDAD.Withoutlocality,ex hb. Mitten(NY).
VENEZUELA.
BOLivAR:
CerroJana,Steyermark
109788(NY).IslaMargarita,
Sugden1200(NY).
GUYANA. Mt. Membaru, Maas & Westra4269
(NY);Mt. Latipu,5?37'N,60?38'W,Maas& Westra

4219 (U);UpperMazaruni:
5?57'N,60?45'W,Gradstein5742 (U).
SURINAM.Lelygebergte,
Florschiitz4844 (NY);
Florschiitz
4664 (NY).
Brownsberg,
FRENCHGUIANA.Cretedes MontsBakra,Cremers4213 (hb.Onraedt).
Campylopuscubensis resembles C. arctocarpus and differsmainly by the more robustplants
with spreadingleaves, sharplytoothedleafapices
and the more elongateupperlaminalcells (which
may be due to the longer leaves). It therefore
wouldbe takenfora subspeciesofC. arctocarpus,
and has probablyevolved from the widespread
neotropical C. arctocarpusvar. arctocarpusby
isolation on the CaribbeanIslands. The records
of C. cubensison continentalCentraland South
Americaaroundthe CaribbeanSeamaybe caused
by secondaryspreading.Similarly, C. arctocarpus occurs also on the CaribbeanIslands,but is
considerablyrarerthan on the continentand also
rarerthan C. cubensis.
Campylopuscubensiswas placed into synonymy of Campylopusfilifolius
(Bryohumbertiafilifolia) by Mitten (1869). The type of C. cubensis
is, however, identical with what later had been
describedas C. harrisii.Therefore,most records
of this species are named C. harrisiiin herbaria
and publications.Williams (1915) treated both
C. cubensisand C. harrisii in the North American Flora, differentiatingthem by the costa either excurrentor scarcelyexcurrentand the leaf
blade either 3-4 or 2 cells wide, but this is not
sufficientto separatethem.
Specimens examined. PANAMA. PANAMA:Vic.

CerroJefe, Lewis& Dressier7604 (NY, U); Cleefet al.
Cerro
10312 (U); Witherspoon8599 (MO);VERAGUAS:
Tute above ExcuelaAgricola"Alto de Piedra,"Crosby
10813 (MO); mouth of Rio Concepci6n,Lewis et al.
2797 (NY).
CUBA. ORIENTE:
SierraCristal,Samek s.n. (PRC);
SouthernBaracoaregion,Le6n 12011 (NY); regionde
Moa,Borhidiet al. 6083, Jervis1682, 1693, 1685 (NY);
crestof SierraNipe, Morton&Acufna3272, 3914, 3919
(NY); Charrasco,Le6n 12013 (NY); Pinal, Wright87
(NY); near Saguade Tamano, Wright88 (NY); Taco
Bay, Samek et al. 21 (PRC),Ekman 3798 (NY); Cupeyal, Samek et al. 40a (MO); La Brena-Moa,Leon
et al. 22701 (PRC).
JAMAICA. ST. ANDREW
Sir John's Peak, 19. Campylopuscuspidatus
PARISH:
(Hornschuch)MitBritton1166, Crosby3033 (NY);Mainridgegap,MaxLinn.
Soc.
Bot.
12:
90. 1869.
J.
ten,
on 10212 (NY); Cinchona, Britton 24, 53, Harris
11192
New
Haven
Nichols
11025a,
127,
(NY);
Gap,
Harris26 (NY);Blue Mountains,Morce'sGap,Harris
Key to Varieties of
10043, 10040, Nichols 55, Patterson20, 27, Brittton
1218, 1048 (NY);John CrowPeak, Underwood2366a
Campylopus cuspidatus
(NY);nearVinegarHill, Portland,Maxon &Killip834
1 Leaves ending in a hyaline tip. ...............
(NY); HardwarGap, Buck 5736 (NY).
DOMINICAN REPUBLIC. LA VEGA:Vic. Piedra
Blanca, Allard 17120 (NY); ESE of Bonao, Zanoni
..........................
1 Leaftipscucullate......
19a. var.cuspidatus.
19b. var.dicnemioides.
81
(I
\8mm
|~A
Efi
FIG. 51. Campylopuscuspidatusvar. cuspidatus(Griffinet al. 322, FLAS). A. Plant. B. Leaf. C. Leaf-tip.
D. Transversesection of leaf. E. Basallaminalcells. F. Upper laminalcells. Unlabeledscale bar for microscopic
details = 50 um.
19a. Campylopus cuspidatus (Hornschuch) Mitten var. cuspidatus

Figs. 51, 52.
Campylopuscarassensis(Brotherus)Paris, Ind. bryol.

242. 1894. Thysanomitrioncarassensis Brotherus,
Acta Soc. Sci. Fenn. 19(5): 9. 1891. Type. Brazil.
MinasGerais:Carama,Wainios.n. (holotype,H-BR;
DicranumcuspidatumHornschuch,Fl. bras. 13. 1840.
isotypes, FH, JE, S).
Type. Brazil.Sincoraet Rio das Contas,Martiuss.n. Campylopushoffmannii(C. Miller) Renauld & Cardot, Bull. Soc. Roy. Bot. Belgique31(1): 147. 1893.
(holotype,destroyedat B; lectotypusnov., BM; isoDicranumhoffmanniiC. Miller, Linnaea 38: 592.
lectotypesJE, NY).
1872. Type. Costa Rica. Forets du Barba,Bondez
CampylopusbolivarensisBartram,Fieldiana Bot. 28:
4. 1951. Type. Venezuela. Bolivar: Mt. Roraima,
9909 (holotype, destroyed at B; lectotypus nov.,
Steyermark58895 (lectotypusnov., FH; isotype, F).
H-BR).
Flora Neotropica
82
90
80
70
60
50
.0
30
-
-.......
IN
~I.
> ~ i~ ~'~ ) /
~,/. ..."Y--_.
i
100 200 300 400 500 600
mles
Preparedby HendrikR.Rypkema
FIG. 52.
A---t
--
-.
V'
~ I~ )
~--~
20
20
--.
Distribution of Campylopus cuspidatus var. cuspidatus (0) and C. cuspidatus var. dicnemioides (0).
CampylopusnematophyllusRenauld & Cardot, Bull.

Soc. Roy. Bot. Belgique32(1): 148. 1893, nom. inval. in synon. cf. Index Muscorumas syn. of C. hoffmannii (n.v.).
Plants to 15 cm high, erect, yellowish green,
glossy, in herbarium specimens golden, brownish
below. Stems single, erect, erect patent foliate,
the stem tips acuminate. Leaves 5-7 mm long,
lanceolate, the upper leaf margins involute, entire. Costa narrow, /3-5 of the leaf width, with
short side-nerves, ending in a hyaline hairtip, in
transverse section with ventral hyalocysts and
dorsal stereids, weakly ridged at back. Alar cells
reddish brown, inflated, auriculate, with collenchymatous thickenings. Laminal cells strongly
incrassate and pitted, elongate to vermicular, 310 x 38-70 gm, shorter in the upper part of the
leaf, 3-10 x 2248 Mm.
Sporophyte not known (?).
Distribution (Fig. 52). On wet rocks, rotten
wood and base of trees in rainforests, disjunct in
SE Brazil, Mexico, Guatemala, Costa Rica and
Venezuela.
CerroTres
Specimensexamined.MEXICO.CHIAPAS:
Picos, Breedlove30109A (hb. Frahm).
GUATEMALA.Amatitlan,Volcan Pacaya,Kellermann s.n. (DUKE).
COSTARICA.Cerrode las Vueltas,Standley43685
(hb. Frahm).
Paramo de Tama above
VENEZUELA. TACHIRA:
the village of Paez, Griffinet al. 322 (FLAS).
83
C,
'
'
Ai*
*-^
Po
Ns
FIG. 53. Campylopuscuspidatusvar. dicnemioides(Cleef9934, U). A. Plant. B. Leaf.C. Leaf-tip.D. Transverse section of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails
==50
50 CLm.
Im.
BRAZIL. MINASGERAIS:Serra do Ouro Preto, Ule
1364 (H-BR).
Campylopus cuspidatus is macroscopically

similar to C. subcuspidatusand can easily be
confusedwith this species. It is, however, differentiated microscopicallyby elongateupperlaminal cells, narrowcosta (1/3-/5 of the leaf width)
and small ventral hyalocysts in transversesection of the costa. Campylopussubcuspidatushas
in contrastoval upperlaminalcells, a broadcosta
and large, lax ventral hyalocysts in transverse
section of the costa, which are conspicuous in

ventral view of the leaf without sectioning.
Fordifferentiationfrom C. bryotropiisee notes
under the latter species.
19b. Campylopuscuspidatus(Hornschuch)Mitten var. dicnemioides(C. Miller) J.-P. Frahm,
Rev. Bryol. Lichenol. 45(2): 143. 1979.
Figs. 52, 53.
Campylopusdicnemioides(C. Muller)Paris,Ind.bryol.
Suppl.91. 1900. DicranumdicnemioidesC. Miller,
84
Flora Neotropica
Bull. Herb. Boiss. 6: 36. 1898. Type. Brazil. Serra

do Itatiaia,AgulhasNegras, Ule 1788 (holotype,destroyedat B; lectotypusnov., H-BR).
CampylopuscucullatifoliusHerzog,Bibl. Bot. 87: 22.
1916. Type. Bolivia.Tablas3400 m, Herzog2826/a
(holotype,JE).
section of the costa, indicate that both populations are derived from C. cuspidatus and must
be joined in one taxon.
Varieties with cucullate instead of hyaline leaf
tips are present in several other species as Campylopus such as C. atrovirens De Not., C. acuminatus Mitt., C. flaccidus Ren. & Card. and
others. The cucullate varieties occur always in
wetter habitats and mostly in the subalpine to
alpine belt.
Like var. cuspidatusin deep, brownish tufts.

Stems with lightertips. Anatomicallyas var. cuspidatuswith conspicuousnarrowcosta with sidenerves, incrassate, pitted laminal cells and especially the collenchymatouscells of the alars.
Differs only in the cucullate instead of hyaline 20.
Campylopus cygneus (Hedwig) Bridel, Mant.
leaf tips.
musc. 72. 1819. Dicranum cygneum Hedwig,
musc. frond. 148. 1801.
"India
Sp.
occ.,"
Type.
Distribution (Fig. 52). In swamps above the
Swartzs.n. (holotype, G; isotypes, H-SOL, NY,
forest line in SE Brazil and in wet paramos of
S).
Figs. 54, 55, 56.
the Andesin Venezuela,Colombia,Ecuador,Peru
and Bolivia.
Campylopuscacuminis (C. Miiller)Paris, Ind. bryol.
PiraSpecimensexamined.COLOMBIA.BOYACA:
mo de Chita, cabecerasdel Rio Casanare,Cleef 9913,
9934 (U); SierraNevada de Cocuy, ParamoConcavo,
Cleef10.016 (U); Paramode la Rusia, Cleef7355 (U);
N-NW de Duitama, Cleef 7271 (U); Pefia de Arical,
N de Vado Hondo, Cleef 9475 (U); Vado Hondo, SiVolcan Purace, Cleef
beria, Cleef 9267 (U). CAUCA:
2636 (U). CUNDINAMARCA:
Paramo de Cruz Verde,
Cleef & Jaramillo 3091 (U); Paramode Palacio, Lagunas de Buitrago,Cleef 2357 (U); Paramo de Cruz
Verde, LagunaEl Verj6n,Cleef& Duncan 3157, Cleef
3219 (U);ParamoentreCoguay SanCayetano,Laguna
Verde,Cleef6214a, 6498 (U), LagunaSeca,Cleef6147
(U); CarreteraBogota-Coachikm 14, Cleef2939, 3043
(U); Paramo de Sumapaz, Chisaca, Cleef 4951 (U).
HUILA:Parque Nacional Cueva de los Guicharos y
cerro Punta, Cleef 5084 (U). META:Paramo de Sumapaz, Cleef8206 (U).
VENEZUELA. BOLiUAR:
Summit of Apacara-TeJunin,Paramo
pui, Steyermark75877 (NY). TACHIRA:
de Tama above Paez, Griffinet al. 818, 877 (NY).
ECUADOR. NAPO:Road Quito-Baeza, 2 km E of
pass, Frahm 335 (hb. Frahm).
Las Lagunas,Hegewald 6211
PERU. CAJAMARCA:
(hb. Hegewald).
BRAZIL.AMAZONAS:
CerroDuida,3?22'N,65?36'W,
Buck& Brewer15555 (NY). Rio DEJANEIRO:
Serrado
Itatiaia,Frahm, Camp.Bras.Exsicc. 11 (ALTA,B, C,
DUIS, FLAS, GOET,GZU, H, HBG, INPA, JE, KR,
M, MO, NFLD, NICH, NY, PC, S, SP, TENN, U).
The populations in the Andes (C. cucullatifolius) and SE Brazil (C. dicnemioides) differ
somewhat in the length of the laminal cells and

may ultimately representdifferentgeographical
races. However, such charactersas side-nerves
in combination with collenchymatousalar cells
and an incrassate,pitted areolation throughout
the whole lamina, and an identical transverse
Suppl. 90. 1900. Dicranum cacuminis C. Miiller,

Hedwigia37: 227. 1898. Type. Jamaica.Blue Mtns.
Peak, Hansen s.n. (holotype,destroyedat B).
CampylopusekmaniiTheriot,Mem. Soc. CubanaHist.
Nat. 13: 216. 1939. Type. Cuba.SierraMaestra,La
GranPiedra,Ekman 1703 (lectotype,H-BR;isolectotypes, NY, S).
CampylopuselliottiiBartram,Bull.Brit.Mus.Nat. Hist.
Bot. 2: 38. 1935. Type. Dominica. Summit of Mt.
Trois Pitons, Elliott 478a (holotype, FH; isotype,
BM).
CampylopusjamaicensisMitten,J. Linn. Soc. Bot. 12:
82. 1869.Type.Jamaica.Wilson813 (holotype,NY).
Campylopussaxatilis Williams,N. Amer. Fl. 15: 137.
1913. Type. Jamaica.Cinchona,Harris 11143 (holotype, NY; isotypes, DUKE, F).
Plants to 7 cm high, usually smaller, very slender, in loose, light green, glossy tufts. Stems
equally foliate except for comose, perichaetia
bearing plants, sparsely radiculose. Leaves erect
patent, sometimes flexuose, ca. 7 mm long, ending in a very long and fine nearly smooth or only
finely denticulate apex. Costa taking 13 of the leaf
base, excurrent, in ventral view with enlarged
cells, in transverse section with large hyaline ventral cells, a median row ofdeuter cells and groups
of stereids at the dorsal side, smooth at back.
Alar cells very conspicuous, large, inflated, forming round auricles, protruding into the costa, reddish, rarely hyaline. Basal laminal cells incrassate, nearly subquadrate or very short rectangular
(2:1), 13-16 x 38-51 um, at margins in 1-2 rows
elongate and hyaline. Upper laminal cells
subquadrate to shortly rectangular, incrassate, 810 x 16-22 um. Vegetative propagation by deciduous stem tips.
85
.af
$/7z...
yf
]L^^ w
4bf
CY.
FG54Cm
FIG. 54.
cygneyrJus isAll
Campylopus cygneus (isotype, S).
Seta 8-9 mm long, brownish, sinuose or cygneous. Capsule asymmetric, short oblong, furrowed, strumose. Operculumobliquely rostrate.
Calyptraciliate at base, rarelyentire.
Distribution (Fig. 56). On soil, rotten wood
and rocks in montane rainforestsin elevations
between 1500 and 2500 m, confined to the Caribbeanislands Cuba,Jamaica,Hispaniola(Dominican Republic, Haiti), Puerto Rico and the
LesserAntilles,also describedas C.setaceusCard.
from the Azores.
examined.
CUBA.ORIENTE:
SierraMaesSpecimens
tra,Maids.n. (PRC);summitof GrandPiedranear
Santiago,Imshaug24984 (F). SANTAGODECUBA:La
GranPiedra,Buck7627 (NY);SierraMaestra,Pico
Turquiiio, Vilimkova1, 3 (PRC); Mala s.n. (PRC),

Bucher54b (NY); W of SierraMaestra,Morton 9414
(NY).
JAMAICA. ST. ANrDREW
PARISH:Morce's Gap,
18005'N, 76?39'W, Maxon & Killip 1280 (NY); Crosby
3560 (DUKE); Parish of St. Thomas, PortlandGap,
Crosby 3515, 3506 (NY); Cinchona, Maxon & Killip

1300 (NY), Harris s.n. (NY); Sir John's summit, Britton 1145 (NY); St. Catharine's Peak, Nichols 75 (NY);
Blue Mtn. Peak, Petterson 51, Maxon 10003, Jaderholm 47 (NY); Hegewald 8116 (hb. Frahm); Griffin
25452 (FLAS);WhitfieldHall CoffeePlantation,Harris 10053a (NY).
HAITI. Massif de la Hotte, More Formon, Judd
4099 (NY).
DOMINICAN REPUBLIC. LA VEGA: 8 km S of
Constanza, Norris et al. 7515 (NY); vicinity of La Lagunita, Norris et al. 5525 (NY); between Constanza
86
Flora Neotropica
FIG. 55. Campylopuscygneus(Griffin25452, FLAS). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 jim.
and La Nuez, Jones &Norris1294B (NY); 13 km from 14420 (NY); Guayanaroad near Cayey, Steere 5087
Valle Nuevo, road to S. Jose de Ocoa, Norris et al. (NY); between Salinas and Cayey, Steere 5298 (NY);
7245 (NY); between Constanzaand Jayaco, Jones & Sierrade Cayey,Steere6710 (NY); Sierrade Luquillo,
Norris 10470 (NY); 40 km S of Constanza, 18?45'N, Mt. Britton Peak, Steere 7094 (MICH);Monte Gui70?37'W,Reese 15777 (NY); 22 km S of Constanza, larte, Reese 14635 (NY); trail from Mt. Brittonto El
18?49'N,70?37'W,Buck 5449 (NY); INDEPENDENCIA: Yunque, Buck 4036 (NY); trail from Hwy 186 to top
Crest of SierraNeiba, Norriset al. 6585, 6260 (NY); of El Toro, Reese 14420 (NY).
LA ESRELLETA:Sierra de Neiba, Buck 4585, Smith
LESSER ANTILLES. SABA:Mt. Scenery, v. SlaSan Jose de Ocoa, 18?36'N, geren 142M (U).
10249c (NY); PERAVIA:
70?35'W,Zanoni 8743 (NY); 45 km S of Constanza,
Reese 15746, 15720 (NY).
Campylopus cygneus is quite variable and has
PUERTO RICO.CaribbeanNat. Forest,ElYunque,
Steeres.n. (NY), Buck4036, 4048 (NY);ElToro,Reese therefore been described under several names.
87
80
90
60
70
50
40
30
--
---
-- ----o
]. y--<-
Prepared
by Hendrik R. Rypkema
FIG. 56. Distributionof Campylopuscygneus() and C. densicoma(0).
The variationconcernsthe lengthof the leaf tips, 21. Campylopus densicoma (C. Muller) Paris,
the foliation (appressed or comose in female
Ind. bryol. Suppl. 91. 1900. Dicranum densicoma C. Miller, Nuovo Giorn. Bot. Ital. n.
plants)and the lower laminal cells (thick walled
and subquadrateto thin walled and hyaline in
ser. 4: 33. 1897. Type. Bolivia. Cochabamba,
Germain 1120 (holotype, destroyedat B; lecperichaetialleaves). It is, however, easily recognized by its large auricles protrudinginto the
totypus nov., NY; isolectotype, H-BR).
costa, the elongate,smooth leaf tip, the quadrate
Figs. 56, 57.
upper laminal cells and the large ventral hyalocysts in transversesection of the costa, which are Campylopus
cavernosus
J.-P.Frahm,Nova Hedwigia
29:245. 1978.Type.Brazil.Rio de Janeiro:Itatiaia
conspicuousalso in ventral view on the costa.
Flora Neotropica
88
Nat. Park,AgulhasNegras,2760 m, Vital4921 (holotype, SP; isotypes, B, U).
Campylopusfilicuspes Brotherusin Herzog, Biblioth.
Bot. 87: 22. 1916. Type. Bolivia. Comarapa,2600
m, Herzog4192(holotype,JE;isotypes,B, FH, H-BR,
HEREDIA:Las Vueltasarea 18 km N of SanJose, Cros-
by 6455 (MO).
PANAMA. Cerro Fortuna, 8?45'N, 82?15'W,Salazar et al. 637 (NY, PMA).
Cabeceras de la
COLOMBIA. CUNDINAMARCA:
S).
QuebradaChuza,Cleef9658 (U);Paramode Sumapaz,
CampylopusgertrudisHerzog, Biblioth. Bot. 87: 21. vanderHammen 2864, Cleef8336 (U); Boyaca,Pra1916. Type. Bolivia. Sillar, 2000 m, Herzog 2731 mo de Pisva, Cleef9896 (U). Magdalena,ParqueNacional de la SierraNevada de SantaMarta,Rio Burita(holotype,JE;isotypes, B, FH, H-BR, S).
Campylopusspurio-concolor(C. Miiller) Paris, Ind. ca, Rangel et al. 341 (FLAS).
VENEZUELA. ARAGUA:
Manacay,ParqueNaciobryol.Suppl.97. 1900. Dicranumspurio-concolorC.
Miller, Nuov. Giom. Bot. Ital., n. ser. 4: 34. 1897. nal HenryPittier,Onraedt4649 (hb.Frahm).TAcHIRA:
Type. Bolivia. Yungas, Rusby 3115 (holotype, de- Distr. Junin, Paramode Tama above Paez, Griffinet
al. 252 (FLAS);Piramo El Rosil betweenLa Britaand
stroyedat B; lectotypusnov., H-BR).
San Jose, Griffinet al. 741 (FLAS).
ECUADOR.NAPo:CerroSumaco,77?39'W,0?36'S,
Lojtnant& Molau 13059 (AAU, NY).
PERU. ANCASH:Cordillera Blanca, road CatacCampylopusdensicoma
Chavin 4200 m, Frahm, Camp. Peruv.Exsicc. 7 (B,
1 Leaves erect, curledor crispedwhen dry.......
C, EGR,F, FLAS,G, GRO,GZU, H, HBG, KR, KRA,
..........................
21a. var. densicoma. M, MEXU, NAM, NFLD, NY, PC, S, U, UPS, USM).
1 Leaves distinctly hamate and homomallous,
Dep. Cuzco,AguascalientesnearMachuPicchu,Hege21b. var. yungarum. wald8711 (hb. Hegewald);above Pillahuata,13?11'S,
straightwhen dry. .........
71?22'W,Fitzpatrick& Willards.n. (NY);PUNO:Prov.
St. Domingo area, Carroll121 (NY).
21a. Campylopus densicoma (C. Muller) Paris Sandia,
BRAZIL. Rio DEJANEIRO:
ParqueNacional Serra
var. densicoma
do Itatiaia, Agulhas Negras, 2600 m, Camp. Bras.
Exsicc. 12 (Serrado Itatiaia),Naveau 263, 295 (PC);
Plants slender to robust, to 8 cm high, green
AbrigoReboucas,Griffin& Vital26 (FLAS).Rio Mato brownishgreen, equally foliate or slightlyco- romba,Massart3276 (PC).
San Lapata, Cardenas
mose at stem tips. Stems erect or prostratein
specimens from inundated habitats. Leaves to 3236 (NY). LA PAZ:Hot Springsbetween Mina Huiand Rio Glorieta, 67?17'W, 16?57'S,Lewis
10 mm long, narrowlylanceolate, ending in a chincani
87507 (NY).
long, slenderpoint. Costalonglyexcurrent,finely
ARGENTINA. TUCUMAN:
Est. Las Paras, Venturi
serrate at tips, filling l/2--/3of leaf width, in trans- 1088 (FH).
verse section with large ventral hyalocysts and

This species is very variable. It has been dedorsalgroupsofstereids.Alarcells differentiated,
from Bolivia by C. Miiller under two
scribed
reddish.Basallaminalcells rectangularwith narrow cell walls, 10-13 x 48-64
im, narrower at
margins.Upperlaminalcells incrassate,irregular
oblique to oval, 6-10 x 13-19 ,m, ca. 4-6:1.
Sporophytespseudolateral,1-3 per perichaetium. Seta relativelylong, up to 15 mm, straight,
slightlysinuosebut not curved,lightbrown.Capsule 2 mm long, erect, symmetric, light brown
to dark brown in age, furrowed when empty.
Operculumlongly rostrate,nearlyas long as the
capsule. Calyptranot fringedat base.
Distribution (Fig. 56). On trunks of trees, nodes
different names and by Herzog under three different names. The C. Miiller names were published in the same publication and have equal
priority. The epithet densicoma has been chosen
as the legitimate name (Frahm, 1984c). As revealed by identifications of collections made
through the recent years, it is widespread through
the Andes, north to Costa Rica.
Specimens from wet rocks, dripping cliffs or
river banks are more slender. Such plants resemble the types of C. cavernosus, C. gertrudis and
C. yungarum. In contrast, specimens grown as
epiphytes, resembling the types of c. filicuspes,
C. densicoma and C. spurio-concolor, are more
robust. Both, however, cannot be differentiated
anatomically.
of bamboo, on wet rocks, often periodicallyinundated or beside waterfalls, in montane rain

forests from 1000 m to the forest line, also in
subalpine shrubs up to 4200 m in Costa Rica,
Panama, Venezuela, Colombia, Ecuador,Peru,
Bolivia, northernArgentinaand disjunct in SE
Brazil.
21b. Campylopus densicoma (C. Miiller) Paris
TaSpecimensexamined.COSTARICA.CARTAGO:
panti Hydroelectric Reserve, Croat 36097A (MO).
var. yungarum (Herzog) J.-P. Frahm, Nova

Hedwigia 39: 156. 1984.
89
6.mm
FIG. 57. Campylopusdensicomavar. densicoma(Cleef9659, U). A. Plant.B. Leaf.C. Leaf-tip.D. Transverse

50 Am.
Campylopus
yungarumHerzog,Beih.Bot.Centralbl.
2200 m,
26(2):51. 1910.Type.Bolivia.Incacorral,
Herzog1908(holotype,JE;isotype,B).
This variety differs only by the strongly hamate leaves. Such hamate forms are found in a
number of species in the genus Campylopus,
which may represent a genotypical difference,
althoughthere is no differencein the ecology or
distribution.
Distribution (Fig. 56). In the same habitats
scatteredthroughthe rangeof var. densicomain
Panama, Colombia and Bolivia.
Specimensexamined.PANAMA. W ridge of Cerro

Tacaruma,Mori & Gentry44296 (MO).
COLOMBIA. ARAUCA:
Sierra Nevada del Cocuy,
Cleef8897 (U). CUNDINAMACA: Paramo de Sumapaz,
Cleef 8479 (U); Piramo de Cruz Verde, Cleef 3048

(U); CarreteraBogota-Fusagasuga,van der Hammen
& Jaramillo 2847 (U); Paramos entre Cogua y San
Cayetano,Florschiitz3602 (U); Subachoque,Paramo
El Tablazo,Frahm 885239 (hb. Frahm).
BOLIVIA.COCBABnA: Prov. Chapar6,E face of
CerroChuaLaguna,17?13'S,65?53'W,Lewis79-2212A
(F); W face of CerroChua LagunaN of Corani,Lewis
79-2194A (F);Rio KhuriN of Corani,Lewis 79-2350,
79-2379 (F).
90
Flora Neotropica
8rm
3i1
Ii
D
FIG. 58. Campylopus dichrostis (Frahm 1594, hb. Frahm). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
50 tAm.
22. Campylopusdichrostis (C. Miiller) Paris in

Brotherusin Engler& Prantl, Nat. Pflanzenfam. 1(3): 333. 1901. DicranumdichrostisC.
Miiller, Hedwigia39: 265. 1900. Type. Brazil.
Minas Gerais:OuroPreto, Ule 1359 (holotype,
destroyed at B; lectotype, H-BR).
Figs. 58, 59.
Plants in compact tufts, to 3 cm high, reddish
tomentose below, darkgreenabove. Stems erect,
loosely foliate with concave leaves, crispedwhen

dry, at stem tips appressed foliate. Leaves 4-5
mm long, lanceolate,lamina reachingto the leaf
tip. Costa filling 1/3of leaf width, lamellose and
serrateat back, ending in the leaf apex, in transverse section with ventral and dorsal stereid
bands. Alar cells reddish brown, thickwalled,
hexagonal.Basallaminalcells incrassate,rectangular,6-13 x 13-67 ,m, at marginsseveralrows
of shortrectangularor almost quadratecells. Up-
91
80
60500
70
-,~
________
___/_
.,
/1
" I0,,.
o~'"~
"
200
'^7
0
^
^"^^
/
o-^
'''.-^C-:.'
j , "^*y^
.FIG.
59.
600
800 1000km
C^-^
Distribution
400
100 200 300 400 500 600 miles
of
Campylopus
dichrostis
(0)
and
C.
edithae
~-
10
(0).
per laminal cells quadrate or somewhat oblique,

incrassate, 8-16 x 10-16 Am.
Distribution (Fig. 59). On soil and rock crevices in light, dry forests, found only in Brazil.
and therefore it cannot be decided whether this

should be interpreted as an extreme modification
or as a genotype, e.g., an ecotype (from heavy
metal-rich soil).
Specimensexamined.BRAZIL.BAHIA:
Chapadade
Diamantina Nat. Park, Schafer-Verwimp8708 (hb.
12 km NE of Jatai, Vital 6359 (SP).
Frahm).GOLAS:
MINASGERAS:Along road MG 55 betweenMorrodo
Pilar and Sao Sebastiao, Frahm 1594 (hb. Frahm);
Serrado Sao Tome das LetrasnearCaxambu,SchiferVerwimp6807 (hb. Frahm).
23. Campylopus edithae Brotherus, Bot. Jahrb.

49: 175. 1912. Campylopus subjugorum
Brotherus var. edithae (Brotherus) Theriot,
Rev. Bryol. Lichenol 11: 42. 1939. Type. Bolivia. Cordillera Real, Knoche 43 (holotype,
H-BR; isotype, B).
Figs. 59, 60.
This species much resembles Campylopus savannarum, from which it is differentiated by the
lamina reaching to the leaf tip, the non-excurrent
costa and the shorter and broader leaves. It is
known only from a few herbarium collections
Campylopusandicola Th6riot, Rev. Bryol. Lichenol.

11: 43. 1939. Type. Bolivia. Chojnacota, Herzog
2976a (holotype,n.v.; isotype, JE).
CampylopusharpophyllusHerzog, Biblioth. Bot. 87:
22. 1916. Type. Bolivia. Cerrosde Malaga,Herzog
4796 (holotype,JE; isotypes, B, S).
Flora Neotropica
92
!\
C
3.5mm
FIG. 60. Campylopusedithae(Griffinet al. 1064, FLAS).A. Plant.B. Leaf.C. Leaf-tip.D. Transversesection
of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Mm.
Theriot,Rev. Bryol.
Campylopus
thysanomitrioides
Lichenol.11: 51. 1939. Type. Bolivia,Cerrosde
Malaga,Herzog4400 (holotype,JE;isotype,B).
Plants forming dense mats or cushions,
brownish or blackish below, with light green or
yellowish green tips. Stems to 5 cm or more,
tomentose below, equally foliate. Leaves concave, conspicuouslycurved inwards(cf. the synonym "harpophyllus"),appressedat stems tips,
about 5 mm long, narrowlylanceolate.Costafilling half of the leaf width, excurrentin a short
hyaline point, in transverse section with thick
walled ventral hyalocysts and dorsal groups of

stereids,smooth at back.Alarcells differentiated,
hyaline. Basal laminal cells thin walled, short
rectangularbeside the costa, 6-13 x 48-64 zm,
becominglongerand narrowerat margins.Upper
laminal cells oval to elongateoval, 3-6 x 10-26
,tm, ca. 4-8:1.
Distribution (Fig. 59). On gravelly or peaty
soil, in dense mats around ponds, on wet rocks
in high alpine habitatsas wet puna and paramo
vegetation, on moraine material beside glaciers
93
from 3600 (rarely 3000) to 5200 m through the
Andes in Venezuela, Colombia, Ecuador, Peru
and Bolivia.
Sierra
Specimensexamined.COLOMBIA.BOYACA:
Nevada del Cocuy, Cleef 5609, 5918, 8551 (U); Alto
Valle Lagunillas,Cleef5609, 5773, 5947 (U); Paramo
de Pisva, carreteraSocha-La LagunaColorada,Cleef
4638 (U); Paramode la Rusia, Cleef6851 (U). CUNDINAMARCA:
Paramo de Sumapaz,4.5 km al S de San
Juan, Cleef 8449 (U).
VENEZUELA.MERIDA:
Sierrade Santo Domingo,
Paramode Mucubaji,Griffinet al. 1064 (FLAS);Paramo de Mucuchies,Steyermark& Rabe 97160 (NY);
Distr. Miranda,along spurroad above Pico El Aquila,
Griffinet al. 1174 (FLAS); Distr. Rangel, Sierra de
Santo Domingo, Paramo de Mucubaji, Griffinet al.
1055 (FLAS).
ECUADOR. NAPO:3 km E of Cerro Quilindana,
78?21'W,0047'S,Lojtnant& Molau 11592 (AAU, GB);
aroundLagunaYuragcocha,Lojnant& Molau 11556
(NY).
PERU.ANCASH:
LagunaLlacaNE ofHuaraz,Frahm,
Camp. Exsicc. 51 (ALTA, B, BING, BM, BUF, C,
DUKE, EGR, F, FLAS, G, GRO, GZU, H, HBG,
KRA, MEXU, NAM, NFLD, NICH, NY, PC, PRC,
POZ, RNG, S, U); ParqueNacional Huascaran,Laguna Llanganuco,Frahm, Camp. Peruv. Exsicc. 8
(ALTA,B, BM, BUF, DUKE,EGR,F, FLAS,G, GRO,
GZU, HGB, KR, NAM, NICH, NY, MEXU, NFLD,
PC, POZ, RNG, S, U); E of Cahuish,Hegewald7681,
7685 (hb. Frahm);QuebradaTulparaju2.5 miles E of
BeHuaraz,ArmstrongB 37568 (COLO).AYACUCHO:
tween Tambo and Quinoaalong road from Huantato
S. Francisco, Frahm, Camp. Exsicc. 66 (ALTA, B,
BING, BM, BUF, C, DUKE, EGR,F, FLAS,G, GRO,
GZU, H, HBG, KRA, MEXU, NAM, NFLD, NICH,
NY, PC, PRC, POZ, RNG, S, U). Cuzco: Between
Orcosand Juliaca,Frahm823924 (hb. Frahm).JUNiN:
LagunaChaulacochanear La Oroya, Hegewald5407
(hb. Hegewald);Laguna Huaylacanchaand Laguna
Anascocha near Canchayllo, Hegewald 5407, 5436,
5896 (hb. Hegewald).LA LIBERTAD:
Laguna SausacochanearHuamachuco,Hegewald6019 (hb. Frahm).
BOLIVIA. LA PAZ:11 km S of Quime, 67018'W,
17"01'S,Lewis 87233 (MO); LagunaHuichincani, 9
km NE from Quime, 67017'W,17?54'S,Lewis 87440
(MO, NY); CumbreNE of La Paz, Philippis.n. (KR).
Campylopus edithae is characterized by a short
but distinct hyaline hairpoint formed by the excurrent costa and is distinguished from similar
species by the costa, which is not contracted at
leaf base and by the short rectangular inner basal
laminal cells. The latter character distinguishes
C. edithae also from C. subjugorum, under which
Theriot placed C. edithae as a variety.
24. Campylopus flexuosus (Hedwig) Bridel,
Mant. musc. 4: 71. 1819. Dicranumflexuosum
Hedwig,Spec. musc. frond. 145. 1801. Thysanomitrionflexuosum (Hedwig)Arnott, Mem.

Soc. Linn. Paris5: 262. 1827. Type: "145 DicranumflexuosumDill. t. 47 f33." Timm s.n.
(lectotype,G).
Campylopusflexuosus
1 Basallaminalcellsnot pitted;plantslooselyfoliatein loosetufts. ..........
24a. var.flexuosus.
1 Basallaminalcellspitted;plantsappressed
foliate
in densetufts. ............
24b. var.incacorralis.
24a. Campylopusflexuosus(Hedwig)Bridelvar.
flexuosus
Figs. 9A, 61, 62.
Campylopus
Mitten,J. Linn.Soc. Bot.
heterophyllus
12:77. 1869.Type.Colombia."AndesBogotenses,
in montibusinterBucumaranga
et Pamplona,"
Weir
347 (holotype,NY;isotypes,FH,H-BR,S).
CampylopushondurensisBartram,Field Mus. Hist.
Nat. Bot.4: 351. 1929.Type.Honduras.Standley

56157a(lectotypus
nov.,F).
Campylopusmexicanus Th6riot, Smithsonian Misc.
Collect.78: 7. 1926.Type.Mexico.Campanario,
Arsene7576 (holotype,PC).
roelliiRenauld&Cardot,Bull.Soc.Roy.
Campylopus
Bot.Belgique38: 9. 1900.Type.CostaRica.Juan
Viiias3400ft, Sargs.n. (n.v.).
Campylopus
sargiiRenauld& Cardot,Bull.Soc.Roy.
Bot. Belgique38: 8. 1900.Type.CostaRica.Sarg
s.n. (sotype,H-BR).
CampylopusstraminifoliusBartram,Contr.U.S. Natl.
Mus.26:63.1928.Type.CostaRica.Standley51212
(holotype,FH).
Plants on loose, glossy tufts, (1-)2-3(-5) cm

high, densely reddishtomentose below, equally
foliate, often bearing short, flagellate, microphyllousbranchesin the axils of the upperleaves.
Leaves straightor flexuose when dry, the upper
leaves often longerand slightlysecund, 5-7 mm
long, lanceolate, graduallynarrowed,serratein
the upperpart.Costa 1/2-2/3of the leafbase, ribbed
at back,excurrentas a serratepoint, in transverse
section with dorsal stereids and ventral substereids.Alar cells enlarged,hyaline or reddishbrown.Innerbasal laminalcells incrassate,rectangular,6-13 x 16-35 ,m, ca. 4-5:1, towards
the marginsnarrower.Upper laminalcells quadrate to irregularlyrhombic, 9-14 x 8-10 ,mm.
Setae 7-8 mm long, flexuose-curved.Capsules
oblong, slightlyinclined, strumose,greenish,becoming brownish and furrowedin age. Spores
light brown, papillose, 13 ,im in diam. Calyptra
fringedat base.
94
Flora Neotropica
FSt7af
FIG. 61. Campylopusflexuosusvar.flexuosus (CampylopodesExsiccatae 7). A. Plant. B. Leaf. C. Leaf-tip.

details = 50 Am.
Distribution (Fig. 62). On soil, earth covered vicariant species, C. comosus (Reinw. &
rocks, on logs, bark at base of trees in various Hornsch.) Bosch & Lac.
types of forests between 1000 and 3000 m eleSpecimens examined: MEXICO. CHIAPAS:Km 5
vation in Mexico, Guatemala,Nicaragua,Hon- alongroadOcosingo-Palenque,Frahm,Camp.Exsicc.
duras, Costa Rica, Panama, Hispaniola, Puerto 7 (B, BING, BUF, C, DUIS, DUKE, EGR, F, FLAS,
Rico, Cuba, Jamaica,Venezuela,Colombia, Ec- G, GOET,GRO,GZU, H, HBG,JE,KRA, M, MEXU,
NICH, NY, PRC, PRE, POZ,
uador, Peru and Bolivia. Outsidethe Neotropics MO, NAM, NFLD,
NE ofTuxtla-Gutierrez,
RNG,
S,
U,
UPS).
Sharp4073a
in North America, West Europe,tropical mon(TENN);Tapachula,Benito Juarez,CerroTres Picos,
tane Africa and scatteredin Australasia,where Eggers & Frahm 792031 (hb. Frahm);San Crist6bal,
C.flexuosus is commonly replacedby a similar, Arzenis.n. (hb. Frahm);N of Ixapastepec,Sharp4547
,zo
95
9c
so
..*..""
,~i
*
din
Preparedby HondrkR.Rpken
__
:p-
.........>
7c
80
>
)l
6C
SC
'
FIG. 62. Distributionof Campylopusflexuosus (@)and C. gardneri(0).

(TENN); San Crist6bal, Gittins 4610 (TENN); Sharp
3591, Breedlove14513, 15152 (MICH);Yerba Buena
55 km NE of Tuxtla-Gutierrez, Sharp et al. 4072, 4166j,

4174, Smith 2802 (TENN). DURANGO:30 km W of El
Salto, Sharp 1852a (TENN). HIDALGO:Sharp 4143,
5682 (MICH),Bucher54b, Pringle 10491 (NY); 8 km
NE of Zacualtipan, Delgadillo 2713 (MEXU); Pueblo

Nuevo Solistahuacan,
Gittins 4591 (TENN).
Morelos National Park near Morelia,
MICHOACAiN:
Norris & Taranto 15508, 15509 (TENN). OAXACA:
Llano de las Flores beyond Ixtlan de Juarez, Sharp M59169
(TENN); near Zempoatepetl 70 km E of Oaxaca, Sharp
et al. 4097 (TENN); 26 km NE of Oaxaca, Delgadillo
677 (NY); La Cumbre 25 km above Oaxaca, Sharp et
al. 2555 (TENN); Serro de San Felipe on Hwy. 175
through Sierra Juarez, Sharp et al. 2566 (TENN); 61
km NE de Llano de las Flores, Delgadillo 861 (MEXU);
26 km NE de Oaxaca, Delgadillo 677 (MEXU); Loma
Grande 26 km N of Oaxaca, Norris & Taranto 16664
(TENN); Cerro de Cuhuatepec SE, Sharp 1432 (TENN).
PUEBLA:Above Huauchinango on Hwy. 130 to Tulancingo, Sharp et al. 1310b (TENN). TAMAULIPAS:
Ranchodel CieloaboveGomezFarias,Nakanishi3616,
Webster104, 107, Sharp 3635, 8746 (TENN). VERACRUZ:Pedrigalde Las Vigas near La Joya, Sharp &
McFarland8674, 8912 (TENN); 17 mi NE of Jalapa
along Rt. 140, Reese 4594 (LAF);above Poza Rica on
Hwy 130, Sharpet al. 1293 (TENN);nearSan Miguel
del Soldado above Jalapa, Sharp et al. 880 (TENN);
Los Huerfanos near Yecuatla, Sharp et al. 3082c
(TENN); 30 km NW Perote, Dill s.n. (DUIS); Sierra
Madre between Jalapa and Perote, Eggers & Frahm
s.n. (hb. Frahm). XALAPA:Sierra de Chiconquiaco be-
tween Santa Rita and Bella Luz, Hermann 28837

(TENN).
GUATEMALA.
GUATEMALA:
Volcan de Pacaya,
Sharp 2073
Standley 58424 (FH). QUEZALTENANGO:
(MICH).
HONDURAS. Cerro Uyuca, Hutzel s.n. (MICH);
Morazan,Standley 13726, 14709 (MICH).
Km 73 on Inter American
COSTA RICA. CARTAGO:
HighwaySE of El Empalme,Crosby6356 (MO). SAN

Jost: Cerrode la Muerto3 mi NW of Villa Mills,Koch
5088 (NY).
96
Flora Neotropica
PANAMA. CHIRIQUi:Cerro Fortuna, Salazar et al.
649 (MO).
CUBA. Loma del Gato, Sierra Maestra, Clement
14550 (NY); TrinidadMtns., Santa Clara,Aguacate,
Britton5389 (NY); Pinar del Rio, Rio Guao, Britton
et al. 10126 (NY).
DOMINICAN REPUBLIC.Samana,Las Terrenas,
Schuster17a (NY);La Vega,9 km NNW of Constanza,
Buck5361 (NY);roadto SanJose de Ocoa 13 km from
Valle Nuevo, Norriset al. 7098 (NY).
PUERTORICO.MaricaoInsularForest,Steere5482
(NY); between Maricao and Sabana Grande, Steere
5745 (NY); Cayey-Guayanaroad, Steere 5085 (NY).
COLOMBIA. ArNIOQuIA:12 km E de Sons6n,
5?40'N,75?15'W,Churchill& Sastre-DeJests 13007
Near Ziraquira,Schultes11481
(NY). CUNDINAMARCA:
(NY); Guadelupe,Onraedt6309 (hb. Frahm);20 km
SE of Bogota on road to Villavicencio, Steere 1854
(NY).
VENEZUELA.BOLiVAR:
Steyermarkdiv. no. (NY).
CARACAS:
Onraedt 4672 (hb. Frahm). MERIDA:Pira-
mo La Negra above Bailadores, Griffinet al. 2142

(FLAS);Piramo de Don Pedro 20-22 km ESE of El
Morro, Ruiz-Terdn& Lopez-Figueiras8694 (FLAS).
TRUJILLO:
Paramode la Morita,Ruiz-T. & Lopez F.
7445 (FLAS).
PERU. Cushi, trail to Pozuzo, Bryan 680b (F).
AMAZONAS:Road Chachapoyas-Cajamarca
above
Leimebamba,Frahm, Camp.Peruv.Exsicc. 9 (ALTA,

B, BM, BUF, DUKE, EGR, F, FLAS,G, GRO, GZU,
HGB, KR, NAM, NICH, NY, MEXU, NFLD, PC,
km
POZ, RNG, S, U); road Cajamarca-Chachapoyas
413, Frahm et al. 367 (B);km 417, Frahm et al. 585,
353, 375, 360 (B); pass between Balsas and Leimebamba,Frahm1977 (hb. Frahm);above Leimebamba,
Celendin, HegePhilippi 2339, 2330 (B). CAJAMARCA:
wald6642 (hb. Frahm).Cuzco: MachuPicchu,Hegewald 8860 (hb. Frahm).

Prov. de Chapar6, Pampa
Tambo, Hermann25246 (hb. Frahm).
24b. Campylopus flexuosus var. incacorralis

(Herzog) J.-P. Frahm, Bryol. Beitr. 7: 35.
1987.
Figs. 62, 63.
CampylopusincacorralisHerzog,Beih. Bot. Centralbl.
26: 52. 1910. Type. Bolivia. Incacorral,Herzog285
(holotype,JE;isotype, B).
Campylopusreflexus Brotherusin Herzog, Biblioth.
Bot. 87: 23. 1916. Type. Bolivia. Incacorral,Herzog
4988 (holotype,JE;isotypes, B, FH, H-BR).
This variety differs from var. flexuosus by
plants growing in more dense tufts with appressed foliate stems, pitted basal laminal cells,
upper laminal cells arranged very distinctly in
rows and the perichaetial leaves ending in a hyaline tip.
Distribution (Fig. 62). Mostly above the forest
line in subalpine shrubs or paramos in Venezuela, Colombia, Peru and Bolivia. Also de-
scribedas CampylopussubperichaetialisBizot &

Kilbertus,from the East African Mountains.
Sierra
Nevadadel Cocuy,AltoValleLagunillas,
Cleef5515,
8775 (U); CordilleraOriental,carreteraRamiriquiMiraflores,v. d. Hammen4047 (U); Paramode la
Sarna,Cleef9318 (U); Paramode Chisaca,Murillo53
(NY);withoutlocality,Weir347 (NY).
VENEZUELA.
Cumbreo largodel Rio
BOLivAR:
Churum,Steyermark93271 (NY); Kukenantepui,
Brewer4943a (FLAS).MERIDA:
Sierrade Santo Domingo,piramode Mucubaji,
Griffinetal. 893 (FLAS);
SierraNevadade MeridabetweenstationLaAguada
andLaMontafia,
et al. 261 (FLAS);
Piramode
Griffin
los Conejos,Ruiz-Terdn7682 (FLAS);ParcqueNacionalde Mucubaji,Onraedt5034 (hb. Frahm);La
Carbonera
areanearLa Azulita,Griffinet al. 2003,
2237 (FLAS).TACHIRA:
Piramo de Tama above the
village of Paez, Griffinet al. 293 (FLAS).TRUJILLO:
Paramode Cend6,Lopez-F.12965 (FLAS);Paramo
de La Morita,Ruiz-T. & Lopez-P. 7444 (FLAS).

PERU. AMAZONAS:
Road Cajamarca-Chachapoyas
km 403.5, Frahm et al. 386 (B). ANCASH:
Cordillera
Blanca, Laguna Llanganuco,Frahm, Camp. Peruv.
Exsicc.12 (ALTA,B, BM, BUF, DUKE,EGR,F,

FLAS,G, GRO,GZU,HBG,KR,NAM,NICH,NY,
MEXU, NFLD, PC, POZ, RNG, S, U). AYACUCHO:
PassbetweenAynaandTapuna,Hegewald9008(hb.
Hegewald).
Althoughthe type specimensof C. incacorralis

and C. reflexus were both collected in Bolivia
(same locality) at only 2200 m, both species are
regardedas subalpine-alpinevarietiesof C.flexuosus. This is supportedby identical collections
in the northernAndes in habitatsas paramosor
wet punas and also by the presenceof this taxon
in subalpineheath forestsin Tanzania,Uganda,
Zaireand Mauritius.It is not out of the question
whetherthe var. incacorralismightultimatelybe
an alpine growth form of C. flexuosus, but this
can not be decided merely by herbariumspecimens. Since it is usually distinguishableby the
more compact growthform and the pitted basal
laminal cells, it is recognizedhere as a variety.
However, some forms are difficultto interpret.
Flagelliferousbranchesare also found in the var.
incacorralis.
Campylopusflexuosus is closely related to C.

arctocarpus,from which it differsmainly by the
transversesection of the costa (ventralsubstereids vs. stereids).Both species are characterized
by subquadrateupper laminal cells arrangedin
distinct rows, the serrate leaf tips and the incrassate rectangularbasal laminal cells. Both
producesporophytesof similar shape relatively
97
ic
3-
m.,
>
.~~~~~~~~~~~
/Ikll
FIG. 63. Campylopusflexuosus var. incacorralis(Griffinet al. 261, FLAS). A. Plant. B. Leaf. C. Leaf-tip.
details == 50
um.
50,um.
frequently.WhereasC. flexuosus var. flexuosus

has non-pitted basal laminal cells, C. flexuosus
var. incacorralishas slightly pitted and C. arctocarpushas distinctlypitted basal laminalcells.
Whereasthe rangeof C.flexuosus extends to the
temperateregionsin North Americaand Europe,
C. arctocarpusis strictlyof tropicaldistribution.
However, both species show more or less the
same rangein the Neotropics.
Campylopusflexuosus is very variable. Spec-
imens from rain forestscan reach 5 cm, whereas

those from dry oak-pine forest, as for example
from Mexico, reach less than 1 cm. Such small
forms produceflagelliferousbranchesabundantly. They are interpretedas stressed forms in either unfavouredecologicalconditions or seasonal formsproducedin dry seasons.Obviouslysuch
forms are lacking in hygric conditions and annually wet conditions.
Only once (Frahm 1977) have propaguliferous
Flora Neotropica
98
i.
TV
FIG. 64. Campylopusfragilis ssp. fragilis (Onraedt5772, hb. Frahm). A. Plant. B. Leaf. C. Leaf-tip. D.
details = 50 Am.
leaves in the comal tufts similar to those of C.

fragilis been found in C. flexuosus. This demonstratesthatall the speciesof Campylopusseem
to have possibilities for vegetative propagation
such as flagellatebranches,isomorphousand anisomorphous brood leaves or clustersof leaves.
25. Campylopus fragilis (Bridel) Bruch &
Schimper, Bryol. europ. 1: 164. 1847.
Key to the Subspecies of

Campylopus fragilis
1 Upper laminal cells quadrate. .. 25a. ssp.fragilis.
1 Upper laminal cells oval to short rectangular,225b. ssp.fragiliformis.
3:1 ....................
25a. Campylopus fragilis
Schimper ssp. fragilis
(Bridel) Bruch &

Figs. 64, 65.
99
SystematicTreatment
Dicranum
fragileBridel,J. Bot. 1800(2):296. 1801. near El Puerto,Sharp 642 (NY); between Perote and
Type.Withoutlocality,hb. Dicksonno. 40 (lecto- Jalapa,Eggers & Frahm s.n. (hb. Frahm).
GUATEMALA.El Quiche,aboveNibaj,Sharp2484
type,BM).
CampylopusfimbriatusMitten, J. Linn. Soc. Bot. 12:
88. 1869.Type.Ecuador.Quito,Spruce51 (lectotypusnov.,NY).
(FH).
COSTA RICA. La Fuente,Alfaros.n. (NY).
DOMINICAN REPUBLIC. BARAHONA:
Monteada
Nueva nearPolo, Judd 1123 (NY); nearcrestof Sierra

SanCris- de Neiba, Norriset al. 6001 (NY); Sierrade Baronco,
nol.37:119.1910.Type.Mexico.Chiapas,
CampylopussubturfaceusCardot, Rev. Bryol. Liche-
t6bal,Munchs.n. (holotype,PC).
Campylopusretinervis(C. Miiller) Paris, Ind. bryol.
retinerve
C.Miller,Bull.
Suppl.96. 1900.Dicranum
Herb.Boissier5: 552. 1897.Type.Jamaica.Cinat B;isochona,Harris11025(holotype,destroyed
type,NY).
CampylopustunariensisHerzog,Biblioth.Bot. 87: 19.
1916.Type.Bolivia.Tunarisee,
Herzog3430 (lecB, H-BR).
totype,JE;isolectotypes,
Plants in small, dense, bright to yellow green
tufts 0.5-3 cm high, densely foliate, with many
short branchesat the apex forminga thick penicillate comal tuft, with clustersof small deciduous leaves often produced in the comal tufts.
Leaves somewhat flexuose when dry, about 5
mm long, narrowedto the insertionand tapered
from an ovate-lanceolatebase to a more or less
long subula,the lowerhalf of the leafshiny-white
and noticeable when dry. Margins entire, incurved above. Costa excurrent,slightly toothed
at the extreme tip, filling 1/2-2/3of the leaf base,
in transversesection with lax ventralhyalocysts

and dorsal substereids.Alar cells scarcelydifferentiated. Basal laminal cells large, lax, thinwalled, hyaline, 13-16 x 61-74 ,lm, in 3-5 rows
narrowerat margins.Upper laminal cells quadrate, 10-12 x 13-15 ,m.
Setae 5-8 mm long. Capsulesellipsoidal,furrowedwhen empty. Spores 14-16 ,m, brownish,
finely papillose. Calyptrafringedat base.
Distribution(Fig. 65). On soil, earth covered
rocks, in rock crevices, at the base of trees, on
stems of fern trees, on decayingwood and peat
from montane forests to the high alpine belt
(1000-4600 m) in Mexico, Guatemala, Costa
Rica, Cuba, Jamaica, Puerto Rico, Haiti, Venezuela, Colombia, Ecuador, Peru and Bolivia,
also in North America, Europe,tropical Africa
and Asia; in SE Asia replacedby vicariantspecies.
Zanoni et al. 19025 (NY). ESTRELLETA:

Sierra de Nei-
ba 18?41'N, 71?43'W,Reese 15158 (NY). INDEPENDENCIA: 13 km from Valle Nuevo on the road to S.
Jose, Norris et al. 7107 (NY). LA VEGA:La Culata,
Buck5373 (NY);44.7 km S ofConstanza,Steere22709
(NY); 3 km W of La Cienaga,Norriset al. 5377 (NY);
4.7 km S of Constanza,Shaw 5439 (NY); vicinity of
pyramids, 13.8 km S of Valle Nuevo, Steere 22709
(NY); 40 km S ofConstanza, 18045'N,70037'W,Buck
5311, 5323 (NY);PicoAlto Bandera,18?44'N,70038'W,
Mojia 13262 (NY); Sierrade Neiba alongHaitianborder, Norris 6197, 6148, 6000 (NY); between Hondo
Valle and Angel Felix, GasbarroB 10584 (NY); vicinity of Constanza,Allard16572 (NY); 47.5 km N of S.
Jose de Ocoa, 18?43'N,70043'W,Zanoni 16462 (NY);
9 km N of Constanza,Buck 5373 (NY). PERAVIA:La
Nevera 19 km S of Valle Nuevo, Steeres.n. (NY).
HAITI. DEP. DU NoRD:Vic. Marmelade,Leonard
8234 (NY); between Furey and Kanshoff,Mackaness
213 (NY).
PUERTORICO.ReservaFlorestalde Guilarte,Mun.
de Adjuntas,Steinsson3799 (NY);upperRio de Maricao, Steere 5649 (NY).
Sierra Maestra, Pico Turquino,
CUBA. ORIENTE:
Ekman5469, 5405, 5473, 14565, 11204, 11233, 11248,
11250, 11222 (NY); between Rio Oro and Rio Yao,
Ekman 7110 (NY);SierraMaestra,Pico Cuba,Samek
s.n. (hb. Frahm).
JAMAICA. ST. ANDREW PARISH:S of Hardwar Gap
18?04'N,76?42'W,Buck5733 (NY);slopesof SirJohn,
Britton 1169 (NY); Cinchona,Nichols 120 (NY); Sir
John'sPeak,Crosby3039 (MO);BlueMtns.,Patterson
53a (NY); Moody's Gap, Britton905 (NY).
COLOMBIA. ARAUCA:
QuebradaEl Play6n, Cleef 10068 (U). BOYACA:Peia
de Arical N de Vado Hondo, Cleef9476 (U); Paramo
de la Rusia NW de Duitama, Cleef 7264, 7329, 7337,
7357 (U); Vado Hondo, ParamoentrePefiade Amical
y Alto de Mogotes, Cleef 9259 (U); SierraNevada del
Cocuy,Grubb& GuymarB 234 (FH);Paramode Concavo, Cleef 8540 (U); QuebradaBocatoma, Cleef &
Florschitz 5846, 5856, 5885 (U); Alto de la Cueva,
Cleef 9977 (U); Paramode Pisva, carreteraSocha-La
Punta,Cleef4267 (U). CALDAS:Nevado del Ruiz, Cleef
& 'tHart 2377, 2517 (U). CAUCA:Mt. Purac6,Pennell
& Killip6657 (NY); Cleef& Fernandez653 (U); CordilleraCentral,Nevado del Huila, Steere 7911 (NY).
CUNDINAMARCA:
Paramo de Sumapaz, Cleef 1556,
Specimens examined. MEXICO. CHIAPAS:Yerba
1600, 2615 (U); Cleef&Jaramillo236, 254 (U); Para-
al. 4166, 4701c, 4214 (TENN); 11 mi N of Teopisca,

N of Oaxacaat
LeDoux et al. 2164 (TENN).OAXACA:
SierraJuarezGap, Smith et al. 4777a (TENN).VERACRUZ:3 km NE de Coatepec, Vivenos76a (TENN);
6469 (U); Paramode Palacio,Cleef2338 (U); Paramo

de CruzVerde, Cleef2834, 2927, 3320, 3303 (U); Cerrosen el lado orientalde la Represadel Neusa, 60 km
NNE of Bogota, van der Hammen & Jaramillo 3795
Buenaabout55 kmNE of Tuxtla-Guti6rrez,

Sharpet mo entreCoguay San Cayetano,Cleef73, 6235, 6360,
Flora Neotropica
100
IX
90
3f?
^^
80
70
60
so0
'I
--
----
------------30
R.Rypkema
; _Prepared
byHendrik
FIG.65.
:o
'41~M
DistributionofCampylopusfragilisssp.
fragilis
-I-
/-^
and
fragilissspfragiliformis(0).
(U); Bogota, Weir180 (NY, F);Paramode CruzVerde,

CarreteraBogota-Coachi, Cleef 2929, 3344 (U); Boqueronbei Bogota, Ttroll2174 (B);Fermede Sasaima,
onraedt5969 (hb. Frahm);Guadelupe,Onraedt6007,
6301, 6295 (hb. Frahm); Subachocque,Cuchilla El
Tablazo, Linares & Bulla 402, 485 (COLO).HUILA:
Cordilleradel Buey, Paramo de las Papas, Bishop 1
(FLAS).META:Paramode Sumapaz,Cleef7903, 7923,
7963, 8282 (U); Cerro Nevado del Sumapaz, Cleef
1285, 7666a (U).
0?29'S, 77?51'W,Churchill& Sastre-De-Jestis13617

(NY); valley of Rio Chingualeast of El Pun, Steere
8374 (NY); road between Quito and Baeza, 78?09'W,
0?17'S,SteereE 164 (NY); 0llgaard & Balslev 10059
(AAU, NY); N-facingridgeofCerroSumaco,77?39'W,
0?35'S, Lojtnant & Molau 12688 (NY, AAU). PICHINCHA:
E of San Miguel de los Bancos, Steere &
Aguada, Onraedt5772 (hb. Frahm);Paramo El Batall6n, Ruiz-Terdn8328 (FLAS).
Frahm 823892 (hb. Frahm). AYACUCHO:

Aina between
VENEZUELA. MERIDA:Sierra Nevada, environ de
Balslev 25651 (NY); NW corer of volcan Pichincha,
Steere & Balslev 26011 (NY). Cotopaxi, 0?40'N,

78?30'W,Balslevet al. 3380 (NY).
PERU. ANCASH:
Road Catac-Chavinde Huantar,
HuankaandRio Apurimac,Killip&Smith22738 (NY);

Lomas of Atiquipa,Hegewald8561, 8562 (hb. HegePara- wald).CAJAMARCA:
Las Lagunas,Hegewald6190 (hb.
2?46'S,Lojtnant& Molau 14821 (NY). CARCHI:
mo ElAngelon roadElAngel-Tulcan,78?54'W,0?4I'N, Hegewald).Cuzco: Machu Picchu, Hegewald 8810,
Co- 8840, 8847 (hb. Hegewald).JUNiN:Pampa HuicushHolm-Nielsenet al. 5434 (AAU, NY). COTOPAXI:
topaxi Nat. Park,Brako4400 (NY). NAPO:Paramode cancha near Tarma, Hegewald 6278 (hb. Hegewald).
JaramilloSE of LagunaPisayambo,Laegaard53326A PASCO:
Along road from Cerrode Pasco to Lima N of
(NY); km 45 on roadfrom Salcedoto Napo, Laegaard Huayllay, 11?0'S,76?20'W,Vitt21741 (ALTA).
54159F (NY); 5.6 km S of Baeza on road to Tena,
BOLIVIA.LA PAZ:Rio Consata, 15?41'S,68?43'W,
ECUADOR. AZUAY:Laguna Toreadora, 79?13'W,
SystematicTreatment
101
Lewis79-1210(F);Sorata,15?45'S,68?42'W,Lewis
Bryol.Lichenol.2: 442. 1981.Type.
Cryptogamie
Brazil.Serrado Itatiaia,AgulhasNegras,Dusens.n.
79-1263(F);NevadoJankhoUma above MinaSt.
Lewis79-1512A(F).TARI- (holotype,B).
Antonio,1551 'S,68?34'W,
W of EntreRios,21?28'S,64?12'W,
JA: Casteon
Lewis ? CampylopussphagnicolaHerzog,Repert.Spec.Nov.
79-513,79-536(F);Canyon5 mi SWof Tablada,SW
RegniVeg. 21: 28. 1925. Type.Brazil.Serrados
of Tarija,21?36'S,64?48'W,
Lewis79-603(F).
Orgaos,MorroAssu, v. Liitzelburg6760b (holotype,
JE).
Although this species produces sporophytes
Plantsyellowish-green,in loose tufts, 1-2.5 cm
relatively rarely, hooked, boomerang-shaped
brood leaves are mostly frequentin the axils of high. Stems densely foliate, reddish radiculose
the upper leaves, giving the plants a typical ap- below. Leaves 4-5 mm long, concave, from an
pearance.In alpine regions,the plants can form ovate base lanceolate, contractedto a scarcely
very low rosettes producing abundant brood dentatetip. Costafilling1/2of the leaf width, conleaves. Such plants have been describedand re- tracted at base, in transversesection as in ssp.
cordedas C. tunariensis.However, observations fragilis. Alarcells scarcelydeveloped.Basallamin the field show that such plants occur on dry inal cells rectangular,thin-walled and hyaline,
substratesas tussocks of peat or decayedgrasses 10-14 x 70-90 gum.Upper laminal cells oval,
and that there are all possible intergradationsto rhomboid or shortly rectangular,9-10 x 13be found to normallydeveloped plants growing 18(-25) ,m, ca. 2-3:1.
Setae 4 mm long, yellowish,flexuose.Capsule
in more favoured conditions. Tunariensis-like
1
mm
long, ovate.
plants can also be found in the lowlands (even
Distribution
(Fig. 65). Known only from SE
in temperateregions)as an expressionof stressed
has been found in alpine regions
where
it
Brazil,
habitats or unfavourableseasons.
The leaf tips can be elongatedin certainforms on decayingTyphasp., at the base of rockycliffs,
and can even be hyaline, which may cause con- on swampygroundand on humic soil in Drimys
fusionwith other,usuallyhyaline-tippedspecies. scrub.
Such forms have been collected in Jamaicaand
Specimensexamined.BRAZIL.MINASGERAIS:Pico
the Dominican Republic.
da Carapuca,Serrado Caraca,20"07'S,43?31'W,Vital
The costa is conspicuously widened at the 7706 (SP). Rio DEJANEIRO:
ParqueNacional Itatiaia,
broadestpartof the leaf and narrowedto the leaf AgulhasNegras,22?23'S,44?14'W,Frahm1636 (hb.
Vital 7387, 7395 (SP); Vitt 21561 (ALTA);
base. Its largeventralhyalocystsareconspicuous Frahm).
6793(JE,
SerradosOrgaos,MorroAssu,v.Liitzelburg
in ventral view even without dissecting.
PC).
Campylopusfragilisis easily recognisedby the
Due to the lack of Campylopusfragilis ssp.
white leaf bases consisting of lax, hyaline cells,
which function for rapid water uptake. These fragilis in SE Brazil,ssp.fragiliformiscan be inbasal laminal cells are sharply delimited from terpretedas a vicariantraceof ssp.fragilis in this
region. It is mainly differentiatedby the shape
the firm, small, quadrateupperlaminal cells.
The usual matter of vegetative propagationis of the upper laminal cells, which is quadratein
but oval in ssp. fragiliformis. The
by means of anisophyllous brood leaves; how- ssp. fragilis
has alwaysbeen collectedwith
ssp.
fragiliformis
been
have
also
branches
ever, microphyllous
and
the
presenceof broodleaves has
foundin Holm-Nielsenet al. 5434 fromEcuador. sporophytes
not
been
observed.
yet
This is similar to C. flexuosus, where microCampylopus luetzelburgii has been described
phyllous branches usually are produced, but
as
a separatespecies based on (1) short rectanbrood leaves have been found only once.
gular instead of of oval upperlaminal cells and
a transversesection of the costa withoutdor25b. Campylopus fragilis (Bridel) Bruch & (2)
sal stereidsinsteadofsubstereids.However,both
Schimperssp.fragiliformis(J.-P.Frahm)comb.
may fall into the rangeof variationof C.fragilis
nov.
Figs. 65, 66.
ssp. fragiliformis.There might be also a differCampylopus
fragiliformisJ.-P. Frahm,Rev. Bryol. entiation of sterile and fertile plants, for all maLich6nol.45: 147. 1979.Type.Brazil.Rio de Ja- terial named C. fragiliformisbears sporophytes
neiro:ParqueNacionalItatiaia,AgulhasNegras, whereasthe two
specimensof C. luetzelburgiiare
Frahm1637(holotype,hb.Frahm;isotypes,ALTA,
of both taxais also supported
sterile.
The
identity
B, MO,SP,U).
CampylopusluetzelburgiiHerzogex J.-P. Frahm, by the fact that the type localities of both taxa
Flora Neotropica
102
FIG. 66. Campylopusfragilisssp.fragiliformis(Frahm 1636, hb. Frahm).A. Plant. B. Leaf. C. Leaf-tip.D.

Transversesection of leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule.Unlabeled scale bar for
microscopicdetails = 50 Mm.
are the same. It can, howver, not be fully excluded that there may exist two or even more
genotypicallydifferentpopulations,as a resultof
separation from the populations of C. fragilis
s.str. in the Andes.
The type specimenof Campylopussphagnicola
consists of a few stems only, which weregrowing
amongst Sphagnum plants and therefore very
lax. The shape of the basal laminal cells and the
structureof the costa leads to the suspicion that

these plants belong to C. fragilis ssp. fragiliformis, which occursin the same regionbut usually
in drier habitats.
26. Campylopusgardneri(C. Muller)Mitten, J.
Linn. Soc. Bot. 12: 83. 1869. Dicranumgardneri C. Miller in C. Muller & Hampe, Bot.
Zeit. 15: 379. 1857. Type:Brazil.Pernambuco:
103
"A
FIG. 67. Campylopusgardneri(Reese 16452, LAF). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 im.
chaetialleaves with broad sheathingbase. Costa

1/3 of the leaf width, in transverse section with
ventralhyalocystsand a dorsal band of stereids.
Alarcells fragile,hyaline,thin-walled.BasallamPlants very small, in appearancelike a species inal cells rectangular,6-13 x 16-45 Mm,with
of Dicranella. Stems to 1 cm high, reddish to- severalrows of narrow,hyaline,thin-walledcells
mentosebelow. Leaveshomomallous,comose at at margins. Upper laminal cells subquadrateto
stem tips, 3.5 mm long,narrowlylanceolate,end- rhomboid, 6-10 x 10-16 Am, ca. 1.5-2:1.
ing in a long scarcely serrate hyaline tip. PeriSierrade Araripe, Gardner20 (holotype, destroyed at B; lectotypus nov., NY; isolectotype, S).
Figs. 62, 67.
Flora Neotropica
104
:i
FIG. 68. Campylopusgastro-alaris(Vital 7617, SP). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Am.
Distribution (Fig. 62). On siliceous substrates
such as rocks and boulders, preferably sandstone,
on sandy soil in savannah vegetation in the arid
parts of Brazil and also in Paraguay, at low elevations between 250 and 600 m.
Mun.de San
Specimensexamined.BRAZIL.BAHLA:
Desiderio,km 173 on roadBR 020, Vital6095. CEARA:
Parque Nacional de Apodi, Vital 2910 (SP). GOIAs:
Mun. de Tocantinopolis, Vital 2992 (SP). MATO
GRosso: On roadto Cuiaba,Pranceet al. 19341 (NY).
PARA:Serra do Cachimbo, km 774 on Cuiaba-San-
tar6mhighway,9?30'S,54?55'W,Reese 16023, 16006

(LAF);km 887 on Cuiaba-Santaremhighway,8?45'S,
54?57'W,Reese 16487, 16452 (LAF).PIAui:Mun. de
Bor Jesus, 9?28'S,44?34'W, Vital8225 (SP).
PARAGUAY.ALTOPARANA:
ReservaBiologicadel
Itab6,25?05'S,54?44'W,Buck 12385 (NY). AMAMBAY:
ParqueNacional CerroCora, 22?40'S,56'00'W, Buck
12503 (NY). CAAGUAZU:
Coronel Oviedo, Bordas 235
(NY).
Since the sporophyte of this species is not
known, the taxonomic position is not entirely
clear. It vegetatively resembles species of Sphae-
105
II
_.-
'
ec
SC
31
10
*.::..
. . ......
- -- - - - -- -
..........
FIG.
69.Distribu
69.
GoG
-sr-ari
iono
Ds ui
FIG.69.Distributonf
6(0)
70.
s()
.htrs
..
SC
'
...
"o
rothecium or Campylopussubg. Campylopidulum such as C. carolinae, which occurs also in

arid regions.In the field it may be confusedwith
species of Dicranellaor Microcampylopus,from
which it is distinguishedmicroscopicallyby the
presenceof conspicuous alar cells.
(O
amyo
:'':
reachingup to leaf tips. Costa narrow,filling /4

to /3of the leaf width, in transversesection with
a multilayeredbandof ventralstereidsandgroups
of dorsalstereids,ridgedand serrateat back.Alar
cells stronglydifferentiated,hyaline or reddish.
Basal laminal cells elongate rectangular,incrassate and pitted, 10-16 x 54-80 tm, narrowed
towards the margins.Upper laminal cells elongate oval, ca. 6:1, 6-10 x 32-48 um, incrassate
and pitted.
Distribution(Fig. 69). On loose sandalongriver banks, in periodicallyinundatedstreamsand
in rockfissuresin waterfalls,foundonly in gallery
forests in the cerradoregion of Brazil.
27. Campylopusgastro-alaris(C. Miiller)Paris,

Ind. bryol. ed. 2, 1: 310. 1904. Dicranumgastro-alare C. Miiller, Hedwigia 39: 256. 1900.
Type. Brazil. Minas Gerais: Serrade Caraca,
Ule 1361 (holotype,destroyedat B; lectotypus
nov., H-BR).
Figs. 68, 69.
Plants to 3 cm high, in dense tufts, brownish
below, greenishabove. Stems equallyand densely foliate, at tips appressedfoliate formingacute
Plateau
Specimensexamined.BRAZIL.AMAZONAS:
tips. Leaves ca. 6 mm long, broadly lanceolate, of SerraAraca, 0?51'N, 63?21'W,Samuels et al. 290
widest at midleaf, contracted at base. Lamina (NY). GOIAS:N of Alto Paraiso, Irwin et al. 33078
106
Flora Neotropica
FIG. 70. Campylopus gemmatus (Ule 1789, H-BR). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section of
leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule. Unlabeled scale bar for microscopic details =
50 nm.
BetweenSerraand Datas, S of
(NY). MINASGERAIS:
Diamantina, Frahm, Camp.Bras. Exsicc. 10 (ALTA,
B, C, DUIS, FLAS,GOET,GZU, H, HBG, INPA, JE,
KR, M, MO, NFLD, NICH, NY, PC, S, SP, TENN,
U); Mun. de Itamb6, 19?27'S,43?25'W, Vital 7617
(SP). PARA:Serrado Cachimbo,Rio Braco Norte, at
air base, 9?22'S,54?54'W,Reese 16436 (NY).
28. Campylopus gemmatus (C. Miiller) Paris, Ind.
bryol. 82. 1900. Dicranum gemmatum C.
Muller, Bull. Herb. Boissier 6: 34. 1898. Type.
Brazil. Serra do Itatiaia, Ule 1789 (holotype,

destroyed at B; lectotypus nov., H-BR).
Figs. 70, 71.
CampylopusarachnoideusHerzog, Rep. Spec. Nov.
Regni Veg. 21: 29. 1929. Type. Brazil. Serra dos
Orgios, MorroAssu, v. Liitzelburg6688 (holotype,
JE; isotypes, B, S).
Plants 1-2.5 cm high, yellowish green, in loose
tufts. Stems single or divided near the base,
107
90
60
70
80
40
50
30
-K----
y:.:--
20 0 4 00
6 00 60 0 10 00 k m
_ ____
.'
?
______
F 400 500 600 miles1 G0200 300

Prepared
by Hendrik
R.
-RypkemaA
"
%
'' ' '
'
0i_____
\
\
'[
ti.^ __ _
^.
/'
\-10 \
,.
--- s l
.--,2
FIG. 71. Distributionof Campylopusgemmatus(O) and C. huallagensiss. lat. (-).
loosely foliate with distant, patent leaves, comose at tips. Vegetative propagationby often
massesofpropaguliferousplantsproducedin the
axils of the comal leaves, covering the tufts like
spider nets. Leaves 3-5 mm long, from narrow
base longlylanceolate,endingin a subhyalinetip,
entire at margins. Costa filling half of the leaf
width, in transverse section with large ventral'
hyalocystsand a dorsalband of substereids.Alar
cells consistingof few large,brownishcells. Basal
laminal cells elongate, rectangular,thin-walled,
6-16 x 35-80 ,m, narrowerat margins.Upper
laminal cells rhomboid, incrassate, 3-6 x 10-32

JIm.
Distribution (Fig. 71). Epiphytic on branches
of shrubs and nodes of bamboo (Chusquea sp.),
also on rocks in the montane forests of SE Brazil
between 800 and 2300 m elevation.
Parque Nacional de Itatiaia, Agulhas Negras, Vitt21520
(ALTA); Parque Nacional Tijuca, Pico de Papagaio,
22050'S, 43?20'W, Landrum 2208 (MICH).
Flora Neotropica
108
E
A
FIG. 72. Campylopusgriseus ssp. griseus (Vital 5603, SP). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
50 ~m.
Campylopusgemmatusis closely relatedto C. both species are vicariant. Therefore,it can be

asperifolius. Both species are characterizedby postulated that both species have been derived
the fluent transition between the basal and the from a common ancestor. Campylopusasperiupperlaminal cells, alarcells protrudinginto the folius differsby sharplyserrateleaf tips (smooth
costa and the presence of apical brood leaves. in C. gemmatus), longer leaves and narrower,
Withinthe genus Campylopus,especiallythe lack more linear upperlaminal cells, 10 times longer
of a delimitation of the upperand basal laminal as broad or even longer.Becauseof these distincells is a unique character.Besides,the rangesof guishing characters,neither taxa is recognised
109
here as subspeciesof the same species.They can,
however, be undoubtedlyregardedas a species
pair from the Andes and SE Brazil.
Distribution (Fig. 73). In wet places on soil

and rocks along small creeks or in damp grassland (campos),often aroundspringsin southern
29. Campylopus griseus (Homschuch) Jaeger, Brazil,Uruguayand Argentina.
Ber. Thatigk. St. Gallischen Naturwiss. Ges.
GERAIS:
Mun.
Specimensexamined:BRAZIL.MINAS
1870-1871: 443. 1872.
de SaoJoaodel Rei, 21?02'S,44?09'W,Vital8862
(SP);
Serra de Caldas, Mosen 23 (H-BR). PARANA:Mun. de
Rio Negro,km 98 alongBR 116, Vital5603 (SP);Mun.

de Guaira,Sete Quedas, Vital8355 (NY); Mc. de Jaguariaiva,km 212 along road PR 11, Vital5839 (SP).
PIAui:
NacionalSete Cidades, Vital5411 (SP).
1 Upperlaminalcells not pitted. ... 29a. ssp.griseus. RIODEParque
JANEIRO:
Itatiaia, Bueno Rangel, Glaziou 9076,
1 Upperlaminalcells stronglypitted. .............
9077, 7068 (PC);Apiahy,Puiggari975 (G, MO). Rio
............................
29b. ssp. ingeniensis. GRANDE DO SUL: Mun. de Pinheiro
Machado, 31?29'S,
53?40'W,Vital9148, 9153 (SP);Mun. de Pedro Oso29a. Campylopus griseus (Homschuch) Jaeger rio, 31?43'S,52?53'W,Vital9107, 9117 (SP);Mun. de
ssp. griseus
Figs. 72, 73. Vacaria,28?24'S,5051 'W, Vital2049, 9384 (SP);Cacapava do Sul, Guerraet al. 2026 (LBLC);Vacaria,
Campylopusgriseus (Horschuch) Jaeger, Ber. Tha- headwatersof Rio Uruguay, Sehnem 5901 (LBLC);
tigk.St. GallischenNaturwiss.Ges. 1870-1871:443. Rio Touros,Sehnem5965 (LBLC);Rincaodo Inferno,
1872. Thysanomitrion
griseumHorschuch, Fl. bras. Lavrasdo Sul, Sehnem 11911 (hb. Frahm);Mun. de
1: 16. 1840. Type:Uruguay,Montevideo,Sellows.n. Lavrasdo Sul, 30?58'S,53?30'W,Vital9199 (SP);Mun.
(holotype,destroyedat B; lectotypusnov., KIEL). de Sep6,30?19'S,53?14'W,Vital9211 (SP);Caxiasdo
Campylopuslapidicola (C. Miiller)Paris, Ind. bryol. Sul, 29?07'S,51"07'W,Vital9316 (SP);Mc. de Canela,
ed. 2, 1: 315. 1904. DicranumlapidicolaC. Miller, 29022'S, 50?47'W, Vital 9295 (SP). SANTACATARINA:
Hedwigia39: 358, 1900. Campylopusgriseus(Horn- Lajes,km 3978 along road BR 116, Vital5664 (SP);
schuch)Jaegervar.lapidicola(C.Miller)J.-P.Frahm, along road rio Bonito-Lebon Regis, Frahm s.n. (hb.
Bryoph.Bibl. 5: 61. 1975. Type. Brazil.Sta. Catar- Frahm); Orleans, Sehnem 12648 (LBLC);between
ina: Orleans,Ule 701 (holotype,destroyedat B; lec- Campos Novos and S. Miguel del Oueste, Frahm,
Camp. Bras. Exsicc. 3 (ALTA, B, C, DUIS, FLAS,
Campylopusrecurvipilus(C. Muller)Paris, Ind. bryol. GOET,GZU, H, HBG,INPA,JE,KR, M, MO,NFLD,
ed. 2, 1: 325. 1904. DicranumrecurvipilumC. Miil- NICH,NY, PC, S, SP, TENN, U): Mc. de Curitibanos,
ler, Hedwigia39: 259. 1900. Type. Brazil. Sta. Ca- 27?17'S,50?27'W,Vital 9429 (SP);Mc. de S. Cecilia,
26?56'S, 50?27'W, Vital 9442 (SP). SAo PAULO:Camtarina:Novo Friburgo,Peters 1871 (n.v.).
(C. Muller)Paris,Ind. bryol. pos do Jordao,Pico de Itapeva,Schifer- Verwimp6917
Campylopusscabrophyllus
Suppl. 97. 1900. DicranumscabrophyllumC. Miil- (hb. Frahm).
ARGENTINA.Foz du Iguaqu,v. Hiibschmanns.n.
ler, Hedwigia37: 98. 1898. Type. Uruguay,Montevideo,Arechavaleta232 (holotype,destroyedat B, (hb. v. Hiibschmann);El MataderaNE of Cordoba,v.
Hiibschmanns.n. (hb. v. Hiibschmann).
lectotypusnov., NY).
Ber.
Thatigk.
(Hampe)
Jaeger,
subgriseus
Campylopus
Campylopus griseus belongs to a complex of
Dicranum subgriseumHampe, Vidensk. Meddel. species which are characterized morphologically
Naturhist.Foren.Kjoebenthavnser. 3, 4: 47. 1872. by hyaline, recurved leaf tips (not to be confused
Type. Brazil.Petropolis,Glaziou4543 (holotypenot with reflexed hairpoints). In South America this
seen, isotypes, H-BR, NY, PC).
complex is represented by C. laxoventralis HerPlants robust, on loose tufts, blackish below zog ex J.-P. Frahm, C. chilensis De Notaris and
and yellowish green at tips. Stems erect, to 10 C. recurvifolius Dusen, all from Chile. It can be
cm high. Leaves 7-8 mm long, with conspicu- supposed that C. griseus has been derived from
ously recurvedleaf tips, from ovate base grad- a southern temperate to subantarctic ancestor.
ually contractedto a long, fine tip. Costa excur- All species are closely related and cannot be disrent in a long, serrate, hyaline tip, lamellose at tinguished in the field. They differ mainly in the
back, with ventral and dorsal bands of stereids. thickness of the basal laminal cells and the transAlar cells inflated, hyaline or brownish. Basal verse section of the costa. Whereas all Chilean
laminal cells incrassate, elongate, rectangular, 10- species have thin-walled basal laminal cells, the
13 x 26-51 ,um, towards margins narrower, subtropical C. griseus ssp. griseus and the andean
forming a hyaline border. Upper laminal cells ssp. ingeniensis have firm, incrassate basal laminal cells, which function as protection against
oval, incrassate, 4-6 x 6-16 ,m.
Key to the Subspecies of

Campylopus griseus
Flora Neotropica
110
80
~~---.~
k._
I
--------??Lw
s
.
50
60
70
...
."__
0
0
FIG.7.
FIG. 73.
FIG. 73,
Disribtio
o/f
.
Campl-s
40
r--ieu
ss
Distribution of Camplopus grise
200
100
400
200
600
300
400
800 1000km
500
600
miles
p. g
() and C. grses
ssp. ngenenss
20
Distribution of Campylopus griseus ssp. griseus (0) and C. griseus ssp. ingeniensis (0).
shrinkingin open habitats with high evaporation. Campylopusgriseus ssp. griseus and ssp.
ingensiensisdifferonly by the pitted vs. not pitted upperlaminalcells and arethereforeregarded
as vicariantsubspeciesof the same species in the
Andes and in SE Brazil,which is a common disjunction in bryophytes.
This species varies in the color (more or less
blackish)and the size of the plants(robustin wet
grasslandand less robuston damp rocks).Small,
lightercolored plants have been describedas C.
lapidicola, a species, which was previously re-
duced to a variety of C. griseus. However, field

observations have revealed that there is no differentiation possible.
29b. Campylopus griseus ssp. ingeniensis (Williams) J.-P. Frahm, comb. nov. Figs. 73, 74.
CampylopusingeniensisWilliams,Bull.New YorkBot.
Gard. 3(9): 109. 1903. Type. Bolivia. Ingenio, Williams 1772 (holotype,NY; isotypes, B, FH, H-BR).
Plants to 8 cm high, blackish below, yellowish
at tips. Stems equally foliate with leaves con-
111
, . /
FIG. 74. Campylopus griseus ssp. ingeniensis (Campylopodes Peruvianae Exsiccatae 14). A. Plant. B. Leaf.
C. Leaf-tip. D. Transverse section of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for
50 ,um.
MAm.
microscopic details -= 50
spicuously recurvedat tips. Leaves from broad

ovate base graduallycontractedinto a long, slender tip, ending in a long, hyaline, serratehairpoint. Costa filling 1/3of the leaf base, with side
nerves, in transverse section with ventral substereids, ridged at back with lamellae one cell
high. Alar cells conspicuous,reddish,auriculate.
Basal laminal cells short rectangular,incrassate
and pitted, 8-13 x 32-58 Mm,in 2-3 rows elongate and narrowerat margins. Upper laminal
cells elongateoval, incrassate,stronglypitted, 56 x 35-45 um, 6-10:1.

Distribution(Fig. 73). On wet rocks and wet
soil in swampyarea,alongstreamsand in springs
in Peru and Bolivia, the few collections made
between 3300 and 3400 m.
Specimens examined. PERU. HuANuco: Road
Huinuco-Huaraz
betweenLa Unionand Huallanca,
112
Flora Neotropica
g51
4.5Im
|A
D!
?
a7
FIG. 75. Campylopusheterostachys(Croat45622, MO). A. Plant.B. Leaf.C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 um.
Frahm, Camp.Peruv.Exsicc. 14 (ALTA,B, BM, BUF,
DUKE, EGR, F, FLAS, G, GRO, GZU, HBG, KR,
NAM, NICH, NY, MEXU, NFLD, PC, POZ, RNG,
S, U); Huinuco Viejo, Frahm et al. P59 (B).
BOLIVIA. Cerrode Malaga,Herzog 4401 (H-BR,
JE);Tablas, Herzog 2945 (H-BR, JE).
30. Campylopus heterostachys (Hampe) Jaeger,
1870-1871: 421. 1872. Dicranum heterostachys Hampe, Flora 48: 581. 1865. Type. Peru.
Prov. Carabaya, Sandia, Hasskarl s.n. (holoFigs. 69, 75.
type, BM; isotype, S).
Campylopusalopecurus(C. Miiller)Kindberg,Enum.
Bryin. exot. 88. 1889. Dicranum alopecurum C.
Miller, Linnaea43:401. 1882.Type.Argentina."Rio
Secco inter Oran et S. Andres,"Lorentzs.n. (holotype, destroyedat B; isotypes, H-BR, JE, S).
Campylopusannotinus Mitten, J. Linn. Soc. Bot. 12:
80. 1869. Type. Colombia. "Andes Bogotenses,in
sylvis prope Pacho," Weir214 (lectotype,NY; isolectotype,H-BR).
Campylopushellerianus(Hampe)Jaeger,Ber.Thatigk.
Dicranum hellerianumHampe, Verh. K. K. Zool.
Bot. Ges. Wien 19: 507. 1870. Campylopushelleri
Hampe ex Kindberg,Enum. Bryin. exot. 50, 1888.
SystematicTreatment
113
Type.Mexico.Huatusco,Hellers.n. (isotypes,FH,
NY).
NearVolcanTajumulcoaboveEl Provenir,Sharp5368
(F);above SanRafaelPie de la Cuesta,Standley68613,
Campylopusporphyreocaulis(C. Miller) Kindberg, 68572 (F).
Enum.Bryin.exot.89. 1889.Dicranum
porphyreo- NICARAGUA. JINOTEGA: 6 km N of Santa Maria
cauleC. Miller,Linnaea42:472. 1879.Type.Ven- Ostuma, Crosby2746 (DUKE).
ezuela.Fendler34 (holotype,destroyedat B;lectoCOSTA RICA. CARTAGO:
Alajuela, Standley 41661,
typusnov., BM;isolectotypes,
MICH,NY, PC,S). 39629,47460,39793,41657, 43013, Valerio282 (FH);
Plantsin loose, yellow-greentuftsto 5 cm high,

densely whitish or reddish tomentose. Leaves
erect spreading,the upper part curled, 6-7 mm
long, long-lanceolate, subulate, ending in a
roughlydentate awn. Costa filling 1/2or more of
the leaf base, in transversesection showinglarge
ventral hyalocysts (occupying1/2of the section)
and groups of 4-7 very small dorsal stereids,
ridgedand serrateat back. Alar cells hyaline or
reddish, thin walled, intruding into the costa.
Basal laminal cells hyaline, thin-walled,rectangular, 6-13 x 42-67 Am, narrowerat margins.
Upper laminal cells short rectangular to
subquadrate,4-5 x 5-7 inm.
Seta 6-7 mm, sinuose, light brown. Capsule2
mm, slightlycurved and asymmetric,strumose,
light brown, furrowed.
Distribution (Fig. 69). On tree trunks and de-
caying wood in montane forests(1100-3200 m)

of Mexico, Guatemala, Honduras, Nicaragua,
CostaRica, throughthe Andes in Venezuela,Colombia, Ecuador,Peru,and Bolivia, in Paraguay
and NorthernArgentina,and also disjunctin SE
Brazil.
vic. Sta.Mariade Dota, Standley41661, 41657, 43016

(NY); SANJOSE:Valle de Copey, 30 km S of Cartago,
Griffin& MoralesB 95 (NY).
COLOMBIA. BOYACA:
CarreteraVilla de LeivaArcabuco,Cleef7527 (U); Caminode Soataal Alto de
Onzaga,v. d. Hammen & Jaramillo 1951, 2811 (U);
CarreteraRiquira-LaCandelaria,Cleefetal. 3466 (U).
CUNDINAMARCA:Sasaima, finca S. Jose, Cleef& v. d.
Hammen4958 (U). Sabanade Bogota,Schultes11068
Carretera Paramo de Guantiva(NY). SANTANDER:
Onzaga,v. d. Hammen & Jaramillo 1730, 1732 (U).

VENEZUELA. LARA:CarreteraBarbacoas-Hato
Arriba-El Tocuyo, Lopez & Smith 16709 (FLAS).
MERIDA: La Carboneraarea, Griffin17463 (FLAS);
Lopez-F.14061, Griffinet al. 1593, 1677, 2239 (FLAS,
NY). TACHIRA:Paramo La Negra, Griffinet al. 2218
(FLAS);compradoen le MercadoPrincipalde la ciudad de Merida, Ruiz-T. & L6pez-F., 8343 (FLAS);
Paramode Tama above the village of Paez, Griffinet
al. 951 (FLAS). Distr. Libertador,finca El Maciegal
alongLa Pedregosatributaryof the ChamaRiver, Griffin et al. 740, 750 (FLAS).
ECUADOR. LOJA: 48-50 km N of Loja, Luteyn
8050 (NY); 12 km S of YanganaS of Loja, Luteyn
8044 (NY). PICHINCHA:
Volcan Pichincha, Steere &
Balslev26011 (NY).
PERU. AMAZONAS:
km 417.5 above Leimebamba,Frahm, Camp. Peruv.

Exsicc. 11 (ALTA, B, BM, BUF, DUKE, EGR, F,
FLAS,G, GRO, GZU, HBG, KR, NAM, NICH, NY,
MEXU, NFLD, PC, POZ, RNG, S, U), Frahmet al.
Alrede- 368 (B); km 403.5, Frahm et al. 787 (B); Saullamur
Specimensexamined.MEXICO. CHLAPAS:
doresFincaLiquidambarcercade Jaltenangode la Paz, nearBalsas,Hegewald6698 (hb.Hegewald).AYACUCHO:
Delgadillo 3570 (MEXU); Tapachula,Benito Juarez, Along road Huanta-San Francisco 15 km W of San
CerroTresPicos,Eggers&Frahm792033 (hb.Frahm). Francisco, Frahm 823945 (hb. Frahm). CAJAMARCA:
DURANGO:
Along Hwy 40 about 4 miles W of La Ciu- Cutero, Raimondi 3827 (B). PIURA:Palambla near
Between
dad, Bowers et al. 5256 (TENN). GUERRERO:
Canchaque,Lowys.n. (LAF).
Mipillasand Atayade Alvarez,Croat45622 (MO).
BRAZIL.SAOPAULO:
Camposde Jordao,BookerSierraMadre Occidental,Real Alto, Mexia man 5609 (SP).
JALISCO:
Mun.
SierradeChichinautzin,
1595b(NY).MORELOS:
BOLIVIA.Vic. Yungasand La Paz, Jay s.n. (NY).
Be- Florida de San Mateo, Herzog 3629 (B, JE, NY).
de Tepoztlan, Cardenas1058 (MEXU). PUEBLA:
low Huauchinango toward Nexaca, Sharp 3080 Bachschluchtdes Cerro Amboro, Herzog 267 (JE).
(TENN); alrededoresde Xicotepec de Juarez,Delga- TARIJA:Rio Bermejo65 km NW of Pozos Bermejo,
Above Arnow41138 (COLO);entre EntreRios et Tarija,De
dillo 1246, 1385 (MEXU). SANLUISPOTOSi:
Rancho
Xilitla, Sharp 5904a (TENN). TAMAULIPAS:
Sloover75 (hb. Frahm).
del Cielo above Gomez Farias,Delgadillo1408, 2701
PARAGUAY.Withoutlocality, Grosses.n. (MO).
(MEXU), Iwatsuki& Sharp 4756, Sharp 3653, 8652,
La Patronata,Sharp 8961
8772 (TENN). VERACRUZ:
This species has often been confused with C.
(TENN);entreSantaRitay Yecuatla,Delgadillo1054 flexuosus but is easily distinguished by the thin(MEXU); vic Coatepec, Gil 158, 275, 387, 382, 95 walled
hyaline basal laminal cells, the alar cells
(TENN); Banderilla5 km al N de Xalapa, Vienox&
nearYecuatla, occupying part of the costa, the usually strongly
Guzmdn570b(TENN);LosHuerfanos
serrate leaf tip and the large ventral hyalocysts
Sharpet al. 3077a (TENN).
La Montanita,Standley of the costa, which are conspicuous even in venHONDURAS. MORAZAN:
12357 (F).
tral view without transverse section. The leaf tip
BetweenTodos
GUATEMALA.HUEHUETENANGO:
Santos and San Martin, Sharp 4808 (F). SANJOSE: varies somewhat as also less serrate forms have
STANDLEY
68592, 41661, 43016 (FH). SAN MARCOS: been observed.
114
Flora Neotropica
Bartramin his moss floraof Guatemala(1949)

included this species and also C. tallulensisand
C. zygodonticarpusin C. flexuosus.
Sporophytesareproducedextremelyrarelyand
have been found only once, on Griffin17463
from Venezuela.
Jaramillo
DomingodelosColorados,
78?48'W,
0?13'S,
& Zak 8156, 7949A (NY).

BOLIVIA.LAPAZ:AlongRio Zongoat Cahuapower station, 16?2'S,67?59'W,Lewis 79-1746 (F).
Describedfrom the East slope of the Andes of

Peru,this species was known only from the type
locality for 70 years, probablydue to the insufficientexplorationof the andinemontaneforests.
31. Campylopushuallagensis Brotherus, HedIdentifications of recent collections have rewigia 45: 261. 1906.
vealed this species to be scatteredthrough the
Andes from Costa Rica to Bolivia.
Campylopushuallagensisis closely relatedto
Campylopus huallagensis
C. weberbaueri,which occurs in the same habitats and about the same range. Both taxa were
1 Upperlaminalcellselongateoval. ...........
31a. var. huallagensis. describedby BrotherusfromNE Peru,fromabout
........................
200 km distant localities. Both species can be
1 Upperlaminalcellssubquadrate.
.............
........................
32b. var. weberbaueri. distinguishedonly by the shapeof the upperlaminal cells and thereforethey are united here as
31a. Campylopus huallagensis Brotherus var. varieties of one species.
huallagensis
Figs. 71, 76.
31b. Campylopushuallagensisvar. weberbaueri
CampylopushuallagensisBrotherus,Hedwigia45: 261.
1906.Type.Peru.RioHuallaga,
Ule2358(holotype, (Brotherus)J.-P. Frahm comb. et stat. nov.
Figs. 71, 77.
H-BR;isotypes,JE,FH).
weberbaueri
Plants to 5 cm high, robust, dark brown or Campylopus
Brotherus,Bot. Jahrb.56,
Beibl.123:5. 1920.Type.Peru.Loreto:E of Moyoblackish below, yellowish green at tips. Stems
4718 (holotype,H-BR).
bamba,Weberbauer
interruptedlycomose foliate. Leaves ca. 10 mm Campylopus
wurdackii
70: 319.
Robinson,Bryologist
1967.Type.Peru.Amazonas:
long, narrowlylanceolate, graduallycontracted
RioVentilla,Wurdack
1524(holotype,US).
to a serrateapex. Costaexcurrent,filling1/3of the
leaf width, in transversesection with ventraland
Plants to 10 cm, in loose, dark green tufts.
dorsalbandsofstereids.Alarcells reddishbrown,
Stems interruptedlyverticillate foliate. Leaves
conspicuous. Basal laminal cells incrassate, 10-11 mm
long, narrow lanceolate, gradually
stronglypitted, longly rectangular,8-13 x 32- contractedto a
long, finely serratepoint, serrate
48 Aum.
Upper laminal cells incrassate,elongate in the upper third of the leaf. Costa
filling /3 of
oval, 6-10 x 20-42 ,m, ca. 4-6:1.
the leaf base, fillingthe leaf tip or shortlyexcurSporophyte not known.
rent, in transversesection with ventral stereids
Distribution(Fig. 71). On soil, rocks,also epi- and dorsal
groups of stereids. Alar cells differphyticon branchesin moist montanerainforests entiated. Basal laminal cells
rectangular,incrasbetween 1400 and 2300 m elevation, mostly in
5-10 x 13-29 um. Upper
sate,
strongly
pitted,
cloud forestsbetween 1800 and 2200 m in Costa
laminal cells short rectangularoblique, 3-10 x
Rica, Colombia, Venezuela, Ecuador,Peru and 5-12
Aum,reachingas a narrow band nearly to
Bolivia.
the leaf tip.
Specimens examined. COSTA RICA. ALAJUELA:
La
Sporophytesfrom pseudolateralshortbranchPalma de San Ram6n, Brenes 16216 (NY).
es. Seta 7 mm long, light brown. Capsule2 mm
COLOMBIA.HUILA:
ParqueNacional de los Guacharos y CerroPunta, Cleef 5088, 5058, 5047, 5068, long, obliquelyovoid, brownish,smooth, slightly
furrowedwhen emptied. Calpytraciliate at base.
5037 (U). VAUPES:Rio Kananari, affluent of Rio ApaDistribution (Fig. 71). Epiphytic, usually on
pori, Schultes& Cabrera13413 (FLAS).
VENEZUELA. BOLIVAR:
Chimanta massif, Stey- branchesof small treesand shrubsin elfin forests
ermark74980 (NY).
in Colombia and Peru.
ECUADOR. MORONA-SANTIAGO:
General Plaza-
Gualeceoroad km 18-20, 78?30'W,3?0'S,Lojtnant&
Molau 14626 (AAU, GB). PICHINCHA:

Reserva Floris-
tica Rio Guajalitokm 59 carreteraantigua Quito-S.
Specimens examined. COLOMBIA. MAGDALENA:
ParqueNacionalSierraNevadadeSantaMarta,Griffn
et al. 500088 (FLAS).
115
IPKr
I'
FIG. 76. Campylopushuallagensisvar. huallagensis(Cleef 5088, U). A. Plant. B. Leaf. C. Leaf-tip. D.

details = 50 Am.
PERU. AMAZONAS:
Road Chacapoyas-Cajamarca relatively young mountain systems can be rekm 417, Frahm et al. 1979 (B). SAN MARTIN:
Road garded as the youngest evolutionary branch.
Chachapoyas-Moyobambakm 387, Frahm et al. 343 Campylopus
huallagensisvar. weberbauerimuch
(B). Pancarlanto,Jay s.n. (NY).
This species is conspicuous by its verticillate
foliation, its size and the length of the leaves.
The occurrence on branches is met with very
rarely in Campylopus. Compared with the probable Mesozoic age of this genus and an origin in
subantarctic regions, such epiphytic species in
resemblesC. lamellinervisin size, leaf shape,serrate leaf borderand areolation.The only differences seem to be the shortrectangularoutermarginal basal laminal cells in C. huallagensisvar.
weberbaueri(vs. elongatein C. lamellinervis),the
dorsal side of the costa, which shows lamellae 1
cell high in C. huallagensisvar. weberbaueribut
2-3 cells high in C. lamellinervis,and the typical
116
Flora Neotropica
?n_gg
Cap p
a.
FIG. 77. Campylopushuallagensisvar. weberbaueri(Griffinet al. 500088, FLAS).A. Plant. B. Leaf. C. Leaftip. D. Transversesection of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for
microscopicdetails = 50 Am.
verticillate foliation of C. huallagensisvar. weberbaueri.Therefore,the latter may have been

derived from C. lamellinervis,which may have
become epiphyticunder the changingecological
conditions in the Tertiary,when the Andes were
upfolded. The loss of the lamellae can be interpreted as caused by the absence of a need for
waterstoragein elfin-forests(C. lamellinervisoccurs in dry forests)and the verticillate foliation
can be interpretedas an adaptation to a better

gas exchangeunderthe perhumidrainforestconditions.
32. Campylopusincertus Theriot, Rev. Bryol.
Lichenol.11:63.1939. Type. Colombia.Paramo El Boqueronnear Bogota, Troll2161 (lectotype nov., PC; isotypes, FH, JE).
Figs. 78, 79.
117
A
FIG. 78. Campylopusincertus(Cleef1607, U). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of leaf.
E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Am.
Plants to 5 cm high, light green,in loose tufts. showinga gradualtransitionto the elongateoval
Stems loosely and equally foliate with erect (6:1) upper laminal cells, 4-6 x 22-38 Asm.
spreadingleaves. Microphyllousbranchesoften
Distribution (Fig. 79). On soil and and wet
present in the axils of the upper leaves. Leaves
ca. 6 mm long, from ovate base graduallycon- rocks in paramos (3300-3700 m, rarely up to
tracted to a finely serrateleaf tip. Costa excur- 4400 m, also in open habitatsas roadsidebanks
rent, filling 1/2of the leaf base, in transversesec- in montanerainforestsat lowerelevations(1900tion with ventral hyalocystsand dorsal stereids. 3200 m) in Venezuela, Colombia and Peru.
Alar cells large, protrudinginto the costa. Basal
Specimensexamined:COLOMBIA.BOYACA:Prialaminalcells rectangular,incrassate,6-13 x 35- mo de la SarnaentreSogamosoy VadoHondo,
Cleef
70 Mm,becoming shorter towards the leaf tip, 9512(U);SierraNevadadel Cocuy,Paramodel Con-
Flora Neotropica
118
80
70
60
50
'
200
_
.^fc/--'
<\
\^^
o0
FIG 7.
?^>
100
oPrepared
400
200
600
300
400
800
1000km
500
600
miles
10
Dstibuio
FIG. 79. Distributionof Campylopusincertus.
cavo, Cleef8593 (U); Pefiade Arical N de Vado Hondo, Cleef 9474 (U); Paramo al NW de Belen, Cleef
2138 (U); CarreteraSoyamoso-Pajarito,13 km al NW
de Vado Hondo, Cleef9550, 9546 (U). CUNDINAMAR-
CA:ParqueNacional Chingaza,Smeets B12 (U); Paramo de Sumapaz,Cleef 1607 (U); Paramode Palacio,
Cleef& v. d. Hammen 4971 (U); Lagunasde Buitrago,
Cleef 9581 (U); Paramoentre Coguay San Cayetano,
Sierra Nevada de Santa
Cleef4900b (U). MAGDALENA:
Marta,Alto Buritica,v. Reenen & Rangel 394 (U).
VENEZUELA. MERIDA:Carretera entre El Morro
y Aricagua,Ruiz-T. & Lopez-F. 9546 (FLAS).TACHIRA: Paramode Tama above the village of Paez, Griffin
et al. 129 (FLAS).
PERU. AMAZONAS;
between Balsasand Leimebamba,Frahm, Camp. Peruv. Exsicc. 13 (ALTA, B, BM, BUF, DUKE, EGR,
F, FLAS, G, GRO, GZU, HBG, KR, NAM, NICH,
NY, MEXU, NFLD, PC, POZ, RNG, S, U).
Campylopus incertus is conspicuous by the
presence of microphyllous branches in the axils
of the upper leaves, which are similar to those
of C. areodictyon. Such microphyllous branches
are also present in the type specimen, but were
not mentioned in the type description. Also, as
in C. areodictyon, these branches are frequently
distichous foliate in the upper part.
By the elongate oval upper laminal cells, which
show gradual transitions to the rectangular basal

laminal cells and the alar cells protruding into
the costa, this species resembles C. asperifolius.
Since the latter species occurs as epiphyte in
montane rainforests, and C. incertus is a paramo
species, both may be not only closely related but
of the same origin and can be regarded as an
ecologically differentiated species pair. Such altitudinal differentiation is found also in C. albidovirens and C. pittieri, C. flexuosus vars. flexuosus and incacorralis, C. cuspidatus vars.
cuspidatus and dicnemioides, and seems to be a
speciation caused by the Andean uplift.
33. Campylopus jamesonii (Hooker) Jaeger, Ber.
Thatigk. St. Gallischen Naturwiss. Ges. 18701871: 422.1872. Dicranumjamesonii Hooker,
P1.rar. 2: 179. 1837. Type. Ecuador. "in monte
Surruchcho," Jameson s.n. (not seen).
Figs. 80, 81.
Campylopusaltissimus (C. Muller)Jaeger,Ber. Thatigk.St. GallischenNaturwiss.Ges. 1877-1878: 381.
1880.DicranumaltissimumC. Miller, Flora58: 530.
1875. Type. Colombia. Antioquia: Walliss.n. (holotype, destroyedat B; isotypes, NY, S).
119
Inmm
FIG. 80. Campylopusjamesonii (Griffinet al. 946, FLAS). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
50 Mm.
CampylopusstandleyiBartram,Contr.U.S. Natl. Mus.

26: 57. 1928. Type. CostaRica. Cerrode las Vueltas,
Standley & Valerio43939 (holotype, FH; isotypes,
JE, NY).
Plants large, Dicranum-like, to 10 cm high, in
shiny, light-green mats. Stems equally and loosely foliate with patent leaves which are crisped
when dry. Leaves 1 cm long, from decurrent base
gradually long- and fine-acuminate, serrate at
margins in the upper third. Costa broad, filling
of the leaf width, in transverse section with

large ventral hyalocysts and dorsal groups of
stereids,smooth at back.Alarcells reddishbrown
or rarelyhyaline, decurrentand protrudinginto
the costa. Basal laminal cells thin-walled, hyaline, 10-16 x 80-96 Am, rectangular.Upper
laminalcells shortrectangular,small and opaque,
6-10 x 6-13 /tm, arrangedin distinct rows.
Sporophytenot seen.
Distribution(Fig. 81). On soil, litter and, rare2/3
120
Flora Neotropica
100
110
90
70
s0
h,
81.
Distribution
of
------------
Campylopus
1;0
2O 3;0
by
4X*
S0
6 O -1
-l
--
--
--
- - - ---- - - - - -
-t------
5~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
( ) and
C. japonicus
(O).
O~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
Preo~d
50
"B~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~3
01
FIG.
jameson'i
60
-----
-----
--
endh R. Rpkem
J!
FI.
Dstiuto
o Cmylpsjmeod
0)ad
apniu
()
ly, rotten trunks in subalpine forests and shrubs

between 2700 and 3700 m, rarely below (as 1200
m in Costa Rica) through the Andes from Mexico
and Guatemala to Costa Rica, Venezuela, Colombia, Ecuador, Peru, Bolivia to northern Argentina, disjunct in SE Brazil. Also in corresponding habitats and elevations in Africa
(Cameroun, Tanzania, Zaire, Uganda, Rwanda,
Kenia, Reunion, Malagasy, Moqambique, S
Rhodesia, Malawi, Transvaal), where this species has been described and recorded as C. procerus (C. Miill.) Par. and C. echernieri Besch.
s.n. (FLAS);Refugiode vida silvestreTapanti,Arrocha

CA 998, 1177 (hb. Frahm).
VolcanPurace,Lagunade San
COLOMBIA.CAUCA:
Rafael,Cleef&Fernandez579 (U); ParamoLasPapas,
Barclay 5988 (MICH). BOYACA:CarreteraChiquinquira-Pauna, v. d. Hammen & Jaramillo 2876 (U);
Paramo de La Rusia N-NW de Duitama, Cleef 7341
(U); Paramode Pisva, Cleef4618 (U); Paramoal NW
Pass above
Llano de las Flores between Ixtlan de Juarez and Tuxtepec, Sharp et al. 2371d (TENN); N of Oaxaca on
Hwy. 175 at Sierra Juarez Gap, Sharp et al. 4772d
(TENN).
5293 (U). SANTANDER:

Valle de Susa arriba de Santa
de Belen, Cleef1820 (U). CUNDINAMARCA:

Bogota, Re-
gi6n de Monserrate,Zuluaga 188 (U); Subachoque,

CuchillaEl Tablazo,Linares& Bulla 875 (COL);Cordillerade Leonora60 km NNE de Bogota, v. d. Hammen & Jaramillo 3145 (U); Piramo de Palacio, Cleef
5138, Cleef&Jaramillo4058 (U); SanCayetano,Hda.
Portugal,Cleef6571 (U); Paramode Guerrero,Schultes
s.n. (NY);META:
Paramode Sumapaz,Cleef806, 5292,
Barbera,v. d. Hammen & Jaramillo 2881 (U).

VENEZUELA. ARAGUA:
Colonia T6var-La Victoria, 10?22'N,67?20'W,Luteynet al. 8278 (NY). MERIDA:ParqueNacional SierraNevada, la Mucuy,FranCOSTA RICA. SANJOSE:Cerrode la Muerte3 mi sen 1344 (GB);between La Aguadaand La Montafia,
NW Villa Mills, Koch 5090 (NY); 3 mi N of El Em- Fransen 1457 (GB);Campo Elias,Road to La Azulita,
palme,Croat32887B (MO);97 km S ofCartago,Griffin Fransen1238 (GB);Montafiade S. RafaelentreS. Cruz
de Moray El Molino,Ruiz-T. 8413b (FLAS).TACHIRA:erous and evergreenforests between 1500 and
et 2400 m altitude.
Paramode Tami abovethe villageof Paez,Griffin
El RosalbetweenLa
al. 313, 946 (FLAS);Pairamo
et al. 739(FLAS). Specimensexamined.MEXICO.CHIAPAS:
GritaandS. Josede Bolivar,Griffin
Mun. de
TRUILLO:
EntreLlanodel Tigrey Tres Pozos, Riuz-T. Tapachula, Benito Juarez, Volcan Tacana, Frahm,
8425 (FLAS).
Camp. Exsicc. 3 (B, BING, C, DUIS, DUKE, EGR,
BetweenLoja and Cuenca, F, FLAS,G, GOET,GRO, GZU, H, HBG, JE, KRA,
ECUADOR.AZUAY:
Steere25943(NY).Loja:betweenLojaandSaraguro, M, MEXU, MO, NAM, NFLD, NICH, NY, PRC,
Ca. 21 km PRE, POZ, RNG, S, U, UPS).
Steere & Balslev25913 (NY); PICHINCHA:
E of Pifo on roadto Baeza,0?23'S,78?13'W,Churchill
JAuSco:AlongGuadalajara-Tepic
road,Koelz34032
& Sastre-DeJesus 13525 (NY). ZAMORA-CHINCHIPE:
(MICH).
79?W, 4?02'S, Holm-Nielsen et al. 3822 (GB); road
Loja-Zamorakm 24-25, Holm-Nielsenet al. 3497 (GB).
A specieswhichis conspicuousby its subquadCalla Calla near Leimebamba,
PERU. AMAZONAS:
rate basal laminal cells. The remarkabledis-
aboveLeimebamba,
Hegewald6895 (hb.Hegewald);
Philippi2341(B).Cuzco:HuaynaPicchu,Urubamba junctionbetweenAsia and Mexico is met in sevvalley,MenzelP 298 (B);MachuPicchu,Frahm823962 eralotherspeciesofbryophytesand is interpreted
(hb. Frahm).
as a relicof a circum-pacificrangein the Tertiary.
Parque Nacional Itatiaia,
The specimensfromMexicodifferfromthe Asian
AgulhasNegras2700 m, Frahm1412 (hb. Frahm),
in the transversesectionof the costa,
populations
Frahm, Camp. Bras. Exsicc. 24 (ALTA, B, C, DUIS,
FLAS,GOET,GZU,H, HBG,INPA,JE,KR,M,MO, which show ventral stereids throughthe whole
NFLD,NICH,NY, PC,S, SP, TENN,U), Griffin& leaf, whereasthe Asian specimens show ventral
Vital 167 (FLAS).
substereidsin the basalpartof the leaf and stereids in the upperpartof the leaf.This mayindicate
34. CampylopusjaponicusBrotherus,Hedwigia that the Mexicanpopulationmay be a vicariant
38: 207. 1899. Type. Japan,without detailed geographicalrace.
locality, Ankarcronas.n. (holotype, H-BR).
Campylopusjaponicusis knownonly sterilein
Figs. 81, 82. Mexico, which may be the reason for the small
and isolated range.
Campylopussaint-pierreiTheriot, SmithsonianMisc.
El
Mexico.
1931.
3.
Collect.85(4):
Hidalgo,
Type.
Chico, Saint Pierre 1589 (holotype,PC).
35. Campylopusjugorum Herzog, Beih. Bot.
Centralbl.26(2): 51. 1910. Type. Bolivia. CoPlants in loose, green tufts to 3.5 cm high,
AbraSanBenito,Herzog3356 (hochabamba:
reddish tomentose below, equally foliate or
Figs. 83, 84.
JE;
isotype, B).
lotype,
Leaves
sometimes interruptedlycomose.
loosely
a
from
acuminate
mm
5-7
erect,
long, slenderly
Plantsin compact,lightto yellowishgreentufts,
lanceolatebase and (in sunny places) ending in to 5 cm
high. Stems equally foliate with apa short or long, hyaline, toothed awn, tubulose
Leaves ca. 6 mm long, narrowly
leaves.
in the upper1/3.Costa 1/2or more of the leaf base, pressed
narrowed into a long,
lanceolate,
gradually
slightlyribbedat back,in transversesectionwith smooth subula. Costa filling 3/4of the leaf-base,
ventral substereidsin the basal part of the leaf
excurrent,in transversesection with ventralhyand ventral stereidsin the upperpart. Alar cells
alocystsand a bandof dorsalsubstereids,smooth
conspicuouslydifferentiated,reddish or hyaline at back. Alar cells lackingor poorly developed.
(sometimes reddish within and hyaline at the Basal laminal cells rectangular,thin-walled, 6margins).Innerbasal laminal cells thick-walled, 12 x 29-58 jm, at marginsin 7-9 rows narrowsubquadrateto short rectangular(1-3:1), the er, elongate, forming a hyaline border. Upper
marginalcells longer,narrowerand hyalinein 3- laminalcells elongateoval, incrassate,3-6 x 167 rows. Upper laminal cells oval-elongate,with 22
2tm, ca. 6-8:1.
rounded ends owing to thickened covers, 13Sporophyteas in C. nivalis.
26 x 5-7 gm, 3-6:1, those at marginsshorter.
Distribution(Fig. 84). On rocks in the high
Sporophytes unknown.
belt of the Andes between 3900 and 4400
alpine
Distribution(Fig. 81). An Asian species rang- m elevation in
Colombia, Peru and Bolivia.
ing from Japanto S China,PeninsularMalaysia,
Sierra
Tahiti and Queensland.In the Neotropicsfound
only in Mexico on soil and rocks in wet conif-
Nevada del Cocuy,ParamoConcavo, Cleef8696 (U).
122
Flora Neotropica
ai
,~
l
09C)-
OOOOoOOO o
oooo
....)
00
FIG. 82. Campylopusjaponicus(CampylopodesExsiccatae3). A. Plant. B. Leaf.C. Leaf-tip.D. Transverse

50 Am.
PERU.JUNI:LagunaChaulacocha
nearLaOroya, smaller ventral hyalocysts in transversesection
Hegewald5440 (hb.Frahm).
of the costa and longer upper laminal cells.
BOLIVIA.Tunarisee,
Herzog4902(JE).
Campylopusjugorumresembles C. edithae in 36. Campylopusjulaceus Jaeger, Ber. Thatigk.
the elongateoval upperlaminalcells and growth
St. GallischenNaturwiss.Ges. 1877-1878:384.
in dense tufts in high alpine habitats,but is dis1880. Dicranumjulaceum Hampe, Viddensk
Meddel. Naturhist. Foren. Kjoebenhavn ser.
tinguished by the lack of a hyaline hairpoint,
123
"
"
F
.'
8cm
4mmn
.'
?-^B
FIG. 83. Campylopusjugorum(Cleef8696, U). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of leaf.

E. Basal laminal cells. F. Upper laminal cells. G(H). Capsule(s).Unlabeled scale bar for microscopicdetails =
50 tm.
Plants in loose tufts, yellowishgreen.Stems to

6 cm high, filiform, with densely appressedfoliate leaves. Fertile plants ending in nearly globose perichaetia. Leaves 4-6 mm long, entire,
lanceolate,endingin a shorthyalinetip, the stem
(C. Miller) Paris,Ind.bryol. leaves narrowlylanceolate,the perichaetialleaves
Campylopusmicrojulaceus
Suppl.94. 1900. Dicranummicro-julaceumC. Miillanceolateto ovate. Costa filling1/3of the
ler, Bull. Herb. BoissierVI: 37. 1898. Type. Brazil. broadly
Serrado Itatiaia, Ule 1796 (holotype, destroyedat leaf width, excurrent,in transversesection with
ventral hyalocystsand dorsal groupsof stereids,
B; isotype, H-BR).
3, 4: 45. 1872. hom. illeg. Type. Brazil. Rio
de Janeiro: Serra dos Orgaos, Glaziou 4554
(holotype, n.v.; isotypes, H-BR, PC).
Figs. 84, 85.
124
Flora Neotropica
.0
200
400
600
800 1000km
0 100 200 300 400 500 600 miles

10
10
FIG. 84. Distributionof Campylopusjugorum() and C. julaceus (0).
lamellose at back. Alar cells not or scarcelydifferentiated,hyaline or brownish. Basal laminal

cells rectangular,hyaline, thin-walled, 6-12 x
25-70 ,um,in several rows narrowerat margins.
Upper laminal cells oval, 5-10 x 10-16 gm,
incrassate.
Sporophytenot known (?).
Distribution(Fig. 84). On open sandy or gravelly soil along roadsides, rarely on sandstone
rocks,from SEBrazilto Paraguay,Argentinaand
Bolivia, in elevations from 300 m to the forest

line.
Specimens examined. BRAZIL. BAHIA:4 km E of
Morrodo ChapeualongroadBA 052, Vital6045, 6046

(SP). MINASGERAIS:
ParqueNacional Itatiaia, Brejo
do Lapa, Vital4927 (FLAS,SP). PARANA:
Along road
fromGuarapuavaand Prudent6polis,Frahm 1572 (hb.
Frahm); Serra da Esperanqa25?22'S, 51?18'W, Vitt
21383 (ALTA). Rio DEJANEIRO:Serra do Itatiaia, Dusen s.n. (H-BR). SANTACATARINA:Km 213.5 along
road BR 135, Frahm 1579 (hb. Frahm);N of Curiti-
125
-i
C,C/7~
iF
::^-^
:''
FIG. 85. Campylopusjulaceus(Glaziou4554, H-BR). A. Plat. B. Leaf C. Leaf-tip.D. Transversesection
of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details = 50 cm.
banos 26?46'S,50?15'W,Vitt21017 (ALTA);km 198

alongHwy.BR 116betweenCuritibaandLages,Frahm,
GOET,GZU, H, HBG, INPA,JE,KR, M, MO,NFLD,
NICH, NY, PC, S, SP, TENN, U); km 213.5 along
Hwy. BR 116, Frahm 1579(hb. Frahm);km 142 along
Hwy. BR 282 between Varegio and Faxinal, Frahm,
GOET,GZU, H, HBG, INPA,JE,KR, M, MO,NFLD,
NICH, NY, PC, S, SP, TENN, U).
BOLIVIA. Eastern Cordillera,QuebradaCunucu,
Herzog 274 (JE, B).
PARAGUAY.
AMAMBAY:
Parque Nacional Cerro
Cora,trailup to CerroMuralla,22?40'S,56?00'W,Buck
12480 (NY).
ARGENTINA.W of Cordoba,v. Hiibschmanns.n.
(hb. v. Hiibschmann).
This species much resembles C. introflexus anatomically and is perhaps derived from the latter
species, which is widespread through the temperate and subantarctic regions of the Southern
Hemisphere. It differs only by the non-reflexed
Flora Neotropica
126
hairpoints, the appressedfoliate stems and the
blunt perichaetialleaves.
In Africa this sepcies occurs as ssp. arbogastii
(Renauld & Cardot)J.-P. Frahm, which differs
only by shorterlamellaeat the back of the costa.
This subspecies occurs in South Africa, Malagasy, Reunion and the Seychelles,conspicuously
at about the same latitude as ssp. julaceus, also
in the SE part of this continent and even in the
same habitats(however, down to sea level).
Campylopusjulaceus has a noticeably broad
altitudinalrange. Whereasit has been found in
SE Brazilnot below 700 m, it growsdown to 300
m in Paraguayand again up to 3500 m in the
Andes of Argentina.The locality in Bolivia is at
700 m, which might be the reason for the continuous range, connecting the Andes with SE
Brazil.
Only male plants,producingconspicuousbudlike perichaetia,have been found.
(hb.Frahm);km 181 alongHwy. BR 116 fromCuritiba

to Lages,Frahm 1578 (hb. Frahm);Serrado Espigao
17 km N of Dr. Pedrinho,Frahm 1895 (hb. Frahm).
SXo PAULO:
Serrado BocainaNW de Camposda Cunha, Frahm 1584 (hb. Frahm).
38. Campylopuslamellinervis(C. Miiller) Mitten, J. Linn. Soc. Bot. 12: 82. 1869.
Campylopus lamellinervis
1 Leaves tortuosewhen dry, recurved.Plants light
38a. var. lamellinervis.
green. ..................
1 Leavesstraightwhendry,erectpatent.Plantsdark
38b. var. exaltatus.
green. .....................
38a. Campylopuslamellinervis(C. Miller) Mitten var. lamellinervis

Figs. 88, 89.
Campylopuslamellinervis(C. Miller) Mitten,J. Linn.

Soc. Bot. 12: 82. 1869. Dicranum lamellinerveC.
Miiller, Syn. musc. frond. 1: 390. 1848. Type. Jamaica. Wilson s.n. (holotype, destroyed at B; iso37. CampylopusjulicaulisBrotherus,Denkschr.
types could not be located).
Akad.Wiss.Wienmath.-nat.K1.83:261.1926. Campylopusgiganteus Sullivant, Proc. Amer. Acad.
ArtsSci. 5:278. 1868.Type.Cuba. Wrights.n. (n.v.).
Type. Brazil. Parana:Roca Nova, Dusen 245
CampylopusmultisulcatusDuby in Hampe, Vidensk.
(holotype, H-BR; isotypes, FH, JE, NY).
272. 1870, nom. inval., cf. Index Muscorum.
Figs. 86, 87.
Campylopuspenicillatus (Horschuch) Jaeger, Ber.
Plants slender,in loose tufts,lightgreen.Stems
Thatigk.St. GallischenNaturwiss.Ges. 1870-1871:
442. 1872, hom. illeg.DicranumpenicillatumHornerect, to 4 cm high, densely reddish tomentose,
schuch, Fl. bras. 1(2): 13. 1840. Type. Brazil.Serra
equally foliate giving the plants a filiform apdos Orgaos,Gardner25 (holotype, destroyedat B;
pearance. Leaves appressed to the stem when
lectotypusnov., B; isolectotypes,GOET,JE).
erect
when
5
mm
lanceowet,
dry,
patent
long,
CampylopuspseudodicranumHerzog,Beih. Bot. Cenlate, concave, with canaliculatetips, serrate at
tralbl.26(2):53. 1910.Type.Bolivia.CerroAmboro,
tips. Costa filling2/3of the leaf width, excurrent,
Herzog s.n. (holotype,JE; isotypes, B, H-BR).
in transversesection with ventralhyalocystsand Campylopussubpenicillatus(C. Miller) Brotherusin
a ventral band of stereids, ridged at back. Alar
DicranumsubpenicillatumC. Muller,Hedwigia39:
cells hyaline or brownish, thin-walled. Basal
257. 1900. Type. Brazil. Sta. Catarina:Blumenau,
laminal cells rectangular,incrassate,6-16 x 26Ule s.n. (holotype,destroyedat B; isotype, H-BR).
54 um, narrowerat margins.Upper laminalcells Campylopustortuosus(Hampe)Paris,Ind.bryol.Suppl.
98. 1900. DicranumtortuosumHampe,Linnaea25:
small, subquadrate,incrassate, 6-10 x 10-16
361. 1853. Type. PuertoRico (n.v.).
Gim.
Distribution(Fig. 87). For 50 yearsthis species
was known only from the type localityin Parana.
Recently,however,it has beencollectedthroughout SE Brazil, where it occurs on rotten wood,
stems of fern trees and rocks between 300 and
2200 m.
Specimens examined. BRAZIL. BAHIA:Ca. 26 km
N of Seabra,road to Aguade Rega, Irwinet al. 30855
(NY). Rio DE JANEIRO:Itatiaia Nat. Park, Griffin &

Blumenau 7 km
Vital 245 (FLAS). SANTACATARINA:
NW of Benedito Novo, Mun. de Timb6, Frahm 1583
Plants large, to 10 cm high, Dicranum-like, in

loose, yellowish green tufts. Stems erect, unbranched, tomentose, equally foliate with tortuose, recurved leaves. Leaves ca. 1 cm long,
from ovate base graduallycontractedinto a long

subula,stronglyserratein the upperthird. Costa
filling
1/3-V2 of
the leaf base, excurrent, in trans-
verse section with ventral and dorsal rows of

stereids, ridgedat back in the basal part, lamellose at back in the upperpart,the lamellaebeing
serrate.Alarcells large,inflated,thin-walled,reddish-brown. Basal laminal cells longly rectan-
127
FIG. 86. Campylopusjulicaulis (Dusn 245, H-BR). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of
leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 ,um.
gular, incrassateand pitted, at margins shorter

(2-3:1), not pitted. Upper laminalcells subquadrate, incrassate.
Setae in bundlesof 3-4 arisingfrom the tip of
the stems, ca. 9 mm long, brownish. Capsules
curved, inclined, brownish, 2 mm long. Operculum 1.5 mm long, darkercoloured, obliquely
rostrate.
Distribution(Fig. 89). On sandy or loose mineral soil in shaded places in dry forests, such as
restingas and campos cerrados, also on grassy
slopes from sea level to the forest line, lacking
in rain forests regions, in Puerto Rico, Jamaica,

Haiti, Cuba, Venezuela, Colombia, Ecuador,
Peru, Bolivia and Brazil. In SE Brazil frequent,
elsewhere rare.
Specimens examined. CUBA. ORENTE:Sierra Nipe,
near Woodfred,Shafer3337 (PC).

GUYANA.UpperMazaruniRiver,5?45'N,60?35'W,
Boom & Gopaul7620 (NY).
ECUADOR. 8 km W Cuchumani,Hinz s.n. (hb.
Frahm).
PERU. Cuzco: Urubamba, Vagas2700 (MICH).
Km 241 alongroadBA-052, Mun.
BRAZIL.BAHIA:
de Morrodo Chapeu, Vital6026 (SP).MINASGERAS:
128
Flora Neotropica
/f
______
!___
yg
_ \ ô^ -V
-~
................
"-~ A ~-~0
"^'^
^^
Jo^
70
FIG. 87.
600 800 1000km
by Hendrik R.Rypkema
^Prepared
80
^ L
200 400
100 200 300 400 500 600 miles
-10
.1
60
50
L.0
Distribution of Campylopusjulicaulis (0) and C. longicellularis (0).
Along road MG 55 between Morro do Pilar and Rio

Peixe, Frahm 1426 (hb. Frahm);along road BR 135
between Itabirito and Belo Horizonte, Frahm 1431
(hb. Frahm),Frahm, Camp. Bras. Exsicc. 19 (ALTA,
KR, M, MO, NFLD, NICH, NY, PC, S, SP, TENN,
U); km 392 along road BR 135, Mun. de Ouro Preto,
Frahm 1427 (hb. Frahm);SierraMoeda NW of Ouro
Preto, Frahm, Camp. Bras. Exsicc. 18 (ALTA, B, C,
DUIS, FLAS, GOET, GZU, H, HBG, INPA, JE, KR,
M, MO, NFLD, NICH, NY, PC, S, SP, TENN, U);
SerraMoeda, Mun. de Itabirito, Vital5529 (SP);along
road Belo Horizonte-Juizde Fora, Vital6501 (SP);N

of Barbacenaalong road BR 135, Frahm 1430 (hb.
Frahm);Caldas, Widgrens.n. (HBG, H-BR, H-SOL).
Km 98 along road BR 116, Mun de Rio NePARANA:
gro, Vital 5603 (SP); 5 km S of Jaguariaiva,24?15'S,
49044'W,Vital10636 (SP);Ponte Grossa, ParqueNacional Vila Velha, Frahm 1416 (hb. Frahm). Rio DE
JANEIRO:Corcovado, v. Lutzelburg s.n. (JE); Apiahy,
Puiggari 390, 975 (JE);Serrado Itatiaia, Dusen s.n.

(NY); Parque Nacional Itatiaia, Rio Campo Belo,
22?30'S,44035'W,Landrum2182 (NY);nearRio Lago
Azul. Vital4857 (SP), Schiffner615, 616, 436 (H-BR);
Serrados Orgaos, Goeldi s.n. (U). SANTACATARINA: lamellosecosta,this taxonwas placedas a variety
Piloes,Palhoca,Reitz&Klein3235(FH).RiodeBrito, of C. cubensis(Frahm, 1985c). The latter much
Yano2278(SP).SXoPAULO:
Santos,Horeaus.n.(JE),
Mosen 351 (H-BR);Corumbatai,Vital 4905 (SP); 37 resembles C. lamellinervisvar. exaltatus with
kmE of Brotas,Vital4841(SP);km62 alongroadSao straightand erectpatent,only crispedleaveswhen
Barras,23?58'S,47059'W,Vital dry, but never with tortuose leaves. Therefore,
MiguelArcanjo-Sete
7716 (SP);Serrado Bocaina,alongroadCamposda it seemed much more reasonableto place C. exFrahm1425(hb.Frahm); altatus in C. cubensis.
Cunha-Sao
JosedosBareiros,
Campylopus cubensis,
IlhaComprida,
2502'S,47'55'W,Vital6754(SP);Ilha
has
a
border
of
narrow,elongate cells
however,
do CardosoMun. de Cananeia,Frahm 1429 (hb.
at the basal marginof the leaf and a non-lamelVital8767(SP).
Frahm);Mun.de Paraibuna,
lose costa, whereasC. lamellinervishas a border
Campylopuslamellinervismuch resemblesC. of small rectangularcells and a lamellose costa.
huallagensisvar. weberbaueriin the largesize of The plants referredto C. exaltatus were, howthe plants and the leaves and especiallythe areever, large expressions of C. cubensis.Theriot,
olation, which is identical in both species. Dif- who combined C. exaltatuswith C. penicillatus,
ferencesare discussed under the latter taxon.
might have seen the type of C. exaltatus.Therefore it seems to make more sense to follow the
39b. Campylopus lamellinervis var. exaltatus concept of Theriot to place C. exaltatuswith C.
(C. Muller) J.-P. Frahm, Bryoph. Bibl. 5: 78. lamellinervis.This is supportedby the existence
1975. Campylopuspenicillatusvar. exaltatus of specimens with a borderof short rectangular
(C. Miller) Theriot, Mem. Soc. Cubana Hist. cells at the basal part of the leaf and a lamellose
Nat. 13: 117. 1939. C. cubensisSulliant var. costa, which show identical relations as in C.
exaltatus(C. Muller)J.-P. Frahm,Nova Hed- lamellinervis.
wigia 41: 275. 1985. Dicranumexaltatum C.
Miller, Linnaea 42: 472. 1879. Campylopus
39. Campylopus longicellularis J.-P. Frahm,
exaltatus (C. Miller) Par., Ind. bryol. 246,
CryptogamieBryol. Lichenol7(4): 443. 1986.
1894.Type.Venezuela.Withoutlocality,FendType. Colombia, Meta: Paramo de Sumapaz,
could
lers.n. (holotypedestroyedat B, isotypes
Figs. 87, 90.
Cleef777a (U).
not be located).
Plants to 6 cm, blackish,erect, equallyfoliate
Plants resemblingvar. lamellinervisin all anwith
erectspreadingleaves.Leavesto 8 mm long,
atomical structures,but with erect patent, dark
narrow
lanceolate. Costa filling 2/33/4of the leaf
green, not tortuose, recurved and light green
in a long, smooth or only in the
excurrent
width,
leaves. The stems are taller, up to 15 cm, and
serrate
awn, in transversesection
apex
slightly
the leaves longer,to 15 mm.
bandsofstereids, smooth
with
and
ventral
dorsal
Distribution (Fig. 89). On soil in Pinus and
or
at
back
slightlyridgedin the upperpartof the
Podocarpusforest between 500 and 900 m in
leaf.
Lamina
reachingonly l/3 of the leaf length.
Cuba and Puerto Rico, also in northernVeneBasallaminalcells thin-walled,
Alar
cells
lacking.
zuela.
shortlyrectangular(ca. 3:1), ca. 45-55 x 15-20
Sierrade la ,um.Upperlaminalcells elongate,incrassate,pitSpecimensexamined.CUBA. ORIENTE:
GrandPiedra,sobreSanSebastian,P6cs 9119A(EGR); ted, ca. 70-80 x 8-10 ,m (8-10:1).
SierraCristal,Samek et al. s.n. (PRC).
PUERTORICO.MaricaoStateForest,km 15.8along
Distribution (Fig. 87). On soil in paramos,
Allen
6371
road 120,
(MO).
Peninsula de Paraguana, hithertofound only in Venezuelaand Colombia
VENEZUELA. FALCON:
Cerrode SantaAna,L6pez-F.19452(FLAS).
in altitudesbetween 3000 and 4000 m.
As in some other C. Miller species, the type
ParaSpecimensexamined.COLOMBIA.BOYACA:
specimen was destroyed at the Botanical Mu- mo de la Rusia,Cleef7056a(U);Paramosal NW de
seum in Berlinand no isotypes could be located. Belen, Cleef2098 (U). CAUCA:CordilleraCentral,La19109(US).
Therefore,this remainsa dubioustaxon. Due to gunitade las Casitas,Cuatrecasas
VENEZUELA. TACHIRA:
Paramo de Tama, Steythe lack of authenticmaterialand based on ma- ermark
98636(US).
terialkept at NY, which shows a borderof elonThe systematic position of this species is not
gate cells at the basal part of the leaf and a non-
130
Flora Neotropica
mm C
I/
^
y A
FIG. 88. Campylopuslamellinervisvar. lamellinervis(Vital 6026, SP). A. Plant. B. Leaf. C. Leaf-tip. D.
details = 50 urm.
fully clear, for elongate upper laminal cells are CampylopussubfulvusTheriot, Rev. Bryol. Lichenol
11: 45. 1939. Type. Bolivia. Yungasof La Paz, Troll
rarelymet with in this genus.
38a (holotype, PC).
40. Campylopusluteus (C. Muller) Paris, Ind.

bryol. 254. 1894. Thysanomitrionluteum C.
Miiller, Linnaea 42: 470. 1879. Type. Venezuela. Tovar, Fendler39 (holotype, destroyed
at B; lectotypus nov., H-BR; isotype,
H-SOL).
Figs. 91, 92.
Plants to 3 cm, in loose yellowishgreen,glossy

tufts. Stems equally foliate, fertile plants ending
in comal tufts. Leaves to 7 mm long, from ovate
base contractedinto a long, narrow,subtubular

subula,the subulaas long or longerthan the leaf
base, more or less toothed at margins and the
80
X\
131
70
60
i?
50
9
----------
-------------
- ---
200
40
400
600
800 1000km
100 200 300 400 5600 600 miles
FIG. 89. Distributionof Campylopuslamellinervisvar. lamellinervis(-) and C. lamellinervisvar. exaltatus

(0).
Flora Neotropica
132
1II
"'B
FIG. 90. Campylopuslongicellularis(Cleef 7036a, U). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails = 50 tim.
back. Costa filling 1/2of the leaf base, excurrent

in the subula, in transverse section with large,
ventral hyalocystsand dorsalgroups of stereids,
smooth or slightlyridgedat back.Alarcells, large,
inflated,reddish,protrudinginto the costa. Basal
laminal cells rectangular,incrassate,ca. 1:5, ca.
6-13 x 19-48 Mm,at marginsnarrower,forming
a border. Upper laminal cells incrassate,
subquadrate,shortrectangularto oblique, 4-6 x

10-16 Mm.
Vegetativepropagationby deciduousleaves or
stem tips.
Seta 8-10 mm, flexuose. Capsule 2 mm,
curved. Operculumobliquely rostrate. Calyptra
fringedat base, pale yellowish.
Distribution (Fig. 92). On soil, soil covered
133
FIG. 91. Campylopusluteus (Fendler39, H-BR). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails = 50 MAm.
rocks and trunks of trees in montane rainforests

between 800 and 3200 m altitude in Guyana,
Venezuela, Colombia, Peru and Bolivia.
de Cali 32?22'N,76?43'W,Hartman 243a (COLO).
Specimensexamined.GUYANA.N slope of Mt. Roraima, 5?17'N,60?43'W,Gradstein5336 (U).

CordilleraLeonora
COLOMBIA.CUNDINAMARCA:
60 km NNE de Bogota, v. d. Hammen & Jaramillo
3147, 3182 (U); Represa del Neusa, Cordillerade la
cubaji,Griffinet al. 1031 (FLAS).LlanoComponedero

arribade la Piedrade Pirela,Ruiz-T. & Lopez-F.6555
Piramo de Tama, Ruiz-Terdnet al.
(FLAS).TAcHmRA:
8372 (FLAS);Paramo de la Negra entre los estados
Meriday Tchira, Ruiz-Terdn8404b(FLAS).TRUJILLO:
Senda, Cleef 6659 (U). VALLEDE CAUCA:Farallones

VENEZUELA. FALC6N:Cerro Santa Ana, Steyermark & Brown 94570c (MO); Ocalite, Pittier 453 (MO).
MERIDA:Sierra de Santo Domingo, Piramo de Mu-
Flora Neotropica
134
80
70
60
50
40
30
^0
\"1~~~~~~
200
*?
?-S--
0
-
--
-_____
400
600
800 1000km
l
100
200 300 400 500 600 miles
Prepared
o0o
'1,
FI
ostbo ofCmy0u
FIG.0 92.
Paramode LaCanada,Ruiz-Terdn9186 (FLAS);above

the village of Santa Rita, distr. Bocon6, Griffinet al.
1194 (FLAS);Paramode Guaramacalabove Bocon6,
Griffinet al. 979, 1092 (FLAS).
ues
Distribution
of
Campylopus uteu
.10
41. Campylopus nivalis (Bridel) Bridel, Bryol.

univ. 1: 477. 1826.
Pampa Tambo, Hermann
25250, 25252 (hb. Frahm).
A species which is conspicuous by its long,

narrow leaf tip which is as long or longer as the
leaf base, the subquadrate upper laminal cells
and the alar cells protruding into the costa.

Campylopus nivalis
1 Upperlaminalcells oval, ca. 4:1. ..41a. var. nivalis.
1 Upperlaminalcells shortrectangular,
ca. 2:1. ....
........................
41b. var. multicapsularis.
135
FIG. 93. Campylopusnivalisvar. nivalis(Frahmet al. P 236, B). A. Plant.B. Leaf.C. Leaf-tip.D. Transverse
50 im.
41a. Campylopus nivalis (Bridel) Bridel var. nivalis Weisia nivalis Bridel, Spec. musc. frond.
1: 123. 1806. Thysanomitrion nivale (Bridel)
Arnott, Mem. Soc. Linn. Soc. Paris 5: 263.
1827. Type. Ile Bourbon. Bory St. Vincent s. n.
(holotype, B).
Figs. 5A, 93, 94.
CampylopusarseneiTheriot, SmithsonianMisc. Collect. 78(2): 5. 1926. Type. Mexico. CerroAzul, Arsene 4981 (lectotypusnov., PC).
Campylopuschrismarii(C. Muller) Mitten, J. Linn.
Soc. Bot 12: 88. 1869. DicranumchrismariiC. Mil-
ler, Bot. Zeit. 13: 761. 1855. Type. Mexico. Michoacan, Chrismars.n. (holotype, destroyed at B;
lectotypusnov., NY; isolectotypes,H-BR, S).
Campylopusdestructilis(C. Miiller)Jaeger.Ber. Thatigk.St. GallischenNaturwiss.Ges. 1870-1871:430.
1872. DicranumdestructileC. Miiller,Bot. Zeit. 17:
220. 1859. Type. Mexico. Schiede s.n. (holotype,
destroyedat B; isotypes could not be located).
Campylopusfriabilis(Hampe)Jaeger,Ber.Thatigk.St.
Gallischen Naturwiss.Ges. 1870-1871: 432. 1872.
Dicranumfriabile Hampe in C. Miller, Bot. Zeit.
17: 220. 1859. Type. Costa Rica. Las Nubes, Wendland s.n. (holotype,n.v.; isotype, GOET).
Flora Neotropica
136
100
110
90
so
HO103
60
70
80
80
70
60
30
20
.0
200
400
600
800
1000k

100
FIG. 94.
90
00
70
60
Distribution of Campylopus nivalis s. lat.
Campylopuskrauseanus(Hampe & Lorentz)Jaeger,

Ber. Thatigk.St. GallischenNaturwiss.Ges. 18701871: 427. 1872. DicranumkrauseanumHampe &
Lorentz, Bot. Zeit. 27: 434. 1869. Type. Ecuador.
Krauses.n. (holotype,n.v.; isotype, GOET).
Campylopuspoasensis Renauld & Cardot, Bull. Soc.
Roy. Bot. Belgique3(1):148. 1893.Type.CostaRica.

VolcAnde Poas, Pittiers.n. (n.v.).
CampylopussubfimbriatusTh6riot,Rev. Bryol.Lichenol. 11: 40. 1939. Type. Bolivia. Illampu, Troll 69
(lectotype,PC).
CampylopussuboblongusHerzog,Beih. Bot. Centralbl.
137
61b: 585. 1942.Type.Ecuador.Withoutlocality, nearUruapan,Rejmdnek4, 6, 51, 52 (PRC);Mun. de
Tlalmanalco,3 km E de San Rafael, Cruz Cisneros

Espinoza 29 (holotype,JE).
CampylopustolucensisBartram,Rev. Bryol.Lich6nol. 1655 (FLAS);Sierra del Ajusco, 19?12'N, 90?12'W,
15: 21. 1946. Campylopuschrismariivar. tolucensis Vivas259, 260 (MEXU);Gap at Nevado de Toluca,
J.-P. Frahm,Bryologist80: 120. 1977. Sharp et al. 1547b (TENN);southernrim of Nevado
(Bartram)
Type.Mexico.Nevadode Toluca,Bartram4114 de Toluca,Frahm, Camp.Exsicc. 8 (ALTA,B, BING,
BM, BUF, C, DUKE, EGR, F, FLAS,G, GRO, GZU,
(holotype,FH).
andfurthersynonymsfromAfrica(Frahm,1985a). H, HBG, KRA, MEXU, NAM, NFLD, NICH, NY,
PC, PRC, POZ, RNG, S, U); summit of Nevado de
Plants to 5 cm high, usually 1-3 cm, yellowish Toluca, Vitt 17919 (ALTA); Volcan Popocatepetl,
green, in tufts. Stems tomentose below, erect. Rickett76 (NY); Cleef&Delgadillo10236, 10248 (U);
Leaves 5 mm long, lanceolate,ending in a more Volcan de Ixtaccihuatl,Eggers & Frahm s.n. (hb.
MICHOACAN:
Puerto Garnica E of Morelia,
or less elongatechanelledpoint, serrulateat the Frahm).
Sharpet al. 2673 (TENN);Morelia,Campanario,Arextreme apex or smooth. Costa filling 2/3of the sene 7573 (NY); 13 km W of Hidalgo, Sharp 8774
leaf width, narrowedat base, wider above, ex- (TENN);PuertoCerrode GarnicaNat. Park44 km W
current,in transversesection with large ventral of Hidalgo,Iwatsuki&Sharp2947 (TENN).MoRELOs:
M6xico-Cuemavaca,Schwab SN 93
hyalocystscoveringmore than half of the trans- Along highway
34 mi E of Mitla on road to
(hb. Frahm).OAXACA:
verse section, and dorsal groups of stereids or Tamazulapam,
Sharp et al. 2973 (TENN); between
even small groups of stereids, ridgedat back in Tuxtepecand Oaxaca,Sharp et al. 2316 (TENN);Sithe upper part of the leaf, smooth below. Alar erraSan FelipeNE of Oaxaca,Iwatsuki& Sharp5162
cells not or slightlydifferentiated,hyaline. Basal (TENN);10 km NE Llanode las Flores,Delgadillo769
N of Ixtlan,Hermann26197 (NY);Llano
laminal cells thin-walled,rectangular,10-16 x (NY); 29 mi
Grande, Sierra Juarez, Sharp & McFarland 859a
26-54 Am,in severalrows narrowerat margins. (TENN);pass above Llano de las Flores on the road
Upper laminalcells shortrectangularto oval, 3- betweenIxtlanand Tuxtepec,Sharpet al. 2369b, Iwatsuki & Sharp 5267, 5383, 5424, 5449 (TENN);N of
10 x 10-19 Am,incrassate.
Seta 7-8 mm long, yellowish to brownish. Oaxacaon Hwy 175, Sharp 4777b (TENN). PUEBLA:
15 mi S of Chignahuapan,Hermann 26451 (F); PoCapsule 1.5 mm long, erect, symmetric, cylin- pocatepetlsouth of Tlamacas,den Held & van Rhijn
dric, furrowedwhen dry or empty. Operculum HM 6 (U); below Tlapacoyin, Sharp et al. 1121
(TENN);Ixtaccihuatl,Sharp4298 (TENN).VERACUZ:
long-rostrate.Calyptraciliate at base or not.
Distribution(Fig. 94). A predominantlyalpine Tlacotepec, 16 km NE of Huayacocotla,Juarez 1000
Pass along road between Orizaba and Tespecies on a large variety of substratessuch as (TENN);
huacan,Eggers & Frahm 792036 (hb. Frahm).
dry sandy soil or volcanic ashes in dry, tussock

grass punas, wet humic soil or peat in paramos
or wet punas,on rocksandboulders,on wet cliffs,
in open habitatsor in alpineforestssuch as Polylepis forests, from the tree line to 4400 m; also
in open habitats as open slopes, banks of trails
and roadsbelow the forestline down to 2000 m.
In Mexico also in light forestssuch as oak-pineforests, where this species occurs also on tree
trunks and rotten wood. From Mexico, Guatemala, Costa Rica, Venezuela, Colombia, Ecuador, Peru and Bolivia to northern Argentina.
Found only once on the CaribbeanIslands on
Hispaniolain an open pine foreston soil at 2700
m elevation, perhapsdue to an occasionalspore
dispersal.Also in alpine to subalpineregionsof
tropicalAfrica.
GUATEMALA. HUEHUETENANGO:
Road beyond La
Volcan
Pradera,Standley81776 (NY). SAN MARCOS:
Tajumulco,Shannon 3698 (NY). Quezaltenango,Los
Vahos, Standley86125, 86121 (NY);Volcande Agua,
Godman&Salvins.n. (NY);VolcanSantaMaria,Steyermark34182 (NY); E of Totonicapan,Sharp 2595
(NY); Volcan de Zunil, Standley 85890, Steyermark
34846 (NY); mountainabove Totonicapan,Standley
84502 (NY).
COSTA RICA:ALAJUELA:
Volcin de Poas, Griffin
et al. 20000 (NY); Griffin&Araya47 (NY). SAN JOSE:
Cerrode la Vuelta,Standley43656, 43929 (NY); Volcande Poas,Alfaro55 (NY);Griffin&Araya47 (FLAS);
Cordillerade Talamanca,ChirripoMassif,Kuhbier624
(BREM);Volcan Irazu,Schultes11845 (FLAS).
COLOMBIA. ARAUCA:SierraNevada del Cocuy,
Pefiade Arnical,Cleef9492b
Cleef8870 (U). BOYACA:
(U); Paramode Pisva, Cleef4564 (U); SierraNevada
del Cocuy, QuebradaBocatoma, Cleef & Florschiitz
5651 (U); Alto Valle Lagunillas,Cleef & Florschiitz
5546 (U); Paramode la Rusia, Cleef6759, 7180, 7404
Specimensexamined.MEXICO.CHIAPAS:4.5 mi W (U); Paramo de Chita, Cleef 9949 (U). CAUCA:Mt.
of San Crist6balde las Casas,Hermann26422 (F); Purac6,Killip6579 (NY). CUNDINAMARCA:
Paramode
VolcanTacana,Mun. de Union Juarez,Breedlove Sumapaz, Cleef 4930 (U); Chisaca, Cleef 3615 (U);
About 9 mi W of La ParamoentreCoguay SanCayetano,Cleef4875, 6485
24339, 2334 (MO). DURANGO:
Ladera (U); Represadel Neusa, Cleef& Jaramillo4169a (U);
Ciudad,Bowerset al. 5097b (TENN).JALISCO:
Norte del Nevadodel Tolima, 19?34'N,103037'W, Paramode Sumapaz,Cleef 4930 (U); Paramode PaVolcanParicutin lacio, Cleef5256A(U);Subachoque,CuchillaEl TablaDelgadillo4629 (MEXU).MEXIco:
138
Flora Neotropica
zo, Linares & Bulla 745, 854, etc. (COLO);Bogota,
Hegewald7275 (hb. Hegewald);between Contumaza
Paramo de Monserrate, Sturm 13 (NY). MAGDALENA: and Cascas, Hegewald 7351 (hb. Hegewald).Cuzco:
Bues 1097a, 1478 (NY); 1 km N of passbetweenUrcos

and Juliaca, Frahm 824000, 823982 (hb. Frahm).
JuNiN:LagunaChaulacochanearLa Oroya,Hegewald
5407a (hb. Frahm);30 km E of Huancayo,Hegewald
9184 (hb. Frahm).LIBERTAD:
691, 715 (U). SANTANDER:
Otuzco,Sagasteguiet al.
Edge of Paramo de las Vegas, Killip & Smith 15754a (NY); vic. Vetas, Killip & 11647 (NY); CerroCampananear Trujillo,Hegewald
7263 (hb. Hegewald);Pampasde la Julia near QuiruSmith 17370 (BM).
Sierra de Merida, Grif- vilca, Hegewald5997 (hb. Hegewald).PAsco: Along
VENEZUELA. LIBERTADOR:
fin 17471 (NY). MERIDA: Sierrade Santo Domingo, road from Cerrodo Pasco to Lima, 11?00'S,76?20'W,
Paramode Mucubaji,Griffinet al. 1053, 1074, 1113 Vitt21727 (ALTA).
1 km S of LagunasTahua
BOLIVIA.COCHABAMBA:
(FLAS);Paramode Timotes, Lopez-F. 18952 (FLAS);
Quebradade El Padre,L6pez-F. 15451 (FLAS);Sierra Cruz, 17?15'S,66?24'W,Lewis 79-2477 (F); area of
de La Culata,Paramode Los Conejos,Ruiz-T. 6959 LagunaTarucani,17?14'S,66?24'W,Lewis 79-2443D
(FLAS);SierraNevada de Merida,Lagunade Los An- (F). LA PAZ: 1 km below Escoma-Charasaniroad,
teojos, Griffinet al. 298 (FLAS);Telefericode Merida, 15?16'S,69?02'W,Lewis 79-900, 922 (F); 5 km NE of
Iguada, Onraedt5782, 5797 etc. (hb. Frahm);Loma Milluni, 16?18'S,68?06'W,Lewis79-1648 (F);Nevado
Paramo Jankho Uma above St. Antonio, 15?51'S,68?34'W,
redonda,Onraedt8237 (hb.Frahm).TACHIRA:
de TamaaboveVillaPaez,Griffinet al. 938,212 (FLAS). Lewis 79-1425, 1573 (F); Nevado Huayna Potosi,
Cerro
Paramo La Estrella above La Mesa de EsTRUJILLO:
16?17'S,68?08'W,Lewis 78-1938 (F). POTOSI:
et al. 1410 (FLAS);cumbredel Paramo Kari Kari SE of Potosi, 19'36'S, 65?42'W,Lewis 79nujaque,Griffin
283 (F). TARIJA:
5 mi NE of Iscacayachi,21?27'S,
de La Cristalina,Ruiz-Terdn9026 (FLAS).
Around LagunaToreadorain 64?54'W,Lewis 79-654, 701 (F).
ECUADOR. AZUAY:
ARGENTINA.W of Cordoba,v. Hiibschmanns.n.
ParqueNacional de Cajas, 79?13'W,2?46'S,Lzjtnant
& Molau 14821 (GB). CARCHI:Paramoabove Tufiio, (hb. v. Hibschmann).
Steere8760A (NY);Valle de Maldonadokm 53 on the
According to the broad ecological amplitude
roadTulcan-Maldonado,78?3'W,0?50'N,Holm-NielMarand
San
sen et al. 5817 (AAU, GB). IMBABURA:
Lago
geographical range, this species is quite varicos, CayambeMt., Cazalet& Pennington63, 73 (NY). able with respect to the density of the tufts, presLOJA:Krauses.n. (NY). NAPO:Paramode Quilindana, ence of a
tomentum, size of the plants, length of
LagunaYurac Cocha, Kieftet al. 180 (U); Paramode the leaves and
length of the leaf tip. The leaf tips
E267
Steere
GuamaniN of Quito-Baezaroad,
(NY);
LagunaParacocha,78?9'W,0?16'S,Lcjtnant& Molau can be relatively short (and usually serrulate) or
11086 (NY);CerroSumaco,77?39'W,0?35'S,Lsjtnant much elongate into a long, smooth acumen. Such
Paramo de Gua& Molau 13010 (NY). PICHINCHA:
forms previously had been separated as var. submani along Quito-Baeza road, 78?10'W, 0?17'S,
J.-P. Frahm by studies of herLojtnant& Molau 11081, Balslev1623A(NY);Volcan oblongus (Herzog)
de Pichincha,Steere&Balslev26016 (NY); N slope of barium material. Field studies revealed that this
volcan Pichincha,Balslev 1748 (NY). Quito, Jameson is a modification of more shaded habitats (such
s.n. (NY).
as Polylepis forests) and there are all possible
PERU. ANCASH:Near Llanganuco, Hegewald 7554
All these modifications also apintergradations.
(hb. Frahm);near Laguna Safuna NE of Alpamayo,
to
the
African
ply
populations. The forms with
Bunin B 49872, 49876 (COL);along road from Catac
to Chavin,Frahm, Camp.Peruv.Exsicc. 6 (ALTA,B, longer leaf tips are correlated with longer upper
BM, BUF, DUKE, EGR, F, FLAS, G, GRO, GZU, laminal cells and are expressions of more humid
HBG, KR, NAM, NICH, NY, MEXU, NFLD, PC, habitats. Notably, forms with short leaf tips and
POZ,RNG, S, U). LagunaLlaca27 km NE of Huaraz, shorter
upper laminal cells have been collected
Frahm, Camp. Exsicc. 69, 79 (ALTA, B, BING, BM,
on dry, volcanic mountains.
frequently
BUF, C, CBG, F, FLAS, G, GRO, GZU, H, HBG,
In SE Asia, the Afro-American C. nivalis is
KRA, MEXU, NICH, NY, PC, PRC, S, U); Frahm et
al. P 235, P 236 (B); along road from Huanuco to replaced by a very similar species, C. austroHuaraz between Huallanca and Chiquian, Frahm subulatus Geheeb &
Brotherus, found in the al823954 (hb. Frahm). AYACUCHO:
Along road from
belt
of
New
Guinea.
This is the main area
pine
Huantato San FranciscobetweenTambo and Quinoa,
Frahm 824020 (hb. Frahm);at pass Huamina,Hege- with large, extensive alpine habitats in SE Asia.
wald8996 (hb. Frahm);along road from Ayacuchoto
Campylopus schimperi Milde is a similar speOcros, Frahm, Camp. Exsicc. 57 (ALTA, B, BING, cies which occurs in alpine habitats through the
BM, BUF, C, DUKE, EGR, F, FLAS,G, GRO, GZU, Holarctic. It differs from C. nivalis
by rectangular
H, HBG, KRA, MEXU, NAM, NFLD, NICH, NY,
and
not
oval
In America it
laminal
cells.
upper
Las
CAJAMARCA:
LaguPC, PRC, POZ, RNG, S, U).
nas, Hegewald 6210 (hb. Hegewald); Contumaza, ranges from the Arctic to Colorado.
ParqueNacional de la SierraNevada de SantaMarta,
Rangel&vanReenen975, 1012, 1035, 1058 (U). META:
Paramode Sumapaz,Cleef 1194, 1384 (U); RISARALDA:Mun. de Pereira,Volcin de Sta. Rosa, vanReenen
139
000
FIG. 95. Campylopus nivalis var. multicapsularis (Frahm 824001, hb. Frahm). A. Plant. B. Leaf. C. Leaftip. D. Transverse section of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for
microscopicdetails = 50 mm.
41a. Campylopus nivalis var. multicapsularis (C.

Muller) comb. et stat. nov.
Figs. 94, 95.
Campylopusmulticapsularis(C. Miller) Schimperin
Paris, Ind. bryol. Suppl. 94. 1900. DicranummulticapsulareC. Miller, Prodr.bryol. boliv. 34. 1897.
Type. Bolivia. Larecaja,Mandon 1611 (holotype,B;
isotype, NY).
Campylopusinsignis Herzog, Bibl. Bot. 87: 23. 1916.
Type. Bolivia. Abra San Mateo. Herzog 3721 (holotype, JE).
CampylopusleucobasisHerzog, Biblioth. Bot. 87: 23.

1916, nom. inval. cf. Index Muscorum.
CampylopussubannotinusTheriot& P. de Varde,Rev.
Bryol. Lichenol. 4301 (holotype, PC; isotype, JE).
Plants resembling much those of C. nivalis var.
nivalis, to 3 cm high, in tufts, light green above,
reddish tomentose below. Stems equally foliate.
Leaves to 5 mm long, lanceolate. Costa filling /3
of the leaf width, excurrent in a serrate tip, in
140
Flora Neotropica
transversesection with ventralstereidsand dorsal groups of substereids,smooth at back in the

lower part and ridged in the upper part of the
leaf.Alarcellslackingor scarcelydeveloped.Basal
laminal cells elongate rectangular,thin-walled,
6-12 x 12-26 um, at marginsin 5-6 rows narrower.Upper laminalcells incrassate,shortrectangularto oblique, 6-10 x 10-19 ,m, ca. 1:2.
Sporophyte as in C. nivalis var. nivalis.
Distribution (Fig. 94). On moist soil in para-
mos and wet punas between 3300 and 4500 m,

rarelyin open places at lower altitudes,throughout the Andes from Venezuelato Colombia,Ecuador, Peru to Bolivia.
ParaSpecimensexamined.COLOMBIA.BOYACA:
mo de la Rusia N-NW de Duitama, Cleef 6785 (U);
SierraNevada de Cocuy, Alto Valle Lagunillas,Cleef
& Florschiitz5770 (U); Vado Hondo, Siberia, Cleef
9353 (U); Paramode la Sama entreSogamosoy Vado
Paramo entre
Hondo, Cleef9212 (U). CUNDINAMARCA:
laris is combined here as a new variety of C.

nivalis. The taxa seem to differ in habitat: C.
nivalis has a broader ecological amplitude, extendingeven to dry habitats(as dry grasspunas,
even volanic ash), whereas var. multicapsularis
preferswet habitats, such as peaty soil around
small lakes or drippingcliffsin paramosand punas. It has, therefore, a smaller range than C.
nivalisvar. nivalis,not extendingbeyondColombia to the north or Bolivia to the south.
The shorter upper laminal cells of var. multicapsularisseem to indicateonly a modification,
perhapsinfluencedby the differenthabitat.However, measurementsof the length-widthratio of
the upperlaminal cells revealedno overlapping,
thus indicatingthat the taxa seem to be genetically distinct.
42. Campylopusoblongus Theriot, Rev. Bryol.
Coguay San Cayetano,LagunaVerde,Cleef6348 (U);

Lichenol.11: 62. 1939. Type. Colombia,above
Paramode Sumapaz,Chisaca,Cleef3613 (U); Paramo
Bogotaat 3000-3200 m, Troll2082 (holotype,
de Cruz Verde, Cleef 3378 (U).
PC).
Figs. 96, 97.
VENEZUELA.MERIDA:
SierraNevada de Merida
betweenla Aguadaand La Montania,Griffinet al. 256
bolivianus
Theriotex J.-P.Frahm,LindCampylopus
(FLAS); Teleferico, Loma Redonda, Cleef & Huber
7:
28.
1981.
Bolivia.Sillunticava,
Yunbergia
Type.
4789 (U). TRUJILLO:
QuebradaEl PajaritoentreTuiiagasdela Paz,Troll122a(holotype,PC;isotype,JE).
me y Las Mesitas,Ruiz-Terdn7537 (FLAS).
Road up N slope of volcan
ECUADOR.IMBABURA:
Plantsin loose tufts,to 4 cm high,green.Stems
de Cayambe,Steere26881 (NY).
tomentoseat base, equallyfoliatewith erect
erect,
PERU.ANCASH:
LagunaLlanganuco,Hegewald7533
somewhatflexuoseleaves.Leaves6-7 mm
patent,
road
from
Huanuco
to
on
(hb. Hegewald); pass along
Huaraz,Frahm, Camp. Peruv.Exsicc. 15 (ALTA, B, long, lanceolate, narrowedin a long, chanelled,
BM, BUF, DUKE, EGR, F, FLAS, G, GRO, GZU, serrateacumen. Costa filling3/4of the leaf width,
HBG, KR, NAM, NICH, NY, MEXU, NFLD, PC, excurrent,in transversesection with lax ventral
POZ, RNG, S, U). Cuzco: Pass between Urcos and
hyalocystsand a band of dorsal stereids,ridged
Juliaca,Frahm 824001 (hb. Frahm).HUANCAVELICA:
Ichoquenus,Hegewald 9157 (hb. Hegewald).JUNiN: at back in the upper part, smooth below. Alar
Laguna Caulacochanear La Oroya, Hegewald 5406 cells inflated,auriculate,conspicuous,hyalineor
(hb. Frahm);Anascocha near Canchayllo,Hegewald reddish,protrudinginto the costa. Basal laminal
5403 (hb. Hegewald);30 km E ofHuancayo,Hegewald
9175 (hb. Hegewald).Pass Huaytallapallana50.5 km cells rectangular,incrassate,9-15 x 40-45 ,m.
E of Huancayo, Hegewald 9157 (hb. Hegewald).LA Upperlaminalcells subquadrateor short-rectanLIBERTAD:
HuancamarcanearOtuzco,Hegewald5164 gular,partly oblique, 5-8 x 9-13 Am.
(hb. Hegewald);CerroChiputur,Hegewald7450 (hb.
Seta 8 mm long. Capsule2 mm, oblique, with
Hegewald).PUNO:7 km E of Pass La Raya, Hegewald rostrateoperculum.Calyptraentire or ciliate at
5495 (hb. Hegewald).
BOLIVIA.LAPAZ:Head of Rio ZongoN of La Paz, base.
Distribution(Fig. 97). On moist soil, soil-cov16?17'S,68?06'W,Lewis 79-1816, 79-1817 (F). ORURO:AguasTermalesde Rio JunthumaKhuchu,18?6'S, ered rocks, dead tree stumps, fallen decayinglog
69?2'W,Lewis 79-2128, 79-2136 (F). TARIJA: 5 mi NE in montane and especiallysubalpinerain forests
of Iscayachi,21?27'S,64?54'W,Lewis 79-698 (F).
Mexico in
between200 and
(in
pine-oakforests)
3300 m elevation; most frequentbetween 2500
and 3000 m, more rarelydown to 1000 m and
only one record on the lomas of Peru between
oblique. Since the range of C. multicapsularis 200 and 600 m. In Mexico, CostaRica, Panama,
falls into the range of C. nivalis, C. multicapsu- Venezuela,Colombia, Peru and Bolivia.
Campylopus multicapsularis seems to differ

from C. nivalis only by the shorter upper laminal
cells, which are not oval but short-rectangular to
141
/''
<p\/~~~~~~
' ,L,,~,m~J
]J
"
;:,-
'
i
.
F
Y
A,,?
|uy 4.5cm
li(
'2o/D i
',:
"
I
I'5mm
(TENN); 23-24 km NE de Llano de las Flores, DelSpecimens examined. MEXICO. CHIAPAS:SW side
of Cerro Mozotal, mun. de Motozintla de Mendoza, gadillo 808 (MEXU); Puerto Gallo, Haagerova s.n.
La Ferreriaabove (PRC).
Breedlove25777 (MO). JALISCO:
COSTA RICA. SAN JOSE:Vic. Santa Maria, StanSierrade
Manantlan,Crum 818 (MICH). MORELOS:
Chichinautzin, mun. de Tepoztlan, Cdrdenas 1047 dley 41661 (NY).
PANAMA. Above Bajo Chorro, D'Arcy 11112F
(MEXU). MICHOACAN:
Along Hwy. 15 about 50 km
W of Cd. Hidalgo,Sharpet al. 2246 (TENN).OAXACA: (MO).
San Cristobal, ApolCOLOMBIA. CUNDINAMARCA:
SierraJuarezgap on route 175 between Oaxaca and
Tuxtepec, Websteret al. 670, Smith et al. 3049, Sharp linaire s.n. (PC); environs d'Alban, Onraedt 846g
et al. 4813, Richards&Sharp7078 (TENN);trailabove (FLAS);Paramo entre Cogua y San Cayetano, Cleef
Tamazulapam70 km E of Oaxaca,Smith et al. 4418 108 (U). SanJose de Guaviare,Schultes11123 (FLAS).
142
Flora Neotropica
-. -oc.
.
'~'
9c
e0
byb-F- RRypkenu-
..
FIG 97... Ditibto
*oo
FIG.I
200
'
soeo
-----
myou
-----.
u -o.(
70
so*0.=..
9.Dsruino
,03oo moo
,ooo~
of Ca
SC
lnu()adC
so
. o
.----
--
60
clu
RRypkema
Prepred
byHendrk
FIG. 97.
Distribution of Campylopus oblongus (0) and C occultus (0).
Frahm).Lomasde GuilmananearLima, Vargas9631

(MICH).
elongateoval upperlaminalcells and thin-walled

basal laminal cells, whereas the upper laminal
cells in C. oblongusare irregularlyquadrate,short
rectangularand oblique, and the basal laminal
cells are incrassate.
Campylopus bolivianus was described from
materialwhichTheriothad namedon herbarium
labels as C. alopecurus(C. Miiller) Paris. var.
bolivianus. These specimens had, however, no
relationship with C. alopecurus (C. heterostachys), since the basal laminal cells were incrassate, whereasin C. heterostachysthey arehyaline
and thin-walled.Therefore,the name bolivianus
was validated by describingit as a new species.
It had, however, alreadybeen describedby Theriot as C. oblongusfrom Colombia.
Campylopus oblongus is similar in appearance

to C. sharpii, which has, however, distinctly
43. Campylopusoccultus Mitten, J. Linn. Soc.

Bot. 12: 86. 1869. Type. Brazil. Parana:Weir
VENEZUELA.
MERIDA:Rio Capaz arriba de La
Azulita, Steyermark97129 (NY); Sierrade Santo Domingo, Paramode Mucubaji,Griffinet al. 945 (FLAS);
La Carboneraarea, Finca San Eusebio, Griffinet al.
1437 (FLAS);Cordillerade Merida,trail from La Escalerato Puentede Escalera,Luteynet al. 7864, 7872
4.6 km E ofZumbador, 7?56'N, 72?3'W,
(NY). TACHIRA:
Luteyn 9855 (NY).
PERU. AMAZONAS:
Along road from Chachapoyas
to Cajamarca,Frahm 825117 (hb. Frahm);on N-exposed grass-roof, Frahm 825114 (hb. Frahm).
AYACUCHO:Aina betweenHuantaand Rio Apurimac,
Killip & Smith 22738, 22790 (NY); along road Ayacucho-SanFrancisco15 km fromS. Francisco,Frahm,
Camp.Peruv,Exsicc. 16 (ALTA,B, BM, BUF, DUKE,
EGR, F, FLAS, G, GRO, GZU, HBG, KR, NAM,
NICH, NY, MEXU, NFLD, PC, POZ, RNG, S, U).
LIBERTAD:E of Huancamarca, Hegewald 7228 (hb.
143
'l
2o
o
5nm
FIG. 98. Campylopusoccultus(Frahm1447, hb. Frahm).A. Plant.B. Leaf.C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails = 50 Mm.
Montevideo, Arechavaletas.n. (holotype destroyed
at B; lectotypusnov., NY).
Campylopuscacti (C. Miller) Kindberg,Enum. Bryin.
exot. 88. 1889. Dicranumcacti C. Miller, Linnaea
XLIII:403. 1882. Type. Uruguay.Concepci6n,LoCampylopusacervatusMitten, J. Linn. Soc. Bot. 12:
87. 1869. Type. "Americaaustralis,"Lobb 122 (horentzs.n. (holotype,destroyedat B; lectotypusnov.,
BM; isolectotypes,JE, PM, S).
lotype, NY; isotypes, C, H-BR).
Campylopus calymperidictyon(Geheeb & Hampe)
Campylopusbrachymitrius(Geheeb& Hampe) KindKindberg,Enum. Bryin. exot. 88. 1889. Dicranum
berg, Enum. Bryin. exot. 88. 1889. Dicranum
brachymitriumGeheeb & Hampe, Flora 64: 343.
calymperidictyonGeheeb & Hampe, Flora64: 342.
1881. Type. Brazil.Apiahy,Puiggari331 (lectotype,
1881. Type. Brazil.Sao Paulo,Apiahy,Puiggari250
H-BR).
(isotypes, FH, S).
Campylopusdentato-acicularisC. Muller,Gen. musc.
Campylopusbrachythysanos(C. Miller) Paris, Ind.
frond. 270. 1900, nom. nud. Material.Brazil. Jabryol. Suppl. 89. 1900. Dicranumbrachythysanum
C. Miller, Hedwigia36: 97. 1897. Type. Uruguay.
guarone, Ule 708 (H-BR).
66 (holotype, NY; isotypes, F, H-BR).
Figs. 5D, 7C, 9B, 97, 98.
144
Flora Neotropica
CampylopusdicksoniaeC. Muller,Gen. musc. frond.

270. 1900, nom. nud. Material.Brazil.S. Catarina,
Ule 660, 1125 (H-BR).
Campylopusdivisus (Geheeb & Hampe) Paris, Ind.
bryol. 245. 1894. Dicranum divisium Geheeb &
Hampe, Flora 64: 345. 1881. Type. Brazil.Apiahy,
Puiggari323 (isotypes,H-BR, S).
CampylopusfuscatusBescherelle,J. Bot. 5: 146. 1891.
Type. Uruguay, Montevideo, Courbons.n. (holotype, PC).
Campylopusglauco-pallidusC. Miiller, Gen. musc.
frond.270. 1900, nom, nud. Material.Brazil.S. Catarina, Ule 703, 836 (H-BR).
Campylopushumifugus(C. Miller) Kindberg,Enum.
Bryin.exot. 88.1889. DicranumhumifugumC. Miller,LinnaeaXLIII:400. 1882.Type.Argentina.Gran
Chaco, Rio Secco, Lorentzs.n. (holotype,destroyed
at B; lectotypusnov., H-BR; isolectotype,FH).
Campylopusjoinvilleanus(Hampe) Jaeger,Ber. Thatigk. St. GallichenNaturwiss.Ges. 1877-1878: 382.
1880. Dicranum joinvilleanum Hampe, Vidensk.
45. 1872. Type. Brazil.Rio de Janeiro,Glaziou4557
(holotype,n.v.; isotype, H-BR).
Campylopusplatyneuron(Hampe) Jaeger,Ber. Thatigk. St. GallischenNaturwiss.Ges. 1877-1878: 382.
1880. Dicranum platyneuron Hampe, Vidensk.
139. 1875. Thysanomitrion
platyneuron(Hampe)C.
Miiller,Gen. musc. frond. 260. 1900. Type. Brazil.
Rio de Janeiro,Glaziou 705 (isotype, NY).
Campylopuspseudobrachmitrius
(C. Miller) Paris,Ind.
bryol. ed. 2, 1: 324. 1904. DicranumpseudobrachymitriumC. Miiller,Hedwigia39: 249. 1900. Type.
Brazil. Minas Gerais: Serra do Itatiaia, Ule 1803
Campylopus restingae C. Miiller ex Brotherus,
Denkschr.Akad.Wiss. Wien math.-nat.K1.83:260.
1926. Type. Brazil.Rio de Janeiro:Maua, Ule 212
(holotype,H-BR; isotypes, JE, KIEL,NY).
CampylopusrufescensBrotherusin Paris, Ind. bryol.
Suppl.96. 1900. DicranumrufescensC. Miiller,Bull.
Herb. Boissier VI: 34. 1898. Type. Brazil. Minas
Gerais: Serrado Ouro Preto, Ule 1348 (lectotype,
H-BR).
CampylopusscabrellusMitten, J. Linn. Soc. Bot. 12:
86. 1869. Type. Brazil. Without locality, Lobb s.n.
(holotype,NY).
C. Miiller,Hedwigia38
Campylopussubbrachymitrius
Beibl.:57. 1899,nom. nud.Material.Brazil.Ule289,
433, 636 (H-BR).
CampylopustortilisetusUle in C. Miiller, Bull. Herb.
Boissier6: 34. 1848. nom. nud. cf. Index Muscorum.
Dicranumexile C. Miller in Kindberg,Enum. Bryin.
exot. 55. 1888, nom. nud. cf. Index Muscorum.
Plants in soft, yellowish green, glossy tufts.
Stems 1-3 cm high, erect, sparsely foliate with
appressed leaves, tomentose, comose foliate at
tips. Leaves about 5 mm long, from ovate base
contracted into a narrow lanceolate tip. Perichaetial leaves with broader, sheathing base. Leaf
tip chanelled. Costa filling 1/3 of the leaf width,

excurrentin a long, toothed tip, in transverse
sectionwith ventralhyalocystsand dorsalgroups
of stereids, ridged at back. Alar cells large, inflated,hyalineto reddishto nearlylacking.Basal
laminal cells thin-walled,shortlyrectangular,610 x 19-38 ,um, narrowerat margins. Upper
laminal cells small, subquadrate,5-10 x 6-15
jim, incrassate.
Seta 5-6 mm long, curved, yellowish, brownish in age. Capsule 1-1.5 mm long, erect, symmetric, furrowedwhen empty, brownishto dark
brown. Operculumhalf as long as the capsule,
reddishbrown, obliquely rostrate.
Distribution(Fig. 97). On wet sandy or peaty
soil, on stumpsof treesand sandstonerocksfrom
sea level to 2000 m elevation in SE Brazil,Uruguay, Paraguayand Argentina.
SANTO:
Specimens examined: BRAZIL. ESPiHRTO
Reserva Florestal "Pedra Azul" E of Venda Nova,
Schafer- Verwimp8810 (hb. Frahm).GoIis: Km 130
along BR 040, mun. de Cristallina, Vital 6262 (SP).
Km 67 along road BR 135 between
MINASGERAIS:
Curveloand Diamantina,Frahm,Camp.Bras.Exsicc.
22 (ALTA,B, C, DUIS, FLAS,GOET,GZU, H, HBG,
INPA, JE, KR, M, MO, NFLD, NICH, PC, S, SP,
Vila Velha NW of Curitiba,
TENN, U). PARANA:
25?13'S,50?6'W,Vitt21458 (ALTA);5 km W of Jaguariaiva,24?14'S,49?45'W, Vital 10693 (SP); mun.
de Rio Negro,26?3'S,49?45'W,Vital9460 (SP);Guaratuba,Hatschbach38858 (hb. Frahm);ParqueNacional
Vila Velha, Vital 1777 (SP); 9 km E of Sao Joao do
Triunfo,Vital3288 (SP).Rio de Janeiro:Glaziou9093
(NY); Serrados Orgaos,PicadaSaudade,v. Liitzelburg
7044 (B); Teres6polis, Bandeira s.n. (RB); Serra do
Itatiaia,Viscondede Maua,Schafer- Verwimp8393 (hb.
Frahm);AbrigoReboucas,Schifer- Verwimp7564 (hb.
DOSUL:Mun. de Rio Grande,
Frahm).Rio GRANDE
32?15'S, 52?30'W, Vital 8987 (SP); Universitariode
Pelotascampus, Vital2032 (SP);Itapoa,PortoAlegre,
Sehnem 8173 (hb. Frahm);Taimbe, Sao Franciscode
Paula,Sehnem5378 (ASSL);mun. de Canela,29?22'S,
Serra do
50?47'W, Vital 9296 (SP); SANTACATARINA:
Espigao 11 km N of Moema, Frahm, Camp. Bras.

Exsicc. 20 (ALTA, B, C, DUIS, FLAS, GOET,GZU,
H, HBG, INPA, JE, KR, M, MO, NFLD, NICH, NY,
PC, S, SP, TENN, U); along road from LauroMiller
to Bom Jardin, Vital2009 (FLAS);Porto Belo, Praia
de Bombas, Yano 2433 (SP); N of Curitibanos,km
215.3 ofBR 116,26052'S,50?17'W,Vitt21033 (ALTA);
NW of Blumenau,37 km NW of Dr. Pedrinho,26?36'S,
49?45'W, Vitt20960 (ALTA);6 km E of Palma Sola
on road to Campo Ere, 26?24'S,53013'W, Vitt21242
(ALTA);km 175 of BR 282,6 km E of Xaxim, 26?55'S,
52?30'W,Vitt21170 (ALTA);along road SC 453 between Rio Bonito and Lebon Regis, Frahm 1489 (hb.
Frahm);Serrado Espigao 30 km NW Dr. Pedrinho,
Frahm 1527 (hb. Frahm);alongroadfrom Pinheiroto
SystematicTreatment
145
FIG. 99. Campylopusoerstedianus(Oersteds.n.,H-BR).A. Plant.B. Leaf.C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 jm.
CamposNovos, Frahm1576 (hb. Frahm);SerraGeral,

Ule 50 (NY);Tuberao,Ule 52 (NY). SAo PAULO:
Serra
do Bocaina,alongroadfrom Camposda Cunhato Sao
Jose, Frahm 1450 (hb. Frahm);km 128 along road
from Itanhaemto Peruibe,Frahm 1493 (hb. Frahm),
FLAS,GOET,GZU, H, HBG, INPA, JE,KR, M, MO,
NFLD, NICH, PC, S, SP, TENN, U).
PARAGUAY. CORDILLERA:
Tobati, Bordas 216
(NY). Guaira:Villarica,Bordas65B (NY). PARAGUARI:
ParqueNacionalYbycui,26?5'S,56?53'W,Buck12009,
12012A (NY).
Campylopuspyriformis
(Schultz)Bridelis similar to C. occultusbut has no comal tufts and has
rectangularinstead of quadrate upper laminal
cells. It is of southernhemispheric distribution
with disjunctoccurrencein W Europeand reaches north to SE Brazil (Rio de Janeiro), from
whence it has been describedas C. beyrichianus
Duby, C. erythrodontiusJaeger,C. subreconditus
(Geheeb & Hampe) Kindberg, and C. tijucae
Brotherusin Paris. It has the same range as C.
146
Flora Neotropica
Herb
7a
FIG. 100.
...
Campylopus oerstedianus (isolectotype, H-BR).
Plants in olive-green tufts, lighter above,

brownishbelow. Stems to 3 cm high, tomentose.
Leaves4-5 mm long, appressedto the stem when
dry, lanceolate, graduallynarrowedinto a subtubulosesubula.Costa filling /2of the leaf width,
excurrentin a short, mostly hyaline tip, in transverse section with small ventral hyalocysts and
dorsal groups of stereids, ridged at back. Alar
cells not or slightly differentiated.Basal laminal
cells hyaline, rectangular,8-13 x 45-60 um,
abruptly changing to the upper laminal cells,
44. Campylopusoerstedianus(C. Miiller) Mit- which are subquadrateto short rectangularor
ten, J. Linn. Soc. Bot. 12: 81. 1869. Dicranum oblique, 6-9 x 13-23 sjm.
oerstedianumC. Miiller,Bot. Zeit. 9(14): 259.
Sprophytenot known.
Distribution (Fig. 101). On soil and decom1851. Type. Costa Rica, "in regionealpina de
Irasu 11000'," Oersted s.n. (holotype, de- posed tussocks,from the lowlandsto the treeline
stroyed at B; lectotypus nov., NY; isolecto- in Costa Rica and Jamaica. Outside the neotypes, FH, H-BR).
Figs. 99, 100, 101. tropics in North Carolinaand Europe(southern
introflexus,whichalso goes fromthe subantarctic
to southernBrazil(see note under C. pilifer).
Campylopusacervatusand C. scabrellushave
equal prioritywith C. occultus.The latter name
has been chosen becauseit has been more commonly used whereas C. scabrellusand C. acervatus were known only from the type material.
Besides, neither type specimen represents the
most common expression of this species but
small, depauperateforms.
147
100
110
60
70
80
90
30
-?-
0*
200
0
400
100 200 300
6?0
800
10OO0km
-- I
400 500 600 mi
FIG. 101.
...
Distribution of Campy/opus oerstedianus (@) and C. pauper s. lat. (0).
Alps in Italy, Pyrenees, Vosges Mountains and

Massif Centralin France).
(surroundingsof the Caribbeanand Mediterranean Sea) and the recordsfrom the U.S.A. and
Europeare from localities, which are known for
Specimen examined. JAMAICA. ST. THOMAS Tertiaryrelicts.
PARISH:Blue Mtn. Peak, 18?3'N, 76?35'W, Crosby 3372
Campylopusoerstedianuscan easily be taken
(MO).
for a shade-grown C. pilifer, with accordingly
This species has a widespreadbut very scat- shorterhair-tip.It differs,however,in the ribbed
tered distributionand is known worldwidefrom and not lamellose back of the costa, the transonly a few localities, probablydue to the lack of verse section of the costa with relatively small
sporophytesand any special manner of vegeta- ventral hyalocysts,which are as largeas the metive propagation.Therefore,the existing records dian deuter cells (but larger in C. pilifer), the
can be interpretedas relicts from a originally dorsal groups of stereids, which consist of only
denser population. The total range is Tethyan 2 cells (about 4 cells in C. pilifer), the shorteror
Flora Neotropica
148
abruptlynarrowedinto a long, narrowawn. Costa filling 1/3of the leaf base in the appressedstem
leaves (narrowerin comal leaves), excurrentin
a serrateawn, in transversesection with ventral
hyalocystsand dorsalgroupsof stereids,smooth
at back. Alar cells reddish. Basal laminal cells
thick-walled,rectangular,sometimesslightlypitted, 5-8:1, 6-13 x 26-67 #m, those of the comal
leave thin-walled,narrowerand smallerat margins. Upper laminal cells subquadrateto short
rectangularor oblique, 5-10 x 13-26 Am.
Vegetative propagation(rarely)by deciduous
comal
leaves.
45. Campylopus pauper (Hampe) Mitten, J. Linn.
Seta
4-8 mm long, pale. Capsule1.5 mm long,
Soc. Bot. 12: 74. 1869.
light brown. Calyptrafringedor entire at base.
Distribution(Fig. 101). On soil, soil covered
rocks and rotten log in open places below and
Campylopuspauper.
above the tree line in elevations between 2200
and 3600 m in Mexico, Venezuela, Colombia,
1 Basal laminal cells rectangular;capsule oblongPeru and Bolivia.
var.
45a.
...........
strumose.
pauper.
cylindric,
even lacking hyaline hairpoint of the leaves and
shorter, subquadrate or short rectangular (not
oval) upper laminal cells. All these characters
concern "minor" differences and it can be supposed that both species have a phylogenetic relationship.
Several collections from Jamaica have been
named C. oerstedianus, which, however, with the
exception of one specimen, must be referred to
modifications of C. fragilis with hyaline excurrent costas, or to C. pilifer.
to subquad1 Basallaminal cells short-rectangular

Yerba
Specimens examined. MEXICO. CHIAPAS:
rate;capsule ovoid, not strumose. ............
45b. var. lamprodictyon. Buena, Pueblo Nuevo, about 55 km NE of Tuxtla,
......................
Sharpet al. 2898 (TENN).
COLOMBIA. ARAUCA:
45a. Campylopus pauper (Hampe) Mitten var. QuebradaEl Play6n, Cleef9026b, 9140 (U). BOYACA:
Figs. 5F, G, 101, 102. Paramode Pisva,careteraSocha-LaPunta,Cleef4624b
pauper
(U); Mun. de Sogamoso, Lagunade Tota, Aguirre&
DicranumpauperHampe,Linnaea32:137. 1863.Type. Rangel 464 (FLAS).CAUCA:
ParqueNacional de PuParamo de SuColombia. Manzanos,Lindig 2013 (holotype,n.v.; race, Cleef 4990 (U). CUNDINAMARCA:
mapaz, Cleef8484 (U); CarreteraS. Cayetano-C6gua,
isotypes, H-BR, H-SOL, GOET,JE, NY).
Campylopuschrysodictyon(Hampe) Mitten, J. Linn. Cleef 6633 (U); Piramo entre C6guay San Cayetano,
Soc. Bot. 12: 78. 1869. Type. Colombia. Bogota, Cleef4900 (U); Paramode CruzVerde,Cleef3019 (U);
Tequendama,Lindig s.n. (holotype, n.v.; isotypes, San Crist6balnearBogota,Apollinaires.n. (PC);above
DECAUCA:
Farallones
Bogota, Troll2081 (PC).VALLE
H-BR, H-SOL, GOET, NY, S).
Campylopusfendleri (C. Miller) Kindberg, Enum. de Cali, 3022'N,76043'W,Hartman 56634 (COL).
VENEZUELA.MERIDA:
Bryin.exot. 88. 1889. DicranumfendleriC. Miller,
Linnaea42: 472. 1879. Type. Venezuela.Fendler41 Paramode Mucubaji,Griffinet al. 1076 (FLAS);Paramo de La Negra entre los limites entre los estados
(holotype,destroyedat B; isotypes, H-SOL,NY).
CampylopuskingiiRobins.,Bryologist70(1): 18. 1967. Tachiray Merida,Ruiz-T. et al. 8402b (FLAS);above
Type. Colombia.King& GuevaraC-1132 (holotype, Bailadores,Griffinet al. 2114, 2140 (FLAS);Paramo
de Don pedroentreEl Morroy Aricagua,Ruiz &Lopez
US; isotype, COLO).
Campylopuspropinquus(Hampe)Mitten,J. Linn.Soc. 9575 (FLAS);SierraNevada de SantoDomingo, ParaBot. 12:75.1869. Dicranumpropiquum
Hampe,Ann. mo de Los Granates,Lopez-F. 15427 (FLAS);along
Sci. Nat. Bot. ser. 5, 3: 367. 1865. Type. Colombia. the trailof the telefericobetweenAguadaand La MonParamoEl
Lindig2119 (holotype,n.v.; isotypes,H-BR, H-SOL, tafia, Griffinet al. 1786 (FLAS).TACHIRA:
RosalbetweenLa GritaandSanJosede Bolivar,Griffin
GOET,JE, NY).
Campylopusrosulatus(Hampe) Mitten, J. Linn. Soc. et al. 693 (FLAS).TRUJILLO:Paramode Guaramacal
Bot. 12:79.1869. DicranumrosulatumHampe,Lin- above Bocon6, Griffinet al. 1051 (FLAS);arribade la
naea 32: 139. 1863. Type. Colombia. Bogota, La ColoniaT6var,Ruiz &Lopez 10160 (FLAS);trailfrom
Penna et Pacho, Lindig 2012 (holotype, n.v.; iso- Humocaroto Buenos Aires, 9?40'N, 70?5'W,Liesner
et al. 8076 (MO).
types, H-BR, GOET, NY).
Km 403.5 along road from CaPERU. AMAZONAS:
Plants to 5 cm high, yellowish-green, in loose jamarcato Chachapoyas,Frahm,Camp.Peruv.Exsicc.
tufts. Stems erect, appressed foliate below, end- 21 (ALTA, B, BM, BUF, DUKE, EGR, F, FLAS, G,
GRO, GZU, HBG, KR, NAM, NICH, NY, MEXU,
ing in a comal tuft with erect spreading leaves,
NFLD, PC, POZ, RNG, S, U); Las Palmas near Leimm
ca.
6-7
radiculose
at
base.
Leaves
slightly
mebamba,Hegewald6971 (hb. Hegewald).JuriN: Belong, the lower stem leaves narrow lanceolate,
tweenHuatapallanaandChilifruta,Hegewald9245 (hb.
the comal leaves with broad sheathing base Hegewald).
149
4.5rmn
FIG. 102. Campylopuspauper var. pauper (Cleef 8484, U). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
details = 50 um.
BOLIVIA. Near Yungas,Rusbys.n. (NY). COCHA- 5 cm) and entire or ciliate calyptras. This all
BAMBA:Antahnacana 7 miles SSW of Locotal, 48 miles
NNE of Cochabamba,Hermann 25240 (F); Pampa

Tambo, Hermann 25247 (F); Incacorral,Herzog 279
(JE).
This species has been describedfourtimes under differentnames by Hampe aftermaterialcollected by Lindig in the surroundingsof Bogota.
The type specimens differ slightly by the pitted
or not pitted basal laminal cells and the shape
of the capsule, size of the plants (between 1 and
seems, however, to fall into the variation of one

speciesas all these charactersvary independently
of each other.
Also specimenscollectedby Lindigin the same
area as Campylopus chionophilus (C. Muller)
Mitten belong to this species. The type of C.
chionophilus,however, belongs to Dicranodontium denudatum.
Robinson (1967) cited C. brachyphyllulus
Wilson ex Mitten as synonymouswith C. rosulatus.
Flora Neotropica
150
eF~~I
V/
\?_\
5fw
Ds
FIG. 103. Campylopusperexilis (Cleef 7116, U). A. Plant. B. Leaf. C. Leaf-tip. D. Transversesection of
leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails = 50 Am.
However, this species is placed here as a synonym of C. flexuosus.
46. 1939.Type.Bolivia.Yungasvon La Paz, Troll

111 (holotype,PC;isotype,B).
Differsfromvar.pauperespeciallyby the short

basal laminal cells, which are only 1-2 times
longeras broad.The capsuleis shorterand ovoid,
Campylopuslamprodictyon(Hampe) Mitten, J. Linn. not strumose,the calyptrais entire at base.
Soc. Bot. 12: 78. 1869. Dicranum lamprodictyon
Distribution(Fig. 101). In similar habitats as
Hampe,Ann. Sc.Nat. Bot. ser. 5, 5: 339. 1866. Type. var.
pauper in Mexico, Costa Rica, Venezuela
Colombia. Bogota,La Penna,Lindig s.n. (holotype,
and
Colombia.
The collections from Mexico are
BM; isotype, S).
Campylopustrollii Theriot, Rev. Bryol. Lichenol. 11: from lower altitudes, 1100-2000 m.
45b. Campylopus pauper var. lamprodictyon

(Hampe) J.-P. Frahm, Bryol. Beitr. 7: 58. 1987.
151
Specimens examined. MEXICO. CHIAPAS:Barrio of
Banabi, Mun. de Tenejapa, Breedlove 15351 (MICH);
near San Crist6bal de las Casas, Arzeni 30b (PC). JALISCO:W of Autlan on Hwy. 80, Sharp et al. 2728
(TENN). PUEBLA:Alrededores de Xicotepec de Juarez,
EntreSantaRita
Delgadillo1390 (MEXU);VERACRUZ:
y Yecuatla,Delgadillo 1017 (MEXU).
Km 73 on InterAmerican
COSTARICA.CARTAGO:
q
70
e0o
60
HighwaySEof ElEmpalme,9?37'N,83?50'W,Crosby
6351 (MO).
Paramode La Canada,
VENEZUELA.
TRUJILLO:
Ruiz-T.9193b(FLAS).
Paramode LaCristalina,
SanCarlosbeiMapiri,Buchtien282 (B).
BOLIVIA.
The taxonomic position of this taxon is not

clear. It has been described from material collected by Lindig in the same area as C. pauper
and threemore synonymsand occursin the same
rangeas var. pauper.It can be recognizedby the
short basal laminal cells. This characteris combined with short, ovoid, not strumose capsules.
Such capsules, however, are also found in C.
chrysodictyon, one of the synonyms of var. pau-
per, but are correlatedin this species with long

rectangular,pittedbasallaminalcells. Therefore,
it can not be excludedthat the basallaminalcells
may also vary and that var. lamprodictyonhas
to be includedin C. pauperwithout infraspecific
rank.
With respect to the short basal laminal cells
and also the presenceof comal tufted stems, C.
pauper var. lamprodictyon much resembles C.
zygodonticarpus, which has, however, erect,

symmetric capsules, whereas C. pauper has
asymmetric,inclined capsules.
46. Campylopusperexilis(C. Miiller)Paris,Ind.
bryol. Suppl. 95. 1900. Dicranum perexile C.
Miller, Nuov. Giorn. Bot. Ital. n. ser. 4: 34.

1897. Type. Bolivia. Cochabamba:Germain
1006 (holotype,destroyedat B;lectotypusnov.,
H-BR; isotypes, JE, NY).
Figs. 103, 104.
Plantssmall,to 1 cm high, in light-green,dense
tufts. Stems equally foliate with erect patent
leaves. Leaves 3-4 mm long, lanceolate, narrowedto a tubulose,relativelyblunt apex, which
is finelyserrateat tip. Costa fillinghalf of the leaf
width, in transverse section with large ventral
haylocysts and dorsal stereids, smooth at back,
endingin the leaf tip. Basallaminalcells hyaline,
thin-walled,rectangular,8-13 x 38-58 um. Upper laminal cells subquadrateto short rectangular(1:1.5-2), 5-10 x 13-25 tsm, incrassate.
200
400
600
800 10OOkm-
0 100 200 300 400 500 600
FIG. 104. Distributionof Campylopusperexilis.

Seta 4 mm long, curved. Capsule 1 mm long,
ovoid, symmetric. Calyptra fringed at base.
Distribution (Fig. 104). On soil and rocks in
the alpine belt of the Andes, rarely below in open
habitats, in elevations between 2700 and 4100
m in Colombia, Peru and Bolivia.
ParaSpecimens examined. COLOMBIA. BOYACA:
mo de la Rusia, Cleef7116 (U).

PERU. ANCASH:
CordilleraBlanca, LagunillasPatoccocha, Hegewald 7640 (hb. Hegewald); Laguna
Querococha, Hegewald 7674 (hb. Hegewald);Jutopampa near Catac, Hegewald 7760 (hb. Hegewald).
HUANCAVELICA:
Ichoquenus, Hegewald 9156 (hb.
Hegewald). JUNIN: Pampa Huicushcancha near Tarma, Hegewald 6269 (hb. Hegewald). LA LIBERTAD:
HuancamarcanearOtuzco,Hegewald5129 (hb. Hegewald); LagunaSausacochanear Huamachuco,Hegewald 6035 (hb. Hegewald).

This is a critical species, known only from a
few localities. It is similar to another Andean
species C. nivalis var. C. multicapsularis, but
characterized by a short, canaliculate leaf tip.
The upper laminal cells, which are more or less
subquadrate, resemble those of C. nivalis var.
multicapsularis, and therefore it can not be excluded that C. perexilis may ultimately be a mod-
Flora Neotropica
152
ification of the latter-drier habitats causing a
shorter leaf tip. This can, however, be decided
only by field studies. In these characters it resembles, furthermore, C. subulatus Milde, a species, which is known from scattered localities in
W Europe and single localities in California and
India, occurring however, at low altitudes.
dot,Bull.Soc.Roy.Bot.Belgique31(1):149. 1893.
(n.v.).
andabout30 moresynonymsfromotherpartsof the

rangeof C. pilifer.Synonymyof taxaof whichthe
typeshavenot beenseenaccordingto Wijket al.,
1959.
Plants in rigid, green to olive-greentufts 0.55 cm high, sparselyreddish tomentose, equally

foliate, only fertile plants forming comose tufts
47. Campylopuspilifer Bridel, Mant. Musc. 72. of perichaetialleaves. Leaves 5-7 mm long, lan1819. Type. Italy.Ischia,Bridels. n. (lectotype, ceolate,graduallyacuminate,subtubuloseabove.
B).
Costa filling 1/2-2/3of the leaf base, excurrent in a
hyaline, serratehairpoint, in transversesection
with ventralhyalocystsand dorsalgroupsof narKey to the Infraspecific Taxa of
row stereids,lamellose at back with lamellae 3Campylopus pilifer
4 cells high. Alarcells not or scarcely,rarelywell
1 Plantswith ventralhyalocysts,usuallyolive green.
differentiated.Basal laminal cells hyaline, thin..............................
47a. ssp. pilifer.
walled,long-rectangular,8-13 x 60-70 ,m, nar2 Dorsal lamellae of the costa consistingof 3-4
cells. .......................
47a. var. pilifer. rower at margins. Upper laminal cells shortly
2 Dorsal lamellae of the costa consistingof 5-6
oval to oblong-rhombic,9-16 x 7-8 ,m.
cells. ...................
47b. var. lamellatus.
Seta 3-5 mm long. Capsules 1.5 mm long,
1 Plants with ventralsubstereids,usuallyblackish.
brownish. Calyptrafringedat base.
ovoid,
........................
47c. ssp. galapagensis.
Distribution(Fig. 107). On exposed rocks and

boulders,
dry soil on banks of roads and trails,
47a. Campylopus pilifer ssp. pilifer var.
with a broad ecological amplitude from semi105-107.
pilifer.
Figs. 3,
deserts to rain forests and an enormous altituCampylopusangustifoliusWarnstorf,Hedwigia57:81. dinal rangefrom sea level to 4800 m, widespread
1915. Type. Brazil.Sao Paulo, Usteris.n. (holotype, and
frequentthroughthe Neotropicsfrom MexB).
ico
Guatemala,Honduras,El Salvador,
through
Campylopuscinchonae Paris, Ind. bryol. Suppl. 90.
1900. Thysanomitrionjamaicense C. Miiller, Bull. Nicaragua,Costa Rica, Panama, Cuba, HispanHerb. Boissier5: 552. 1897. Type. Jamaica. Wilson iola, Jamaica,PuertoRico, Colombia,Venezue813 (n.v.).
la, the Guianas, Ecuador,Peru, Bolivia, Brazil
Campylopuserectus(C. Miiller)Mitten, J. Linn. Soc. to northern
Argentina.Also in North America
Bot. 12:89. 1869.DicranumerectumC. Miiller,Syn.
musc.frond.1:408.1849. Type.Colombia.Galipan, (Arizonato SEUnited States),W Europe,Africa,
India and Sri Lanka. (WorldwidedistributionMoritz 145 (n.v.).
Campylopusliebmannii(C. Miiller)Schimperin Be- map, see Frahm, 1984a.)
scherelle,M6m. Soc. Nat. Cherbourg16: 167. 1872.
DicranumliebmanniiC. Miiller, Syn. musc. frond.
2: 601. 1851. Pilopogon liebmannii (C. Muller)
Brotherus,Nat. Pflanzenfam.1(3):336. 1901. Type.
Mexico. Hahn s.n. (holotype, destroyed at B; lectotypus nov., H-BR).
CampylopusluridusSchimper ex Bescherelle,Mem.
Soc. Nat. Cherbourg16: 167. 1872. Type. Mexico.
Orizaba,F. Miller s.n. (n.v.).
Campylopuspilosissimus Schimper ex Bescherelle,
Mem. Soc. Nat. Cherbourg16: 167. 1872. Type.
Mexico. Mirador,Sartoriuss.n. (n.v.).
Campylopusproliferus
(C. Miller) Mitten,J. Linn.Soc.
Bot. 12: 85. 1869. DicranumproliferumC. Miiller,
Syn. musc. frond. 2: 602. 1851. Type. Venezuela.
T6var, Wagners.n. (n.v.).
CampylopusstrictusSchimper ex Bescherelle,M6m.
Soc. Nat. Cherbourg16: 167. 1872. Type. M6xico.
Rio de Orizaba,F. Miillers.n. (n.v.).
C. Miller ex Renauld& CarCampylopussubproliferus
Selected specimensexamined.MEXICO. CHIAPAS:
BenitoJuarez,30 km S of Tapachula,
Frahm,Camp.
Exsicc.12 (B, BING,BUF,C, DUIS,DUKE,EGR,
F, FLAS,G, GOET,GRO,GZU,H, HBG,JE,KRA,
M, MEXU,MO, NAM, NFLD,NICH,NY, PRC,
Puerto de Rio
PRE, POZ, RNG, S, U). CHIHUAHUA:
Urique, Bowers et al. 5373a (TENN); on road from
Creel to La Junta, Bowers et al. 5397a (TENN).
DURANGO: 4 miles W of LaCiudad,Bowerset al. 5271a
Above Real del Monte, Sharp534
(TENN).HIDALGO:
(TENN); 11 km NE de Acaxochitlfn,Delgadillo 1362
NearPuertolos MazosW of Au(MEXU).JALISCO:

tlfn, Sharpet al. 2696 (TENN);MExico:7 km E de
Amecameca,Cdrdenas3615 (MEXU);BotanicalGarden of Mexico City, Schwab s.n. (hb. Frahm);pass

along road F15 betweenMexico City and Toluca,Eggers & Frahm 792028 (hb. Frahm);Volcan Paricutin,
near Uruapan,Rejmanek 12,13 (PRC). MICHOACAN:
28 km W de CiudadHidalgo,Delgadillo446 (MEXU);
153
?:;~~~~~~~
M,c
i .. ,,..
, .?'i
?
FIG. 105. Campylopuspilifer ssp. pilifer(Crosby7216, MO). A. Plant. B. Leaf. C. Leaf-tip.D. Transverse
details = 50 Am.
25 km E of Morelia,Iwatsuki& Sharp2808 (TENN);
41 km W of Cd. Hidalgo, Sharp et al. 2189 (TENN);
PuertoMorelos 53 km E of Morelia,Iwatsuki& Sharp
3034 (TENN); along Hwy. 15 near Zacapu,Norris &
Taranto15462 (TENN). MoRLOs: Sierrade Chichinautzin,mun. de Tepoztian,Cdrdenas1057b(MEXU);
betweenTres Cumbresand Cuernavaca,Dull s.n. (hb.
Dill); Cuernavaca,Pringle 10505 (JE).NAYARIT:4 mi
E of La Cienaga,Norris & Taranto14298 (TENN); 3
miles W of Mesa del Nayar, Norris& Taranto14658
(TENN); near LabraW of Jesus Maria,Norris& Taranto 14115 (TENN);along road to Jalcacatan,Norris
& Taranto 13192 (TENN). OAXACA:

Km 146 along
road F 190 from Tehuantepecto Oaxaca, Eggers &
Frahm 792031 (hb. Frahm);2 km E of Ixtlan, Delgadillo 713 (MEXU);km 140 along highwayF 190 N
of Oaxaca,Frahm, Camp.Exsicc. 11 (B, BING, BUF,
C, DUIS, DUKE, EGR, F, FLAS, G, GOET, GRO,
GZU, H, HBG, JE, KRA, M, MEXU, MO, NAM,
NFLD, NICH, NY, PRC, PRE, POZ, RNG, S, U,
UPS); SierraJuarezgap on Hwy. 175, Websteret al.
542 (TENN);near Nochixtlan 65 km NW of Oaxaca,
Websteret al. 406 (TENN);40 mi NW of Oaxacaon
Hwy. 190, Smith et al. 470A, 477 (TENN); 1 mi W of
154
Flora Neotropica
et al. 2823, 2826 (TENN). ZACATECAS:
7 km S de la
Laguna Grande, 22?27'N, 103?33'W, Cdrdenas 3023
(MEXU).
HONDURAS. MORAzAN:El Hatillo 10 km NE of
Tegucigalpa,Pilz 1421 (MO).
..
....
C:
EL SALVADOR. Volcan de San Vicente, Winkler

s.n. (hb. Winkler); Cerro El Pilon near Los Naranjos,
Winkler s.n. (hb. Winkler). CHALATENANGO:
Cayaguanca, San Ignacio, Winkler 20 (MO). SANTAANA:
Volcan de S. Ana, Winkler 29 (MO).
NICARAGUA. MASAYA:Parque Nacional Volcan
Masaya, Stevens 5289 (MO).
Km 80 along Pan AmerCOSTA RICA. ALAJUELA:
ican Highway SW of San Ramon, Crosby 3753 (MO).
PUNTARENAS:Quaker settlement of Monte Verde,
Crosby2531b (MO). SANJOSE:17 km SE of El Em-
..... '..3* n__,

g.~c*.
r^_ ;.,"^ ^ *"
palme, 9?37'N, 83?48'W, Crosby 9759 (MO); La Asun-
' ,': *:..:
cion, Cima del Cerrode la Muerte,Delgadillo2962b

(MEXU); Cordillera de Talamanca, Valle des los Conejos, Kuhbier 616 (BREM).
PANAMA. CHImQUI:9 km WNW of Boquete, Da-
r-ab-/ta
vidse& D'Arcy10330 (MO).
*: 3
'a
.'
'
..*
e,
:'.'
CUBA. La Gran Piedra, Buck 7635 (NY).

JAMAICA. ST. THOMASPARISH: Between Portland
gap and Blue Mountain Peak, Hegewald 8103 (hb.
Frahm); Sir John's Peak, Crosby 3046 (MO).
DOMINICAN REPUBLIC. LA VGA: 40 km S of
Constanza, 18?45'N, 70?37'W, Reese 15783 (NY).
GUYANA. Upper Mazaruni District, N slope of Mt.
Roraima, 5017'N, 60?43'W, Gradstein 5293 (U).
COLOMBIA. ANTIOQUA: 10.5 km E de Sons6n,
'
5?40'N, 75?15'W,Churchill& Sastre-De Jesuis13068
(NY); 25 km NE de Medellin hacia Santa Fe, 6?20'N,
75?40'W,Sastre-De Jesus et al. 1464 (NY); 17.6 km
N de Copacabana, 6?25'N, 75?25'W, Sastre-De Jesius

& Churchill 942 (NY); 14.5 km SE de Medellin, 6?10'N,
75?30'W,Sastre-DeJestis & Churchill889 (NY); Rio
Negro, ferme San Cayetano, Onraedt 9927 (hb. Frahm);

Medellin-Sons6n road, 5-6 km SE of La Union, Lu-
teyn 7111 (NY). ARAUCA: SierraNevada del Cocuy,
'
i^.?-:
6:?.
./
j
'1
Paramo de Sumapaz,
Cleef9181 (U). CUNDINAMARCA:
Cleef4935A (U). NARI&O:Mun. Picaurte, La Planada,
Luteyn6837 (NY).
VENEZUELA.
FIG. 106. Campylopuspilifer(lectotype,B).

Rosa Blanca,Norris& Taranto 16732 (TENN); trail
above Tamazulapamnear Zempoatepetl,70 km E of
Oaxaca, Sharp et al. 4543 (TENN); 22 miles N of
Pochutla on Hwy. 175, Norris & Taranto 15969
(TENN);67 kmN ofIxtlande Juarez,Norris& Taranto
16195 (TENN);nearS. Pablo Ayutla, 70 km E of OaNearOcostoc
xaca,Sharpet al. 4133 (TENN).PUEBLA:
below Tezuitlan,Sharp 3976 (MEXU). VERACRUZ:
5
mi W of Mirador,Sharp89220 (TENN);cercade Pacho
al N de Coatepec,19?29'N,96?54'W,Juarez304 (NY);
4 miles W of Jalapa,Norris& Taranto16848 (TENN);
Pedrigalde las Vigasabove La JoyanearJalapa,Sharp
ARAGUA: Parque Nacional Henry
Pittier.MarquezMeza 20 (NY). MERDA: Road to La

Azulita(LaCarbonera),
Fransen1171, 1203 (GB);along
Merida-LaAzulitaroad,Griffinet al. 92 (FLAS);Parque
Nacional SierraNevada, around La Aguada,Fransen
1416 (GB);Mucuy, Onraedt5586 (hb. Frahm).
ECUADOR. COTOPAXI:
Quevedo-Latacunga road,
79"9'W,0?52'S, Holm-Nielsen et al. 3146 (GB); PiCHINCHA: Carretera Nono-Nanegalito, Balslev 2491
(NY, QCA). TUNGURAHUA: 3 km W of Banos, 1?24'S,
78026'W, Churchill& Sastre-De Jesfs 13739 (NY).
GALAPAGOS
ISLANDS:Isla San Crist6bal, Gradstein &
Weber 118 (U); Isla de Santa Cruz, Weber 13673 (COL);
Chatham Island, Weber 14280 (COL).
PERU. AMAZONAS:
between Balsas and Leimebamba, Frahm 823989 (hb.
Frahm). AYACUCHO:
Ayna, Hegewald 9073 (hb. Hegewald). CAJAMARCA:Quebrada Cavilan, Hegewald6546
(hb. Hegewald). Cuzco: Aguascalientes, Hegewald8778
(hb. Hegewald); Machu Picchu, 2400 m, Frahm, Camp.
"O
10
155
13;
90
80
70
60
*C
so
-------- -------- .
O 100200 00 4050 eOOl6s
.1
.'
y %
<
!i
RRpkCym.
Prpared
be, Hendrl0..
,f ,
FIG. 107. Distributionof Campylopuspilifers. lat.

BR 135, Mun.de OuroPreto,Frahm1438 (hb.Frahm);
Serra do Cabral, Mun. de Joaquim Felicio, 17?44'S,
44?12'W, Vital 7898 (SP); Serrado Caraca,Mun. de
SantaBarbara,20?7'S,43?31'W, Vital7664 (SP);along
road Belo Horizonte-Juiz de Fora, Vital 6502 (SP);
alongBR 116, Mun.de PadreParaiso, Vital5896 (SP).
MATOGROSSO:
28 km S de Guiratinga, 16?27'S,
53?47'W, Vital 9879 (SP). Rio de Janeiro: Serra do
Mar between Paratiand Cunha,Frahm, Camp. Bras.
Exsicc. 9 (ALTA, B, C, DUIS, FLAS, GOET, GZU,
LA LIBERTAD:
Sausacocha near Huamachuco, Hegewald 6023 (hb. Hegewald);Otuzco, Hegewald 7223 H, HBG, INPA, JE, KR, M, MO, NFLD, NICH, PC,
Lomasde Lachay,Lowy175 (LAF). S, SP, TENN, U); Teres6polis,ParqueNacional Serra
(hb. Frahm).LIMA:
PUNo:7 km E of pass La Raya, Hegewald5498 (hb. dos Orgaos,Frahm, Camp.Bras. Exsicc. 7 (ALTA, B,
C, DUIS, FLAS, GOET, GZU, H, HBG, INPA, JE,
Hegewald).
BRAZIL. ALAGOAS:Mun. de Monte Alegre, Vital KR, M, MO, NFLD, NICH, PC, S, SP, TENN, U);
1967 (SP). BAHIA:Mun. de Leucois, Boom & Mops ParqueNacionalItatiaia,Frahm,Camp.Exsicc. 42 (B,
1079 (NY). MINASGERAIS:Along road MG 2 between
BING,BM,BUF, C, DUKE, EGR,F, FLAS,G, GOET,
Serroand Datas S ofDiamantina, Frahm,Camp.Bras. GRO, GZU, H, HBG, JE, KRA, M, MEXU, NAM,
Exsicc. 8 (ALTA, B, C, DUIS, FLAS, GOET, GZU, NFLD, NICH, NY, PC, POZ,PRC, RNG, S, U, UPS).
H, HBG, INPA, JE, KR, M, MO, NFLD, NICH, PC, Corcovado,Frahm, Camp.Bras. Exsicc. 6 (ALTA, B,
S, SP, TENN, U); along MG 2 N of Conceiaode Mato C, DUIS, FLAS, GOET, GZU, H, HBG, INPA, JE,
Dentro, Frahm 1397 (hb. Frahm);km 392 along road KR, M, MO NFLD, NICH, PC, S, SP, TENN, U);
Exsicc. 56 (ALTA, B, BING, BM, BUF, C, DUKE,
EGR,F, FLAS,G, GRO,GZU, H, HBG,KRA,MEXU,
NAM, NFLD, NICH, NY, PC, PRC, POZ, RNG, S,
U); Vargas14398 (MICH).HUANUco:
Alongroadfrom
Huanucoto Huaraz,Frahm, Camp. Peruv.Exsicc. 17
(ALTA,B, BM, BUF, DUKE, EGR,F, FLAS,G, GRO,
GZU, HBG, KR, NAM, NICH, NY, MEXU, NFLD,
PC, POZ, RNG, S, U). JUNiN: Huasahuasi,Hegewald
8354 (hb. Hegewald);Carpapata,Kunkel1076 (MICH).
Flora Neotropica
156
AbrigoReboucas,Griffin& Vital28 (FLAS).Rio
P. laevisor P. peruvianusor even with P. gracilis,
GRANDE
DOSUL:Mun. de Canela, 29?22'S,50?47'W, which rarely may also have
hyaline excurrent
Summit of SerraParimaS
Vital9293 (SP). RORAIMA:
Near costas. Leaf shape and areolationare much the
of Anari,Pranceet al. 2158 (INPA).SAOPAULO:
Santos,Rose&Russells.n.(NY);Mun.deBrotas,Vital same and can not be distinguished.The only way

to Brotas,Vital to differentiatethese species is by a transverse
1757(SP);alongroadfromItarapina
2072 (SP).
section of the costa, which shows ventralstereids
Vic. Yungasand La Paz, in
BOLIVIA.COCHABAMBA:
Pilopogon but ventral hyalocysts in CampyJay s.n. (NY). CerroPampalanga,
Herzogs.n. (JE);
lopus.
TresCruces,Herzog3927(JE).
Until 1955, when Giacomini (1955) recognized both taxa as separatespecies, Campylopus
pilifer generally was included in C. introflexus
(Hedwig)Bridel.However,the distributiongiven
for both taxa was incorrect and corrected by
Frahm(1974). Gradsteinand Sipman(1978) introduced the older name C. pilifer for C. polytrichoidesDe Notaris as used by previous authors.
Campylopusintroflexusis a southern hemispheric,subantarcticto subtropicalspecies,which
has been introducedin Europesince 1942, where
it is spreadingwidely, and found in California
and Oregon since 1972. This species differsby
reflexed, not straighthairpoints, and dorsal lamellae of the costa only 1-2 cells high. Campylopusintroflexusis found also in the southern
statesof Brazil.It is very frequentin Rio Grande
do Sul and reachesnorththroughSantaCatarina
and Paranaespeciallyalongthe coast to the state
of Sao Paulo. Campylopuspilifercan be regarded
as a vicariantspeciesof C. introflexusin the tropics, adaptedto eitherdryhabitatsby longercostal
lamellae for water-storageor even rain forest
habitatsby even longerlamellaefor a bettergasexchange.The close relationshipof both species
is shown by the occurrence of hybrids in the
overlappingpartsof their ranges,as in Northern
Argentina. These hybrids show the erect hairpoints of C. pilifer but the low lamellae of C.
introflexus.
In contrast to the wide ecological and geographicalrange, C. pilifer is not especially variable morphologically.A variablecharacteris the
presence of alar cells, lacking in moist habitats
(as rain forests), where this species can take up
wateras rain or mist throughthe leaves, or, conversely, well differentiatedin wet, but exposed
habitats such as open, inundated rocks, where
the alar cells function for water transportfrom
the substrate.
Sterile specimens can easily be confused with
hyaline hair-tippedspecies of Pilopogonsuch as
47b. Campylopuspilifer ssp. pilifer var. lamellatus (Montagne)Gradstein& Sipman,Bryologist 81: 119. 1978. Campylopuslamellatus
Montagne, Ann. Sci. Nat. Bot. ser. 2(9): 52.
1838. Type. Bolivia (holotype, PC; isotype,
BM).
Differsfrom var.piliferby lamellaeat the dorsal side of the costa, which are 5-6(-7) cells high.
Furthermorethe ventralhyalocystsin transverse
section of the costa may be smaller as in var.
pilifer, which needs more detailed studies.
Distribution(Fig. 107). On soil and soil-covered rocks in rain forests in elevations between
2000 and 3500 m, recorded from Costa Rica,
Ecuador,Peru and Bolivia, also in tropical Africa, from where it has been describedas C. introflexusvar. altecristatusRenauld & Cardot.
La
Specimens examined. COSTA RICA. CARTAGO:
Estrella,Standleys.n. (JE).
ECUADOR.Zaruma,Espinosa s.n. (JE).
PERU. AYACUCHO:
Along road from Huantato San
Frahm,Camp.
Francisco,15 kmfromSanFrancisco,
Peruv.Exsicc.18 (ALTA,B, BM,BUF,DUKE,EGR,
F, FLAS,G, GRO,GZU,HBG,KR, NAM,NICH,
NY, MEXU,NFLD,PC,POZ,RNG,S, U).
47c. Campylopuspilifer ssp. galapagensis(J.-P.
Frahm & Sipman) J.-P. Frahm, Bryol. Beitr.
7: 60. 1987. Campylopusgalapagensis J.-P.
Frahm& Sipman,J. Bryol.10:61. 1978. Type.
Ecuador.GalapagosIslands:SantaCruz,base
of Mt. Crocker,Gradstein& Weber17 (holotype, U; isotypes, CDRS, COLO, QCA).
Figs. 107, 108.
Plants greenishor usually blackish. Stems 12 cm high. Leaves 2-3.5 mm long, lanceolate,
appressedwhendry,subtubularabove. Costafilling 1/2of the leaf width, longly excurrentin a
hyaline, serratehairpoint, in transversesection
with ventral substereids,a dorsal band of stereids, with lamellae 2-3 cells high. Basal laminal
cells hyaline, 40-60 x 7-15 ,im, narrowerat
157
FIG. 108. Campylopus pilifer ssp. galapagensis (Gradstein & Weber 17, U). A. Plant. B. Leaf. C. Leaf-tip.
D. Transverse section of leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule. Unlabeled scale bar for
microscopic details = 50 jim.
margins. Upper laminal cells elongated rhomboid, 10-20 x 8-10 ,m, sharply differentiated
from the basal ones.
Seta (3-)4(-6) mm long, sinuose. Capsule 11.5 mm long.
Differs from C. pilifer ssp. pilifer mainly by
ventral substereidsin transversesection of the
costa, shorterleaves and a blackish coloration.
GalapagosIslands:Floreana,Isabela,Pinta, Pinz6n, Santiago,San Crist6baland Santa Cruz.
Specimens examined: ECUADOR. GALAPAGOS

ISLANDS:
Isla Santa Cruz, Weber B 13654, 13573
(COLO);van der Werff 1378, 1476 (U); Gradstein,
Bryoph. Neotrop. Exsicc. 4 (U); Floreana, Wittmer
Farm, Gradstein,Bryoph.Neotrop.Exsicc. 5 (U); Isla
Santiago,Pike 41006 (COLO).
The ssp. galapagensis is a "micro-endemic"
of the Galapagos Islands. Conspicuously, ssp. pilifer also occurs on these islands, but in contrast
to this subspecies, ssp. galapagensis inhabits dryer habitats. It can therefore be supposed that
158
Flora Neotropica
0lll
x ;F
\X
;I
11X}0
FIG. 109. Campylopuspittieri(Lajtnant& Molau 13706, NY). A. Plant.B. Leaf.C. Leaf-tip.D. Transverse
section of leaf. E. Basallaminal cells. F. Upper laminal cells. G. Capsule.Unlabeled scale bar for microscopic
= 50
details=
50ymLr.
details
;:m.
Campylopuspiliferhas invaded these young volcanic islands and has, as an adaptation to the
dry volcanic lava flows, developed ventral substereids as a protectionagainstdesiccation.
Vegetativelyit much resembles,in color, presence of ventral substereidsand dorsal lamellae,
Pilopogonperuvianus.The taxa can be separated
clearly only by sporophyticcharacters.A differentiation of sterile materialseems possible only
by the outermostcell walls of the lamellae,which

are equally thickened in C. pilifer ssp. galapagensis but have distinctly thickerterminal than
lateralcell walls in Pilopogonperuvianus.
48. Campylopuspittieri Williams, Bull. Torrey
Bot. Club34: 569. 1908. Type. Colombia.Rio
Lopez, Pittier 1088 (holotype, NY).
Figs. 2A, 109, 110.
159
Paraleucobryum densifolium Th6riot, Rev. Bryol.
Lich6nol. 11: 64. 1939. Bizotia densifolia(Th6riot)
Pierrotin Bizot, Pierrot& Pocs, Rev. Bryol. Lich6nol. 40: 26. 1974, comb. inval. Type. Colombia,
Paramo El Boquer6nnear Bogota, Troll 2144 (holotype, PC).
CampylopusrenneriHerzog,Biblioth.Bot. 88:5. 1920.
CampylopuslatinervisHerzog,Biblioth.Bot. 87: 19.
1916. nom. illeg.Type. Bolivia. MoranentalTorreni,
Herzog 3738 (holotype,JE; isotype, B).
Campylopusrennerivar. latelimbataTh6r.,Rev. Bryol.
Lichenol.Ind. 1-22: 18, 1954, nom. inval. Material.
Bolivia. Herzog 4318b, 4813a (JE).
Plants to 8 cm high, in whitish-green, rarely
blackish loose to dense tufts, variable in size and
appearance. Stems erect, sometimes branched,
densely foliate with imbricate leaves ("leucobryoid"). Leaves 6-8 mm long, from oblong base
narrowed to an acumen of variable length, entire
at margins, or with a few teeth at the extreme
apex. Costa very broad, filling %of the leaf width,
excurrent, narrowed at leaf base, in transverse
section with lax ventral hyalocysts, a median band
of chlorocysts and a band of chlorocysts alternating with leucocysts at the dorsal side, ridged
at back in the upper part of the leaf. Alar cells
lacking. Basal laminal cells hyaline, rectangular,
16-32 x 38-96 #sm, at margins in 5-15 rows
narrower, elongate, forming a distinct border.
Upper laminal cells oval, 5-10 x 10-19 ,tm incrassate. Lamina vanishing below midleaf.
Seta 6-8(-15) mm long, brownish, sinuose.
Capsule 1.5 mm long, erect, symmetric, ovoid,
brownish, furrowed when empty. Operculum
long-rostrate, 1 mm long. Calyptra ciliate at base.
Distribution (Fig. 110). On soil in wet paramos
in elevations between 3400 and 4300 m, found
once in southern Mexico, more widespread from
Costa Rica to Venezuela, Colombia, Ecuador and
northern Peru.
- ...PredbyHndr
. R
k
..
FIG. 110. Distributionof Campylopuspittieri.
Cleef2410, 2548 (U). CAUCA:Volcan Purace, Laguna

de San Rafael, Cleef& Fernandez 579A (U). CUNDINAMARCA:
Paramo de Sumapaz, Cleef1616, 1638, 8315,
10395 etc. (U); Represe del Neusa, Cleef& Jaramillo
4152, Cleef 4170, 4147, 4141 etc. (U); Paramo entre
Cogua y San Cayetano, Cleef 6170, 6118, 6472, 6383
etc. (U); Carretera Paramo de Palacio-Rio Chuza, Cleef
5392 (U); Paramo de Cruz Verde, Cleef3300a, 3228,
3254, 3266 etc. (U); Cabeceras de la Quebrada Chuza,
Cleef 5331 (U); Cordillera de La Leonora 60 km NNE
of Bogota, v.d. Hammen & Jaramillo 3801 (U); Piramo
de Palacio, Lagunas de Buitrago, Cleef 136, 5233, 5245,
4102, 9598 etc. (U); Paramo de Guasca, Cleef 3392
(U); Subachoque, Cuchilla El Tablazo, Linares & Bulla
711 (COLO). META:Paramo de Sumapaz, Cleef 1448,
1464b, 91, 1395, 808, 1257 etc. (U); Cerro Nevado del
Sumapaz, Cleef 7895, 8099a, 8187 (U). SANTANDER:
Near Vetas, Killip & Smith 17531 (NY). RISARALDA:
NevaVolcan Sta. Rosa, van Reenen 709 (U). TOImMA:
do de Tolima, Cleef 3221 (U).
VENEZUELA. MERIDA:Laguna Negra, Paramo de
Summit Mucuchies, Chardon 39 (NY); Parque Nacional Sierra
Specimensexamined.MEXICO. CHIAPAS:
of Volcan Tacana, Mun. de Union Juarez,Breedlove Nevada, between Teleferico stations La Aguada and
La Montana, Fransen 1459 (GB); Griffin et al. 1610
24340 (MO).
COLOMBIA. ARAUCA:Sierra Nevada del Cocuy,
(FLAS); environs de Aguada, Onraedt 5706, 5776 (hb.
Cabecerasde la QuebradaEl Play6n,Cleef8838, 8854,
Frahm); Sierra de Santo Domingo, Paramo de MuSierra Ne8908b, 8909, 8963, 9022 etc. (U); BOYACA:
cubaji, Griffin et al. 858, 1365 (FLAS); Gonzalez-Pevada del Cocuy,Cleef8351, 5691, 8802, 5567 etc. (U); reira 20 (FLAS); Paramo de Piedras Blancas, Griffin et
Boquer6nde Cusiri, Cleef 8806 (U); Paramosal NW al. 1436 (FLAS); Sierra de la Culata, Paramo de los
de Belen, Cleef 1983, 2021, 1983, 1906; Vado Hondo, Conejos, Ruiz-T. 8008 (FLAS); Paramo de San Jose,
Siberia,Cleef9350;Paramode la sarnaentreSogamosa Ruiz-T. et al. 8410 (FLAS); Paramo de Quirora, Mun.
y Vado Hondo, Cleef 9210a, 9205 (U); Paramo de de Estanques, Ruiz-T. et al. 8416 (FLAS); Sierra NePisva, Cleef 4590, 4665, 4727 etc. (U); Paramode la vada de Merida, Pico El Toro, Ruiz-T. 6892 (FLAS).
Rusia, Cleef 7143, 7017, 7461, 6765 (U); Carretera TACHIRA: Paramo El Batall6n above La Grita, Griffin
Vado Hondo-LabranzaGrande,Cleef 9290 (U); Pena et al. 480, 600 (FLAS); Paramo El Rosal between La
Grita and San Jose, Griffin et al. 671 (FLAS).
de Amical, Cleef 9460 (U). CALDAS:Nevado del Ruiz,
160
Flora Neotropica
ECUADOR.AZUAY:
ParamoEl CajasW of Sayausi,
Balslev 1484 (NY). CARCHI:
E side of pass La Piedra,
Steere 8122a (NY); base of volcan Chiles, km 34-36
on road Tulcan-Maldonado,77?57'W,0?47'N,HolmNielsen et al. 5862 (AAU, GB). NAPO:Road QuitoBaeza,Paramode Guamani,01gaard &Balslev10104
(NY); Steere E 232 (NY); S side of Cerra Sumaco,
77?39'W, 0?35'S, Lejtnant & Molau 12976, 12983
(AAU, GB); N side of LagunaVerdecocha,78?21'W,
0?47'S,Lojtnant& Molau 11706 (AAU, GB);Paramo
de Quilindana,Kieftet al. 202, 203 (U); LagunaYuragcocha, 78?21'W, 0?47'S, LGjtnant& Molau 11589
(AAU, GB); Laguna Parcacocha, 78?09'W, 0016'S,
Lojtnant& Molau 11087, 11096 (AAU, GB);San Miguel-PuertoNuevo road, 78?18'W,0?59'S,Lojtnant&
Molau 13786 (AAU, GB). PICHINCHA:
Road QuitoPapallacta, Paramo de Guamani, 78?12'W, 0?17'S,
Holm-Nielsenet al. 6736 (AAU, GB).
PERU.ANCASH:
LagunaLlanganuco,Hegewald7594
(hb. Frahm);LagunaPar6n,Lopezet al. 8372a (HUT).
CAJAMARCA:
Las Lagunas,Hegewald6200 (hb. Hegewald). SAN MARTIN: Pataz, 30 km E of Parcoy, Ham-
ilton & Holligan 49 (MICH).
This species was described as a species ofParaleucobryum by Theriot. Indeed, the structure of
the leaf with the broad costa and the small lamina
and especially the transverse section of the costa
much resemble this genus. However, since the
description of sterile material by Theriot this taxon has been found several times with sporophytes, which clearly indicate the relationship to
Campylopus. It is a matter of speculation whether Campylopus pittieri can be regarded as a
transitional species between Campylopus and
Paraleucobryum indicating a phylogenetic relationship, or whether the anatomical structure of
the leaves have evolved independently as an adaptation to a higher water storage in the costa.
In this regard it is interesting that Campylopus
pittieri (and also C. albidovirens) as well as the
species of Paraleucobryum do not contain flavonoids and that thus are not only structural
homologies but also a conformity in chemical
contents.
Campylopus pittieri is most closely related to
C. albidovirens, which differs mainly by the presence of alar cells and a not so much leucobryoid
appearance. In contrast to C. pittieri, which is
confined to alpine regions, C. albidovirens occurs
mainly in upper montane and subalpine forest
and may be interpreted as an ecological vicariant
species.
Pierrot (in Bizot, Pierrot & P6cs, 1974) based
the creation of a separate genus, Bizotia, on the
presence of pores on the ventral hyalocysts. Such
pores could, however, not be observed by the

author,even after stainingwith methylene-blue.
Placement in the genus Campylopusis, moreover, meanwhile confirmedby the existence of
sporophytes.
Theriot, when describing Paraleucobryum
densifolium, differentiated two varieties, var.
congestumand var. latilimbatum,both collected
at the same locality-the var. congestumat 3400
m elevation, the var. latilimbatumat 2500 m, a
distinctly lower altitude-which were newly
combinedwith C.pittieriby Florschitz-deWaard
and Florschiitz (1979). The var. congestumis,
accordingto the description,probablythe typical
variety, whereas var. latilimbatumis a smaller
modification.There can indeed be two extreme
forms distinguished,one of very robust plants
and swollenfoliation,much resemblinga species
of Leucobryum,and another of slender plants,
resembling(moreor less) specimensof C. nivalis.
The studyof herbariummaterialshowsthat both
extreme modifications can often be distinguished,but arelinkedby intergradingformsand
thus will probablyhave no taxonomic value.
Accordingto Robinson (1967) and Florschiitz
and Florschiitz-deWaard (1974), Campylopus
pittierishould be identicalwith Campylopusargyrocaulon(C. Miller) Brotherus.However, as
shown by the Paraleucobryum-liketransverse
section of the costa, C. pittieri is quite distinct
and differentfrom that species. This transverse
section resembles, in the basal part of the leaf,
exactlythat of Paraleucobryumenerveand in the
upperpartthatofP. longifolium.In Campylopus,
a transverse section of the costa consists of 5
layers:a ventral (stereids,substereidsor hyalocysts),a median(chlorocysts),and dorsallya band
of stereids, substereidsor leucocysts and an epidermallayerof chlorocystsalternatingwith leucocysts. In C. pittierithereare only 3 or 4 layers:
ventraland dorsalhyalocystswith median chlorocystsin the basalpartof the leaf, or alternating
dorsal chloro- and leucocysts in the upper part
of the leaf. Contraryto these discrepanciesin the
gametophyte, the sporophyte is typical for the
genus Campylopus,resemblingmost species of
sect. Homalocarpus. It has sinuose setae (not
straight as in Paraleucobryum),ovoid to short
cylindric capsules (not elongate-cylindricas in
Paraleucobryum),the capsuleshave no stomata
as in Paraleucobryumand the spore size is also
typical for species of Campylopus.Although a
161
SystematicTreatment
paratype of "Paraleucobryum densifolium"
mentioned by Theriot has sporophytes,Theriot
did not mention it in the type description.
Althoughthis species had been recordedonly
a few times until 1975, it is one of the most
common and typical paramo-mosses.It varies
considerablyin the length of the leaf tip, which
can be shortor elongate,smooth or dentate,also
at the back as a rat's tail file, the height of the
stems, varyingfrom 1 to 8 cm, the color, which
is usually pale or whitish green or rarely even
blackish, and the foliation, which may be appressedor even swollen.
There was some confusion with the types of
Campylopusrenneriand C. latinervis.The types
of both species were collected at the same locality, but publishedin differentyears.The syntype
of C. latinervisconsistsof C. argyrocaulon.Other
material in the herbariumof Herzog named as
C. renneri consists of C. nivalis. Furthermore,
Theriotdescribed(not validly)a varietylatelimbata of C. renneri in the same publication as
Paraleucobryumdensifolium, which are both
synonymous.
Seta unusually long, to 15, rarely to 25 mm

long, straightbut cygneouswhen wet. Capsule2
mm long, pale to darkbrown in age, operculum
cylindric,symmetric,long-rostrate.
Distribution(Fig. 112).On moist, shadedcliffs
and treetrunks,on rottenwood in montanerainforests(in Mexico also in humid pine-forestsbetween 1200 and 3000 m), in Mexico, CostaRica,
Venezuela,Colombia,Ecuador,Peru,Boliviaand
Brazil.
PuerSpecimensexamined.MEXICO.MICHOACAN:
to Garnica,Delgadillo572 (NY). OAXACA:
Km 103.3
N on Hwy. 175 near Pedro Ylox, 17?36'N,96?24'W,
Vitt 17707 (MICH);SierraJuarez,gap on Hwy. 175,
McFarland&Sharp8615 (TENN);Llanode las Flores
N of Oaxaca,Iwatsuki&Sharp5461 (TENN);between
Ixtlan de Juarez and Tuxtepec, Sharp et al. 2383c
(TENN); Cima del Cerro Corralde Piedra, 17?10'N,
96039'W,Cardenas4291 (MEXU).
COSTA RICA. ALAUELA:Vic. volcan Poas, 10?11'N,
84?14'W,Crosby6278 (MO). CARTAGO:

Refugio Silvestre Tanpanti,Arrocha963 (hb. Frahm).SANJOSE:
17 km SE of El Empalme,Crosby10885 (MO).
COLOMBIA. CUNDINAMARCA: Environs d'Alban,
Onraedt8581 (hb. Frahm);Subachoque,Paramo El

Tablazo, Frahm 885247 (COL). MAGDALENA:
Parque
Nacional de la SierraNevada de SantaMarta,Rangel

Camino Real entre
49. Campylopusreflexisetus(C. Miiller)Broth- et al. 844, 854 (FLAS). RISARALDA:
erusin Engler& Prantl,Nat. Pflanzenfam.1(3): Hda. La Sierray Termalesde Sta. Rosa, van Reenen
860 (U).
332. 1901. DicranumreflexisetumC. Miller,
VENEZUELA.MERIDA:
Linnaea38: 586. 1874. Type. Colombia. An- Paramode Mucubaji,Griffinet al. 1350 (FLAS);Pratioquia: Paramo de Sonson, Wallis s.n. (ho- mo La Negra above Bailadores, Griffinet al. 2179
Paramode Don Pedro entre El Morroy Arlotype, destroyed at B; lectotypus nov., (FLAS);Ruiz-T.
& Lopez-F. 9547 (FLAS).TRUJILLO:
icagua,
112.
H-BR).
Figs. 111,
Along the old Bocon6-Trujillohighway,Griffinet al.
benedictiiHerzog,Beih.Bot. Centralbl.
Campylopus
Abra
26(2):50. 1910.Type.Bolivia.Cochabamba:
de SanBenito,Herzogs.n.(holotype,
JE;isotype,S).
Campylopus
ptychotheca
Herzog,Beih.Bot.Centralbl.
Inca26(2):49. 1910.Type.Bolivia.Cochabamba:
corral,Herzog284 (holotype,JE;isotype,B).
Slenderplants in loose, greenishtufts to 6 cm
high, slightly tomentose below, equally foliate.
Leaves erect, spreading,7-8 mm long, concave,
narrowlylanceolate,endingin a verylong,smooth
subula(twice as long as the lamina).Costa filling
'/2 of the leaf base, smooth at back, excurrent,in
transversesection with large ventral hyalocysts
(more than half the height of the section), with
dorsal stereidslackingtowardthe margins.Alar
cells differentiated,hyaline or red-brown.Inner
basal laminal cells rectangular,38-60 x 15-18
Jim,those at marginslong, narrowand hyaline
in 10-15 rows.Upperlaminalcells elongate-oval,
20-25 x 6-8 um
1339 (FLAS).
ECUADOR. IMBABURA:
Lago San Marcos, Cazalet
N slope of Volcan
& Penninston47 (NY). PICHINCHA:
Pichincha,Balslev1748 (NY). ZAMORA:
Loja-Zamora
highway20 km E ofLoja, Steere&Balslev25795 (NY).
PERU. ANCASH:
CordilleraBlanca, LagunaLlaca,
Frahm,Camp.Peruv.Exsicc. 19(ALTA, B, BM, BUF,
DUKE, EGR,F, FLAS,GRO,GZU, HBG,KR, NAM,
NICH, NY, MEXU, NFLD, PC, POZ, RNG, S, U).
BOLIVIA.COCHABAMBA:
Rio Khuri N of Corani,
17?12'S,65?35'W,Lewis 79-2373 (F); E face of Cerro
Chua Laguna, 17?13'S,65?53'W,Lewis 79-2230 (F).
LA PAZ:30 km NE of La Paz, 16?19'S, 67?50'W, Crosby
13569 (MO). TARIJA:

5 mi NE of Iscayachi,21?27'S,
64?54'W,Lewis 79-690 (F).

Parque Nacional Itatiaia,
Agulhas Negras, Frahm 1625, 1626 (hb. Frahm).
This species resembles C. densicoma in the

long leaf subula, which is, however, smooth and
not toothed as in C. densicoma.
Var. circinatus Theriot, Rev. Bryol. Lichenol.
11: 42. 1939 described from Bolivia (Herzog
Flora Neotropica
162
14.5mm
:"?
FIG. 111. Campylopusreflexisetus(Herzog284, B). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 /m.
3938, JE)differsby hamateleaves. It is probably CampylopusatratusBrotherus,Trans.Linn. Soc. Bot.

ser. 2, 6: 89. 1900. Pilopogon atratus (Brotherus)
a parallelform to C. densicomavar. yungarum.
Brotherusin Engler& Prantl,Nat. Pflanzenfam.1(3):
As in other hamate foliate forms in this genus,
336. 1901. Type. Venezuela. Mt. Roraima, Mcit is not known whetherthese are geneticallydisConnell&Quelch527 (holotype,H-BR;isotypes,K,
tinct and requireinfraspecificdifferentiation.
NY, S).
50. Campylopusrichardii Bridel, Mant. Musc.
73. 1819. Type. Guadeloupe,Parkers.n. (holotype, B).
Figs. 4D, 113, 114.
Campylopuscaracasanus(C. Miller) Paris,Ind. bryol.

242. 1894. ThysanomitrioncaracasanumC. Muller,
Linnaea42: 471. 1879. Pilopogoncaracasanum(C.
Mller) Brotherusin Engler & Prantl, Nat. Pflanzenfam. 1(3): 336. 1901. Type. Venezuela.Fendler
163
---- 4--3'
-eHed
Pr-oses
G'l5:>::^
A^
2?0_
~ ~
of00 Ca000-u
Distribution0
.
Peo-pareby HendhkR. Rypkem
^:
FG
FIG.
112
112.
Distribution
reglexisetus.*7".. -
of
Campylopus
..
refiexisetus.
36 (holotype, destroyedat B; lectotypusnov., NY; Campylopusrigidus (Hornschuch)Jaeger, Ber. Thatigk.St. GallischenNaturwiss.Ges. 1870-1871: 443.
isotype, S).
1872. Thysanomitrion
rigidumHornschuch,Fl. bras.
Campylopuslaevigatus(C. Miiller) Schimper in Be1(2): 15. 1840. Dicranumrigidum(Hornschuch)C.
scherelle, Mem. Soc. Sc. Nat. Cherbourg16: 167.
Muller,Syn. musc. frond.I: 409. 1848. Type. Brazil.
1872, nom, illeg., non Bridel 1819. Dicranum laeOuro Preto, Martiuss.n. (n.v.).
vigatumC. Muller,Syn. musc. frond.II: 601. 1851.
Type. Mexico. Liebmann s.n. (holotype, destroyed CampylopussteyermarkiiBartram,Bot. Soc. Venez.
Cienc.Nat. 106:35. 1963. Type. Venezuela.Cabeceat B; lectotypusnov., H).
rasde Rio Chicanan,6?5'N,62?W,Steyermark89600
Campylopusmuelleri(Hampe)Jaeger,Ber.Thatigk.St.
Gallischen Naturwiss.Ges. 1870-1871: 442. 1872.
(holotype,FH; isotype, VEN).
muelleriHampe,Ann. Sci. Nat. Bot. Campylopusyunqueanus(C. Miller) Paris,Ind. bryol.
Thysanomitrion
ser. 5,3:363.1865. Type.Colombia. Triana&PlanSuppl. 99. 1900. ThysanomitrionyunqueanumC.
chon s.n. (n.v.).
Muller,Hedwigia37: 225. 1898. Type. PuertoRico.
SierraLuquillo,SinteniusF 29 (holotype,destroyed
Campylopusnigerrimus(C. Miiller)Paris, Ind. bryol.
at B; lectotypusnov., H-BR).
Suppl. 94. 1900. ThysanomitrionnigerrimumC.
Miller, Bull. Herb. Boissier6: 39. 1898. Type. Bra- ThysanomitriummiserumBrotherus,Denkschr.Akad.
Wiss. Wien math.-nat.KI.83: 264. 1926. Type. Brazil. Rio de Janeiro:Tijuca, Ule 1081 (syntype, dezil. Serrado Itatiaia,Schiffner754 (holotype,H-BR).
stroyedat B; lectotypusnov., H-BR).
Campylopuspuiggarii(Geheeb & Hampe) Paris, Ind.
bryol. 259. 1894. ThysanomitrionpuiggariiGeheeb
Plants conspicuously blackish, only lighter at
& Hampe, Flora64: 346. 1881. Pilopogonpuiggarii
to 10 cm high. Stems equally foliate, sterile
tips,
(Geheeb & Hampe) Brotherusin Engler& Prantl,
Nat. Pflanzenfam.1(3): 336. 1901. Type. Brazil.Sao plants with leaves erect spreading when wet or
appressed when dry, fertile plants with comose
Paulo, Puiggaris.n. (isotype, JE).
164
Flora Neotropica
^~F
5.4cm
Ai
14mm
(?A
.U
......a:
FIG. 113. Campylopusrichardii(CampylopodesExsiccatae75). A. Plant.B. Leaf.C. Leaf-tip.D. Transverse

50 Mm.
perichaetiaon appressed foliate stalks, tomentose below. Leaveslanceolate,ca. 7-8 mm long,

ending in a hyaline denticulatehairpoint.Costa
filling /3-1/2 of the leaf base, excurrent, in trans-
verse section with a ventral and dorsal stereid

band, ridged at back in the upper part, smooth
below. Alarcells differentiated,inflated,reddishbrown. Basal laminal cells very incrassate and
strongly pitted, rectangular,3-6 x 10-35 gm,
narrowerat margins,often 1-2 rows of hyaline
cells at the outermost margin. Upper laminal

cells elongateoval, 3-4 x 10-35 um, incrassate,
slightly pitted.
Setae aggregated,sinuose or curved, 7-8 mm
long, dark brown. Capsule 1.5 mm long, symmetric, smooth or furrowedwhen empty, scabrous at base. Operculumshortly rostrate.Peristome teeth very narrow,hyaline, filiform, split
nearly to the base. Calyptraciliate at base.
Distribution(Fig. 114). On seasonalwet rocks
165
100
90
.......~
...
.
.....
2?0
?0
1 ;O 20'0
6?0
80
60
70
-----. - . -- .
------.
50
40
-----2
8?01000k l
3~O4~ 5;0'00,,,.I.
Prepared by Hendrik R.Rypkema5
FIG.114 Ditribtio
|Preparedby HendrikR Rypkema
ofCamylops
rchadii
----
FIG. 114. Distributionof Campylopusrichardii.
and cliffs and wet gravelly soil, from 900 to 3700

m through the Neotropics in Mexico, Guatemala, Costa Rica, Panama, Lesser Antilles, Dominican Republic, Puerto Rico, Jamaica, Guyana, Venezuela, Colombia, Ecuador, Peru,
Bolivia and Brazil. The records from Chile, Juan
Fernandez Islands must be referred to C. clavatus.
Specimens examined. MEXICO. Without locality,
Karstens.n. as C. karstenii nom. dub. (H-BR); Liebmann s.n. (H-BR). Huatusco,Mohr s.n. (FH).
COSTARICA.CARTAGO:
RefugiosilvestreTapanti,
Arrocha1012 (hb. Frahm);CerroAscensi6n,McDaniel
6793 (NY); nearsummit of InteramericanHighwayat

La Ascensi6n, 9?36'N, 83?46'W,Crosby6134 (MO).
SAN JOSE: Paramo Buena Vista 90 km S of Cartago,
Griffin&Eakin 580 (FLAS);CerroDaserarea,9050'N,
84?07'W,Crosby9828 (MO);Cordillerade Talamanca,
Chirip6-massif,Kuhbier612 (BREM);San Pablo de
Dota, Pittier 9913 (H-BR, PC). HEREDIA: SW slopes
of Volcan Barba,Crosby3700 (MO);Las Vueltasarea

12 km fromSanRafael,10?6'N,84?3'W,Crosby10933b
(MO).
JAMAICA. Blue MountainPeak 18?3'N,76?35'W,
Crosby3383 (MO).
LESSER ANTILLES. Without locality, Funck &
Schlim 229 (KIEL). MARTINIQUE:

Mt. Pele, Duss 298
(H-BR); Crosby 4698 (MO). GUADELOUPE: Volcan de
la Soufriere,Funck& Schlim 229 (H-BR);Husnot 132
166
Flora Neotropica
(PC);De Sloover 33827 (NAM); Crosby4858 (MO).

MountScenery,Wiersma& vanSlageren227BM
SABA:
TroisPitons, Wisemans.n. (MO).
(NY, U). DOMINICA:
DOMINICAN REPUBLIC.SANJUAN:Pico Duarte, Buck 8462 (NY).
PUERTO RICO. Serra de Luquillo, Hioram 410
(NY). El Yunque RecreationArea, Buck 4079 (NY).
32.5 km NE de Sons6n
COLOMBIA.ANTIOQUIA:
hacia La Union, 5?50'N,75?15'W,Churchill& SastreDe Jesus 13089 (NY); Paramode Frontino, Mun. de
Urrao, 6?25'N, 76?5'W,Churchillet al. 13344 (NY);
Mun. de Guatape,6?18'N,75?9'W,Sastre-DeJesus et
al. 1338 (NY); 20 km de Medellin hacia Rionegrom,
6?10'N,75?25'W,Sastre-DeJesis&Churchill757 (NY);
Medellin, Mayor s.n. (PC);La Piedra del Penol, Nee
& Mori 4318 (MO). BOYACA:Vado Hondo, Siberia,
Cleef9256 (U); Pena de Arnical,Cleef9448 (U); Paramos al NW de Belen,Cleef9724 (U); Paramode Pisva,
Cleef 4694 (U); Paramode la Rusia, Cleef 7218 (U);
SierraNevada del Cocuy, Cleef8571 (U). CAQUETA:
30 km S of Guadelupe, Gentry et al. 9047 (MO).
CUNDINAMARCA:Near Zipaquira, Schultes&Bell 11463
(NY); El Boquer6nnearBogota, Troll2026 (JE);Paramo entreCoguay San Cayetano,Cleef6470 (U); Paramo de Palacio, Cleef3826a (U); Paramode CruzVerde, Cleef3029 (U); Paramode Guasca,Schultes239b
(FLAS);Sabana de Bogota, Schultes 12193 (FLAS);
Subachoque,CuchillaEl Tablazo,Linares&Bulla 788
(NY); between Bogota and Carchi, Weir176 (H-BR);
ParamoEl Tablazo, Wood4 (hb. Frahm);Bogota,Weir
s.n. (H-BR). META:Paramode Sumapaz,Cleef 7752
(U); CerroNevado del Sumapaz,Cleefl454 (U); Hoya
El Nevado, Cleef 1492 (U).
VENEZUELA.MERIDA:
Lagunade los Anteojosbelow Loma Redonda,Mdgdefrau1609 (hb. Frahm);Sierrade Santo Domingo, Paramode Mucubaji,Griffin
et al. 1035 (FLAS);environsde Iguada,Onraedt5787
Paramode Tama above Paez,
(hb. Frahm).TACHIRA:
Griffinet al. 888 (FLAS);ParamoEl Rosal betweenLa
Grita and San Jose, Griffinet al. 758 (FLAS).
GUYANA. Upper MazaruniDistr., N slope of Mt.
Roraima, 5?16'N,60?43'W,Gradstein5358 (U).
base
ECUADOR.CARCHI:
RoadTulcan-Maldonado,
of volcan Chiles, 0llgaard & Balslev8356 (NY); road
fromPlay6nde SanFranciscoto El Carmelo,77?38'W,
0?38'N, Lijtnant et al. 12290 (AAU, GB); 3 km E of
Frailej6n,77?40'W,0?42'N,Lejtnantetal. 12161(AAU,
8895 (hb.Frahm).GUANABARA:
Verwimp
Brejoda Lapa,
Lowy 1330 BR (LAF).MINASGERAS:Mun. de Ouro
Preto, km 392 along road BR 135, Frahm 1439 (hb.
Frahm);Serrade Cip6,estradaLagoaSanta-Conceiaio
de Mato Dentro, Yano510 (SP);Caraca,Wainio1885,
Ule 1369 (H-BR);Serrado OuroPreto,Damazio 2129
Sio Jose dos Pinhais,25?40'S,49?W,
(H-BR).PARANA:
Landrum2426 (NY); 10 km E of Curitiba,25025'S,
Ti49?10'W,Landrum2323 (NY). Rio DEJANEIRO:
juca, Ule 152 (H-BR, PC); Serra do Itatiaia, Maua,
Bandeira436 (H-BR);ParqueNacionalItatiaia,Agulhas Negras,Frahm,Camp.Bras.Exsicc. 14, 15 (ALTA,
KR, M, MO, NFLD, NICH, PC, S, SP, TENN, U);
17.5 km NE of Uniao, 22023'S,44?41'W, Vitt21470
Piloes, Palhoca, Reitz &
(ALTA). SANTACATARINA:
Klein2514 (FH);SerraGeral,Brixen s.n. (HBG). SAO

Mun. de Paraibuna,Vital8758 (SP);Apiahy,
PAULO:
Puiggari414 (H-BR);Serrado Mantiqueira,Campos
do Jordao, Schdfer-Verwimp6898; Serrado Bocaina
betweenCamposda Cunhaand Sao Jose des Bareiros,
Frahm, Camp.Bras. Exsicc. 13 (ALTA, B, C, DUIS,
FLAS,GOET,GZU, H, HBG, INPA, JE,KR, M, MO,
NFLD, NICH, PC, S, SP, TENN, U).
BOLIVIA.LAPAZ:2 mi from Soratadown to Rio
Consata,15?41'S,68?43'W,Lewis79-1240 (F);Laguna
Verde near Comarapa,Herzog 4208 (JE);Apolo region,Cargadira,Williams1760(H-BR);SanJos6-Apolo
Casteon,
trail, Williams 1763 (H-BR, NY). TARIJA:
along road to Tarija,21?28'S,64?12'W,Lewis 79-539
(F).
Campylopus nigerrimus was described from

two syntypes, the original material of which has
been destroyed at B. Therefore a lectotypification
is proposed here, choosing one of the syntypes
and, additionally, H-BR as new location of the
type material.
Specimens from Mt. Roraima described as C.
atratus and additional collections matching the
type of this species differ by extremely large plants
with longer leaves and more strongly elongate
upper laminal cells; they are interpreted as "giant" forms of C. richardii.
The distribution on the Caribbean islands is
GB). LOJA:Road Loja-Zamora km 12, 79?9'W, 4?S,
Holm-Nielsen et al. 3677 (AAU, GB). NAPO:Cerro
relatively scattered and, surprisingly, C. richardii
Sumaco, 77?39'W,0?35'S,Lojtnant & Molau 12705 has not
yet been found in Cuba or Haiti. Fur(AAU, GB). Gualaquiza,Allioni 8281 (H-BR);Bomall collections from these regions are
thermore,
"Andes
Allioni
8179
boiza,
(H-BR).
Quitenses,"Spruce
53 (NY, PC).
sterile and thus it may be supposed that C. richKm 413 along road from Cha- ardii has reached these islands by occasional long
PERU. AMAZONAS:
chapoyas to Cajamarca,Frahm, Camp. Exsicc. 75 distance dispersal of unisexual spores, which did
(ALTA, B, BM, F, NY, S, U). JUNIN:Chilifrutanear
Huancayo, Hegewald 9208 (hb. Hegewald). SAN not allow for sexual reproduction due to the lack
MARTIN:Road Moyobamba-Chachapoyaskm 387, of the other sex.
Frahm, Camp. Peruv.Exsicc. 20 (ALTA, B, BM, C,
F, NY, S, U).
BRAZIL. BAHIA:Serra do Tombador 18 km E of 51. Campylopus savannarum (C. Miiller) Mitten, J. Linn. Soc. Bot. 12: 85. 1869. Dicranum
Morrodo Chapeu,Irwin et al. 30748 (NY). ESPIRITO
SANTO:Along BR 262 W of Venda Nova, Schafersavannarum C. Miiller, Syn. musc. frond. 2:
167
596. 1851. Type. Surinam. Kegel s.n. (holotype, destroyed at B; lectotype, BM; isolectotypes, H-BR, JE, L).
Figs. 115, 116, 117.
Campylopusarenaceum (Brotherus)J.-P. Frahm, J.
Bryol. 8(2): 258. 1974. Thysanomitrionarenaceum
Brotherus,Denkschr.Akad. Wiss. Wien math.-nat.
KI. 83: 264. 1926. Type. Brazil.Sio Paulo:Faxina,
Schiffner1100 (lectotype,H-BR).
CampylopusbartlettiiBartram,J. Wash.Acad. Sci. 22:
477. 1932. Type. Honduras.Bartlett 12973 (holotype, FH; isotypes, DUKE, F, MICH, NY, S).
Campylopusschiffneri(Brotherus)J.-P.Frahm,J. Bryol.
8(2): 260. 1974. ThysanomitrionschiffneriBrotherus,Denkschr.Akad.Wiss. Wienmath.-nat.KI.83:
265. 1926. Type. Brazil.Sao Paulo:CampoGrande,
Schiffner560 (holotype,H-BR).
CampylopusspruceiMitten, J. Linn. Soc. Bot. 12: 81.
1869. Type. Brazil.Rio Negro,Spruce59 (holotype,
NY; isotypes, F, FH, H-BR, H-SOL).
?CampylopustortilipilusJ.-P. Frahm, Cryptogamie
Bryol.Lichenol.2(4):446. 1981. Type. Brazil.Piaui,
Mun. de Corrente,Vital8232 (holotype,SP).
Additionally,23 synonymsfromAfrica(Frahm,1985a)
and 4 from SE Asia.
Plants robust, 2-5 cm high, in loose, yellowish
green to dark green tufts, reddish tomentose below. Leaves 5-6 mm long, lanceolate, gradually
acuminate, serrate above, either flexuose spreading when dry with a subhyaline hairpoint (mod.
savannarum) or appressed foliate when dry with
a hyaline hairpoint (mod. bartlettii). Costa filling
about 1/2of the leaf width, ridged at back, excurrent into a roughly serrate awn, in transverse
section with ventral and dorsal stereids, the ventral stereids becoming substereidal towards the
leaf base. Alar cells reddish or hyaline, inflated,
rarely inconspicuous. Basal laminal cells 13-16
x 15-45 ,um, short rectangular, 1.5-3:1, thickwalled, those at margins smaller and quadrate in
1-3 rows. Upper laminal cells rhombic or oval,
18-26 x 7-11 zm.
Distribution (Fig. 117). A pantropical species
occurring on sandy and gravelly soil, on rocks or
soil-covered outcrops, at bases of trees in dry
forests, in the neotropics from SE Brazil to the
Guianas and also in Central America in Panama,
Costa Rica, Nicaragua, Guatemala, Honduras
and Mexico in low elevations between sea level
and 1500 m.
Specimens examined. MEXICO. JAusco: Near
Puerto los Mazos above Autla.n,Sharp et al. 3186
2 miles W of Compostela,Norris
(TENN). NAYARIT:
& Taranto15174 (TENN). PUEBLA:
Xicotepecde Juarez, Delgadillo 1255 (MEXU).
FIG. 115. Campylopussavannarum(holotype of

C. sprucei).
GUATEMALA. QUEZALTENANGO:Slopes of volcan
Santa Marta,Steyermark33672 (NY).
NICARAGUA. JINOTEGA: Region of La Montanita,
Standley10353 (F);SierraW of Jinotegaalong trail to

Cerrode la Cruz,Standley 10219 (F).
COSTA RICA. PUNTARENAS:
Along road E of S.
Vito towardSabalito, Crosby2620 (MO).

PANAMA. Cerro Campana below Su-Lin Motel,
Crosby4454 (MO).
ANTILLES. ST. EUSTATIUS & SABA: Wiersma & v.
Slageren272M (U); Stoffers3463A (U).

COLOMBIA. AMAZONAS:
Rio Apaporis, Cachivera
de Jirijirimo,Schultes& Cabrera1198A(NY);SanJose
de Guaviare, Schultes 11126 (FLAS). VAUPES:Rio
Guainia,PuertoColombia,2?40'N,67?30'W,Schultes
et al. 18232 (FLAS).
VENEZUELA.
BOLIVAR:El Frijol, Holmquist 39
(hb. Frahm);cumbrede CerroGuaiquinima,5?44'N,

63?41'W, Steyermark & Dunsterville 113440 (NY).
MERIDA:BetweenMeridaand La Azulita, Griffnet al.
1532 (FLAS).
GUYANA.Upper MazaruniDistr.,Jawalla,5?40'N,
60?29'W, Gradstein4954 (U); trail from Kamarang
riverto Pwipwimountain,5?54'N,60?46'W,Gradstein
5676 (U); E bank of Warumariver, 5?28'N,60?47'W,
Gradstein 4986 (U); Mt. Latipu, 5?57'N, 60?38'W,
Gradstein5544 (U).
Kabalebo area, FlorschiitzSURINAM. NICKERIE:
de- Waard& Zielman 5483 (U); top of Voltzberg,Florschfitz4572 (U).

FRENCH GUIANA. Piton Rocheux Remarquable,
Flora Neotropica
168
FIG. 116. Campylopussavannarum(Vital 2750, SP). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 um.
Cremers7118 (hb. Frahm);massifdes EmerillonsCentre, Cremers6862 (hb. Frahm).
BRAZIL. AMAZONAS:
Estrada Manaus-Caracarai km
to Mato Grosso, Schifer- Verwimp8548 (hb. Frahm).

Mun. de Chapada dos Guimaraes,
MATOGROSSO:
15?25'S, 55?49'W, Vital 9973 (SP); Serra Ricardo Fran-
130, Griffn et al. 51420 (INPA); Igarapenear Icana, co, 15?S,60?W,Windisch1753 (SP);vic. Buriti,Prance
Serrado Moeda,
Sioli 17 (JE);EstradaManaus-BoaVistakm 45, 02?45'S, et al. 19248 (NY). MINASGERAIS:
60?06'W,Frahm, Camp.Exsicc. 27 (B, BM, F, NY, S, Mun. de Itabirito, Vital 5528 (SP); NW Ouro Preto,
U). BAHIA:Serra do Tombador, 5 km S of Morro do
Chapeu,Irwinet al. 32589 (NY); 4-6 km E of Morro FLAS,GOET,GZU, H, HBG, INPA, JE,KR, M, MO,
do Chapeu,Boom & Mori 1280 (NY); Mun. de Mil- NFLD, NICH, PC, S, SP, TENN, U); Mun. de Passos,
agres,along BR 116, Vital 5945 (SP); ca. 50 km NW nearRepresade Furnas,20038'S,46?18'W,Vital7644
de Jequi6, Vital 8734 (SP). CEARA:Morro da Boa Agua,
(SP);ca. 2 km SE of Mendanha,Vital6510 (SP);Mun.
Eugenio 1263 (RB). GoiAs: 25 km N of Alto do Par- de Janauba, 15?46'S, 43?17'W, Vital 7915 (SP). PARA:
aiso, Irwin et al. 32997 (NY); Mineiros at the border Serrado Cachimbo,km 780-820 on Cuiaba-Santarem
169
90
'XC
70
80
60
50
40
6301
,^
'ct'
5-^^^
'
::
^ o
<^^.
A,
r""_-'
.~"~~~"~..-
,- ......
FI.! 7
FIG. 117.
'--------
--'......lI
Disrbtoofapypusanau.
Distribution of Campylopus savannarum.
In accord with the wide range and the broad

highway, 9?22'S, 54?54'W,Reese 16316 (NY); 2 km
W of AMZA camp, Serrados Carajas,6?4'S,50?8'W,
ecological amplitude, Campylopussavannarum
Sperling 5666 (NY); 25-30 km NW of SerraNorte, is variable in
size, habit, presence of a hyaline
Pocinhos,v.
Daly et al. 1750 (NY). PERNAMBUCO:
5100/a(JE);Recife,estradaparaMumbeca, or subhyalinehairpoint,the shape of the upper

Liitzelburg
laminal cells, and color. It can, however, be difseteCidades,Piracuruca,
Vital5411(SP).
ferentiatedby the band of quadratecells along
In Brazil, Campylopus savannarum is frequent the basal leaf marginand the transversesection
in the Cerradoand caatingavegetation.It occurs of the costa with ventralsubstereidsin the lower
also in the Amazon lowland in "campinas"and and sterids in the upper part of the leaf.
Campylopusbartlettiiis an adaptationto drier
may here be interpretedas relict of a wider extension of savannah-likevegetation during the habitats,with appressedleaves and hyaline hairpoints. It has been combined by Florschiitzas a
Pleistocene.
Campylopus savannarum rarely produces spo- subspeciesof C. savannarumand is, indeed, difrophytes.The main distributionin the tropics is ferent in habit and habitat. There are, however,
in the lowlands,wherelong-distancedispersalby all possible intergradationsfound between the
aircurrentsfromone continentto the otherseems type of C. bartlettiiand that of C. savannarum,
to be unlikely.It may be thereforeinterpretedas also in Africa and SE Asia, which makes it iman ancientspecies with a continuousrangein the possible to make any taxonomic differentiation
Mesozoic, which was split by continentaldrift. between the taxa. Transition forms with apAndrade-Lima80-6430 (SP). PIAu: ParqueNacional
170
Flora Neotropica
FIG. 118.
L2aLi
Campylopus sehnemii (Sehnem 2310, FH). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section
of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 ,um.
pressed leaves (but not hyaline tipped leaves),

especially, are found relatively frequently.Valued as modifications,they can, however, characterize differentexpressions of this species on
differenthabitats.Campylopusspruceiis such an
expressionfromthe wettestpartof the ecological
range, occurringin Campina-vegetationin the
Amazon lowland.It cannotbe excludedthatsome
of these expressionsare more than modifications
but are, rather,geneticallydistinct as, e.g., specimens namedC. arenaceumfromSEBrazil.These
plantsarevery homogeneousin appearance,with

narrower,appressedleaves but no distinct hairpoint, and are found only on sandstone.
Campylopustortilipilushas been describedas
a species because of its leaves being suddenly
contractedat apex and a hairpoint,which is up
to /3 as long as the leaf blade and conspicuously
tortuose. The leaves are only up to 3 mm long
and the plants to 1 cm high. However, the anatomical charactersof the leaf are those of C.
savannarumand the small size and long hair-
171
100
__J-^_
0100 200
80
90
300
60
70
50
o0
____\
400 500 100 m --l-
Prepared by Hendrik R.Rypkenao
FIG. 119. Distributionof Campylopussehnemii(@)and C. sharpii(0).
points may be an extreme adaptation to a dry

habitat. It is questionable whether the tortuose
hairpoints are genetically distinct and have a
function for protection against radiation or are
merely the result of a disturbeddevelopment.
52. Campylopus sehnemii Bartram, J. Wash.
Acad. Sci. 42(6): 179. 1952. Type. Brazil.Rio
Grandedo Sul: CampestreMontenegro,Sehnem 2310 (holotype, FH).
Figs. 118, 119.
Plants bright green, 1.5-5 cm high. Stems
equallyand denselyfoliate.Leavesappressedand
slightlycurledwhen dry, erect patent when wet,
6 mm long, broadly lanceolate, serrate at tips,

widest at midleaf and contracted towards leaf
base. Costa filling 1/2of the leaf width, in transverse section with ventral hyalocystsand dorsal
groups of stereids, excurrent.Alar cells weakly
differentiatedor not, often reddish with hyaline
margins.Basallaminalcells thin-walled,hyaline,
6-9 x 40-50 gm. Upper laminal cells small,
subquadrateto short rectangular,5-6 x 9-12
,um.
Distribution(Fig. 119). Usually on sandstone
rocks in shady places, rarely on rotten wood,
confined to SE Brazil.
172
Flora Neotropica
Specimens examined. BRAZIL. PARANA:Parque
Nacional Vila Velha, Frahm 1392 (hb. Frahm). Rio

GRANDE
DO SUL: Sao Leopoldo, Sehnem s.n. (hb.
Frahm);Vacaria, Sehnem 5962 (LBLC).Sao Paulo:
AlongroadfromItirapinato Brotas,Vital2069 (SP).

Withrespectto the areolationandthe structure
of the costa, C. sehnemii stronglyresembles C.
incrassatus C. Milller, a species widespread
throughthe subantarctic.Campylopussehnemii
differs by longer leaf tips and the lack of hairpoints, which is perhapsdue to the differenthabitat on shady rocks in forests, whereas C. incrassatusgrows in open habitats. Therefore,C.
sehnemii is supposedly derived from the subantarctic C. incrassatus.This has a parallel in
SouthernAfrica and Australia,where, at about
the same latitude as C. sehnemii, C. incrassatus
is replacedagain by a closely relatedspecies, C.
catarractilis(C. Miller) Paris.
53. Campylopussharpii J.-P. Frahm,Horton &
Vitt, Bryologist82: 623. 1979. Type. Mexico.
Oaxaca: Near S. Pedro Yolox, 17?36'N,
96?24'W, Vitt 17715 (holotype, ALTA; isotype, TENN).
Figs. 119, 120.
Plantsin loose, gray-greentufts,to 3.5 cm high,
radiculosebelow. Leaves appressedbelow, longer and somewhat curled when dry above, 7-8
mm long, with bases shiny-whitish, slenderly
long-pointed from an ovate-lanceolate base,
slightlytoothed at the extremeapex. Costaabout
2/3of the leaf base, excurrent;in section showing
largeventral leucocystsand small dorsal groups
of 1-2 substereidalcells. Alar cells red-brown,
thin-walled.Innerbasal cells lax, hyaline, 40-48
x 7-9.5 ,m, the outercells small,elongate,forming a border of 6-8 rows. Upper laminal cells
elongate oval (about 4:1), 17-21 x 5 ,m, extending about 2/3along the costa in 2-3 rows.
Sporophytesunknown.
Distribution(Fig. 119). On soil, in soil covered
rocks and epiphytic in montane rainforest, in
Mexico also on oak forests,in altitudesbetween
2500 and 3000 m in Peru, Colombia,Venezuela
and Mexico.
Specimens examined. COLOMBIA. MAGDALENA:
ParqueNacional SierraNevada de SantaMarta,RanMun. de Sta. Rosa

gel et al. 377 (FLAS).RISARALDA:
Cabal,entraHda.La Sierray Termalesde Sta.Rosa,

van Reenen 1070 (U).
VENEZUELA. MERIDA:
Sierra Nevada de Santo
Domingo,LagunaBrava,L6pez-T.15431(FLAS).
ECUADOR. NAPO: Km 214 Quito-Baeza road,
Steere27059 (NY).
PERU. AMAZONAS:
Along road from Chachapoyas
to Cajamarca above Leimebamba, Frahm, Camp. Peruv. Exsicc. 22 (ALTA, B, BM, FLAS, NY, S, U).
54. Campylopusshawii Wilson in Hunt, Mem.

Lit. Soc. Manchesterser. 3, 3: 234. 1868. Type.
Outer Hebrides.Shaw s.n. (holotype, BM).
Figs. 121, 122.
CampylopusbrittonaeWilliams,N. Amer. Fl. 15: 141.
1913.Type.Jamaica.Summitof St. John'sPeak,

Britton1909(holotype,NY; isotypes,DUKE,FH,
H. S).
CampylopuscarreiroanusCardot,Bull. Herb. Boissier
ser.2, 5: 202. 1905.Type.Azores.SanMiguel,Carreiro689 (holotype,PC).
Campylopusunderwoodii
Williams,N. Amer.Fl. 15(2):
142.1913.Type.Jamaica.
SummitofBlueMt.Peak,
Underwood
2539(holotype,NY;isotypesF, H-BR).
Plants tall, to more than 10 cm high, in yellowish green, loose tufts. Stems equally foliate,
tomentose below. Leaves 10-15 mm long, gradually narrowedinto a long, fine, nearly smooth
or slightly serrulatepoint, spreadingor attached
to the stem. Costa fillingmore than 2/3 of the leaf
base, excurrent,in transversesection with large
ventralhyalocystscoveringmore than half of the
transversesection and dorsalgroupsof stereids,
smooth at back. Alar cells conspicuous,forming
large auriclesprotrudinginto the costa, reddish
inside and hyaline outside. Basal laminal cells
incrassate, short rectangular,often slightly pitted, 5-10 x 50-65 ,um, narrowerand paler at
margins. Upper laminal cells small, incrassate,
oval to oblique, 3-6 x 19-26 um.
Seta to 15 mm long, arisingfrom short pseudolateralbranches,sinuose or curved, yellowish
to light brown. Capsule2.5-3 mm long, curved,
strumose,nearlysmooth or furrowedwhen emptied, olive to dark brown. Peristome dark red.
Operculumshortly rostrate,half as long as the
capsule. Calyptraciliate at base.
Distribution(Fig. 122). On soil, rocksand rotten wood in pinecloudforestsandbroadleafcloud
forests between 1300 and 2700 m elevation in
Cuba,Jamaica,PuertoRico and Hispaniola(Dominican Republic), also on the Azores and the
highly oceanic parts of the British Isles.
Specimens examined. CUBA ORIENTE:Pico Turquino, Mald s.n. (MO).
DOMINICAN REPUBLIC. LA VEGA:Vic. La LaLa Nevera
gunita, Norris 5730 (BP, H-BR). PERAVIA:
173
FIG. 120. Campylopussharpii(Vitt 17715, ALTA). A. Plant. B. Leaf.C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Am.
Area 47 km S of Constanza,18?39'N,70?34'W,Reese
15717 (NY).
JAMAICA. ST. THOMAS
PARISH:
Blue Mountain
Peak 1803'N, 76035'W, Crosby3387 (MO); between
PortlandGap and BlueMountainPeak,Hegewald8081
(hb. Frahm).
The disjunction between the Caribbean Islands and the British Isles, connected by the occurrence on the Azores, seems to be remarkable.
This distribution pattern is, however, met also
in other bryophytes such as Cyclodictyon laete-
virens(Hooker& Taylor)Mitten,Adelanthusdecipiens(Hooker)Mitten, Leptoscyphuscuneifolius (Hooker)Mittenand others.In contrastto the

CaribbeanIsland, Campylopusshawii grows in
the BritishIsles in swamps.Generally,it may be
possible that tropical montane bryophyteshave
been broughtto Europeas packingmaterialfor
tropical flowering plants for greenhouses and
parks, from where they escaped and found an
ecologicalniche in the naturalvegetation.However, this possibility must be excluded for C.
174
Flora Neotropica
9.5 mm
'
/.//
4cm 2
.A
l.
f C
C. Leaf-tip. D. Transverse section

121.
br for microscopic
m
Uperwood
lainal cells.
cellsNY).
A.Unlled scale bar
ofofleaf. E.
Basal laminal cells. F. Upper
Unlabeled
details = 50 ,m.
laminal
shawii, because in the British Isles there occurs

a closely related species, C. setifolius Wilson,
which is not found in the Caribbean.This species
may have been derived from C. shawii by occupying a different habitat, such as wet rocks.
The structuraldifferences between the species
thereforeinvolve specialadaptationsto these different habitats, such as large ventral hyalocysts
with lax cell walls in transverse section of the
costa in C. shawii(occurringin swamps)for storing water,and small ventralhyalocystswith firm

walls in C. setifolius(occurringon wet rocks) as
protectionagainstdesiccation.Therefore,the occurrenceof C. shawii in WesternEuropemay be
interpretedas a relict,wherethis species has survived glaciation periods in unglaciatedhabitats
along the west coast of Irelandand Scotland or
regions now drowned by the postglacial rising
175
80
70
60
50
I~~~~~
~~~30
o
..
0Cb20040
00km
00
60
--- - ..........!--
~~~~~
^
~
__ lb
_'^__
i\
1-^
~~ ~~
~~~~
20
O0O
0
--
200
100
400
200
600
300
400
800
1OOOkc
500
600
mI..
'
',
-..
FIG. 122. Distributionof Campylopusshawii (@)and C subcuspidatus(0).
sea level. Sporophyteshave been found rarelyin

the CaribbeanIslands but never in Europe.
Campylopusbrittonaeand C. underwoodiiwere
both describedby Williams from Jamaicain the
same publication. According to Williams they
differ in the length of the leaves, which seems
insufficientto separatethe taxa.Bothspeciesshow
absolutelyno differencesin leaf anatomy,but the
type of C. underwoodiishows smallerplants,with

leaves more or less appressedto the stem, whereas the type of C. brittonae has patent leaves.
Therefore,C. underwoodiican only be regarded
as a modificationof C. brittonae.It is not known
whether such "underwoodii-expressions"indicate a differenthabitat. This seems possible, for
patent leaves in rain forest species of Campylo-
Flora Neotropica
176
pus may be interpretedas an adaptationto poor narrowerat margins, borderedby 4-6 rows of
nutrients in the substrateand a nutrition pre- elongatehyaline cells. Upper laminal cells elondominantly from atmosphericsources, by siev- gate oval, 3-6 x 6-13 um, stronglyincrassate,
ing out organic particles falling down from the slightly pitted.
canopy.
Distribution(Fig. 122).On humic, wet soil and
55. Campylopussubcuspidatus(Hampe)Jaeger, rotten wood in montane rain forests, on peaty
Ber. Thitigk. St. Gallischen Naturwiss. Ges. soil and in swampsbetween600 m (in the south1870-1871: 441. 1872.
ernmost part of the range from sea level) and
3000 m in SE Brazil, the Guianas, Venezuela,
Costa Rica, Hondurasand Puerto Rico.
Specimensexamined.COSTA RICA.HEREDIA:CeCampylopus subcuspidatus
rrosde ZurquiNE of SanIsidro,Standley& Valerio
1 Leaves 8-12 mm long, erect patentor spreading;
upperlaminal cells 4-6:1. ...................
55a. var. subcuspidatus.
.......................
1 Leaves4 mm long;stems appressedfoliate;upper
55b. var. damazii.
laminal cells 3-4:1. ..........
50511, 52141 (NY). SANJOSE:17km S ofEl Empalme,

Crosby 10886 (MO); Cerro de las Vueltas, Standley
43685 (NY).
PUERTORICO.El YunqueRecreation
Area,trail
fromMt.Brittonto summitof ElYunque,Buck4053
(NY).
Cerro Marahuaca,
VENEZUELA. AMAZONAS:
55a. Campylopussubcuspidatus(Hampe)Jaeger 3?35'N,65?20'W,Guarigliaet al. 1655 (NY). BOLIVAR:

var. subcuspidatus Dicranum subcuspidatum Auyan-tepui,
94055(NY);cumbrede CeSteyermark
Hampe, Vidensk. Meddel. Dansk Naturhist. rro Guaiquinima,Steyermark& Dunsterville113558
Foren. Kj0benhavn 1870: 273. Type. Brazil. (NY).
GUYANA.Upper MazaruniDistr., Mt. Latipu,
Rio de Janeiro: Tijuca, Glaziou 7096 (holo5606(U);N slopeof Mt.
5?57'N,60?38'W,Gradstein
type, BM: isotypes, NY, PC). Figs. 122, 123. Roraima,5?16'N,60?43'W,Gradstein
5350(U).
Florschiitz7136
SURINAM.Wilhelminagebergte,
CampylopusbrotherianusParis, Ind. bryol. Suppl. 89. (U).
1900, nom. nud. Materialis the same as in the type
4218
Cremers
FRENCHGUYANA.PicCoudreau,
of C. procerus.
Onraedt).
(hb.
152.
Bowers,
Bryologist77(2):
Campylopusparamoensis
1974, nom. nov. pro C. atratusBartram,Contr.U.S.
Natl. Herb. 26: 61. 1928, hom. illeg. Type. Costa
Rica. Cerrode las Vueltas 3000 m, Standley43686
(holotype,FH; isotypes, H-BR, S).
Campylopuspraealtus (C. Miiller) Paris, Ind. bryol.
Suppl. 96. 1900. Dicranumpraealtum C. Miller,
Hedwigia37: 227. 1898. Type. Puerto Rico. Sierra
de Luquillo,Sintensiss.n. (holotype,destroyedat B;
lectotypusnov., NY; isolectotypes,H-BR, MO).
CampylopusprocerusBrotherusin Paris, Ind. bryol.
Suppl. 89. 1900. Type. Brazil.Rio de Janeiro:Pico
de Papageio, Ule, Bryothec. bras. 110 (holotype,
H-BR).
BRAZIL.Sio PAULO:
JaraguanearTaipas,Schiffner
2677 (H-BR);Ubatuba,Serrado Mar,MakinoH-8

(SP).
Campylopus subcuspidatusmorphologically
resembles C. cuspidatus,which can be distinguished by collenchymatousalar cells, the presence of a hair tip, and incrassate, vermicular,
pitted laminal cells throughoutthe whole leaf.
Perhaps due to the lack of sporophytesthis
species has a scattereddistributionin SE Brazil,
the Guianamassif, CentralAmericaand a single
locality in Puerto Rico. The sole record in the
Plants very tall, to more than 10 cm high, very Caribbeanmay go backto a secondaryextension
robust, Dicranum-like, in loose tufts, green to of its range. It is closely relatedto Campylopus
golden-yellowish, shining, darker brownish or shawiiand differsfrom this speciesony by sharpblackish below. Stems tomentose below, erect, ly serrateleaf tips, elongate oval upper laminal
rarely branched,equally foliate with appressed cells and longer basal laminal cells. Since both
or erect spreadingleaves. Leaves about 15 mm rangesare not (exceptfor this singlerecordfrom
long,graduallynarrowedinto a long serratepoint. Puerto Rico) overlapping,C. shawii may be inCosta filling 2/%-3/of the leaf base, in transverse terpretedas a geographicalvicariant species of
section with largeventral hyalocystsand dorsal C. subcuspidatusin the Caribbean.
stereids, smooth at back, excurrent.Alar cells
reddish, conspicuous,protrudinginto the costa. 55b. Campylopussubcuspidatus(Hampe) JaeBasallaminalcells longly rectangular,5-6 x 32gervar.damazii(Brotherus)J.-P.Frahm,Bryol.
Beitr. 7: 75. 1987.
50 tm, incrassate,with pitted walls, shorterand
177
ii
FIG. 123. Campylopussubcuspidatusvar. subcuspidatus(Buck4053, NY). A. Plant. B. Leaf.C. Leaf-tip.D.

details = 50 ,um.
CampylopusdamaziiBrotherus,Denkschr.Akad.Wiss.
Wien math.-nat. KI. 83: 260. 1926. Type. Brazil.
Minas Gerais: Pico da Serrade Itabira,Damaziou
2147 (holotype, H-BR).
Differs from var. subcuspidatus by shorter
leaves, which are appressed to the stem, and
(probably caused by the leaf length) shorter upper
laminal cells.
Distribution. Known only from the type locality.
The taxonomic position of this variety is not
clear, but the study of this problem is difficult,

due to the lackof any othercollectionresembling
var. damazii.
56. Campylopus subjugorum Brotherus, Bot.
Jahrb.49:174. 1912. Type. Bolivia. Cordillera
Real, Knoche60a (holotype, H-BR).
Figs. 124, 125.
Plants similarto those of C. nivalisbut in more
dense tufts with appressedfoliate stems, upper
Flora Neotropica
178
A
FIG. 124. Campylopussubjugorum(CampylopodesPeruvianaeExsiccatae 24). A. Plant. B. Leaf. C. Leaftip. D. Transverse section of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for
microscopicdetails = 50 Jim.
laminal cells more rectangular than oval and costa distinctly ridged at back in the upper part with
short lamellae one cell high.
Distribution (Fig. 125). At high elevations in
the Andes of Venezuela, Colombia, Ecuador, Peru
and Bolivia from 3800 m up to 4900 m.
Griffinet al. 1261 (FLAS);Sierrade Santo Domingo,

Paramode Mucubaji,Griffinet al. 1176 (FLAS).
ECUADOR. NAPO:Quito-Baeza road, Pframo de
Guamani, Ollgaard& Balslev 10163 (NY).
PERU. ANCASH:
CordilleraBlanca,pass Carnicero,
Kinzl s.n. (JE); pass between Huanuco and Huaraz,
Frahm, Camp. Peruv.Exsicc. 24 (ALTA, B, BM, F,
Specimensexamined.COLOMBIA.ARAUCA:Sierra
Nevada del Cocuy, Cleef8917, 2125a (U).
Hegewald9081, 9087 (hb. Hegewald).
VENEZUELA.
MERIDA: Above Pico del Aguila,
Between Tambo and Aina,

NY, S, U). AYACUCHO:
BOLIVIA. COCABAMBA:Base of Serranias Taru-
cani, 17?14'S,66?24'W,Lewis 79-2468A (F). LA PAZ:
179
90
80
70
60
50
30
I^|.,.y
--
200
400
600
800
'600
100 200 300 40 500
CT^
<^=K/^
*.
-~~~~-~~~---20
j]
1000k-<
m s
Prepared by Hendrik R.Rypkema,
FIG. 125.
:_
Distribution of Campylopus subjugorum (0) and C surinamensis (0).
Between Abra Huallata and Millipaya, 15?51'S,

68'39'W, Lewis 79-1370 (F); 5 km NE of Milluni,
16?18'S,68?06'W,Lewis79-1670A (F); 1 km S of Puer-
to Acosta, Lago Titicaca, 1532'S, 691 5'W, Lewis 79-
719 (F); Escoma, N side of Lago Titicaca, 15?38'S,

69"9'W,Lewis 79-784 (F).
57. Campylopus surinamensis C. Miller, Linnaea 21: 186. 1848. Type. Surinam. Near Paramaribo, Kegel 516 (holotype, destroyed at B;
lectotypus nov., H-BR; isolectotypes, GOET,
PC).
Figs. 2B, 125-127.
CampylopuscatumbensisBrotherus,Bih. Kongl.Svensk
Vetens. Akad. Foerh. 21 Afd. III(3): 8. 1895. Type.
Brazil. Rio de Janeiro:Catfimbez,Mosen 190 (holotype, H-BR; isotype, S).

CampylopuscostaricensisBartram,Contr. U.S. Natl.
Herb. 26: 62. 1928. Type. Costa Rica. Finca Montechristo,Standley& Valerio48579 (holotype, FH;
isotype, US).
Campylopusdonnellii (Austin) Lesquereux& James,
Man. Moss. N. Amer. 79. 1884. Dicranumdonnellii
Austin,Bot. Gaz.4:150. 1879.Type.Florida.Smith,
MusciAppal. 470 (holotype,NY).
CampylopusgracilicaulisMitten,J. Linn. Soc. Bot. 12:
83. 1869. Type. Brazil. Rio Negro, Spruce60 (lectotype, NY; isolectotypes,BM, FH, NY, PC, S).
CampylopusmarmellensisBrotherus,Hedwigia45: 262.
1905. Type. Brazil. Amazonas: Rio Madeira, Ule
246 (lectotype,H-BR; isolectotypes,FH, JE, NY).
180
Flora Neotropica
;iill'li,ril.- ilt!ti1 laxissilllis fuscidulis SelCLeULuc Cu;...l .. .
-ii vuitllcnl leveCn congestis; perlch. multo Ilmajra latiora basi
v.i;ilal;t vel couvuluta, laxissime et awuplissime reticulata, subula
cluniigta flcxuosa grossius serrulata coronata. Caetera ignota.

I' a t r i a. Alabama, Mobile, raro sed semper sterile: C.Mohr.
D. leucogaster Mexicanum proximum raptim distinguitur:
caule magoisflexuoso crassiore multo longius setoso, cellulis alaribus omnino albidis laxioribus, -eateris minoribus,
.
FIGS~~~~~~~~~~.
FIG. 126. Campylopussurinamensis(lectotypeof C subleucogaster,NY).

CampylopusminarumParis,Ind. bryol. ed. 2, 1: 318.
1904, nom. nud. Materialis the same as in the type
of Dicranumlaxobasis.
DicranumlaxobasisC. Miller, Hedwigia39: 252. 1900.
Type. Brazil. Minas Gerais: Serra de Caraca, Ule
1353 (holotype,destroyedat B; lectotypusnov., HBR).
Campylopussubleucogaster(C. Miiller) Jaeger, Ber.
381. 1880. Dicranumsubleucogaster

C. Muller,Bull.
TorreyBot. Club 5: 49. 1879. Type. Alabama:Mobile, Mohr1868 (holotype,destroyedat B;lectotypus
nov., NY).
Plants to 3 cm high, with a rosette of basal
leaves, appressed stem leaves and a terminal comal tuft, yellowish green. Stems erect, not
seci; tgla.E aallmnl
181
cel.F
pe
aia
elUlble
cl
a o
irsoi
eal
50~..
cm..
;...:,
FIG. 127.
Campylopus surinamensis (Florschfitz 4790, U). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
50 im.
branched.Leaves about 5 mm long, lanceolate,

ending in a serratetip. Costa filling /2 of the leaf
base, in transverse section with ventral hyalocystsand dorsalgroupsof stereids,ridgedat back,
ending in the leaf tip or excurrent,especially in
comal leaves. Alar cells weakly developed, hyaline or brownish. Basal laminal cells rectangular, incrassate, 10-16 x 32-60 ,um,narrowerat
margins. Upper laminal cells short rectangular,
5-6 x 15-20 Am,varyingin length between 1:2
and 1:4,rarelyto 1:6.Involucralleaveswithbroad

ovate base, suddenlycontractedinto a long fine
stronglydenticulate subhyaline point, with hyaline thin-walledbasal laminal cells.
Seta about 9 mm long, sinuose or curved.Capsule curved, 1.5 mm long, asymmetric,brownish
as the seta. Operculumlongly rostrate.Calyptra
ciliate at base.
Vegetative propagationby small hooked propaguliferousleaves in the comal tufts.
Flora Neotropica
182
Distribution (Fig. 125). On open acidic sandy
soil, rarely on rotten wood, especially along rivers or in savannah vegetation in the Amazon
lowland up to 100 m elevation in Brazil, extending to the Guianas, Venezuela, Peru and Bolivia, in SE Brazil from sea level up to 1500 m.
Also collected in Cuba and Honduras. Frequent
in the coastal lowland of SE United States.
MonSpecimensexamined.HONDURAS. TOLEDO:
key River, Gentle3840 (MICH).
CUBA.PINAR
DELRio: La Mulata,Borhidi&Duarte
5439 (EGR);Rio SantaRita,Hazen 12238 (NY);Loma
SanJuan,Horeaus.n. (NY);SantCoblacion,El Valdes,
Samek et al. s.n. (PRC).
VENEZUELA. AMAZONAS:
Cucuritalde Caname,
67?22'W,3?40'N, Davidse et al. 17017 (MO); Cassiquiare, v. Liitzelburg22589a (JE).
TRINIDAD.Valencia,Toco Road,Brittonet al. 1815
(NY).
GUYANA. EASTDEMERARA:
Dakara Creek, 6?28'N,
East
58?15'W, Gradstein 4735 (U). UPPERMAZARUNI:
bank of Waruma river, Gradstein 5077 (U); Upper
Mazaruni River, Leng 329 (NY).
SURINAM. Moengotapoe,Arnoldo3502 (U).
PERU. LORETO:Quistococha near Iquitos, Hege-
wald 6366 (hb. Frahm);LowyPBR 209 (LAF).
BRAZIL: AMAZONAS:
Campus do INPA, Griffin et
48?4'W,Davidseet al. 17716 (NY). PIAUi:ParqueNa-
cionalSeteCidades,Vital5396(SP).RONDONIA:
Santa
120kmS of PortoVelho,9?10'S,6007'W,
Barbara
Fife
et al. 4215 (NY);firstrapidsof Rio PacfasNovos,
SW of
11?S,64?W,Reese 13610 (NY). Sio PAULO:
JuquiaalongSP 116,Frahm1593 (hb. Frahm);Estaci6nBiol6gicaMogiGuacu,Vital6985 (SP);Serra

do BocainaNE Camposda Cunha,Frahm1591(hb.
Frahm);km 209 betweenRegistroand Jacupiranga,
24032'S,46053'W,Vitt20753 (ALTA);Rio Javarynear
Esperanca,
(NY);CampoGrande,SchiffKrukoff7530
ner654, 2158 (NY).
BOLIVIA. BENI:27 km NW, 5 km W, 6 km S of
Reese12817,13003,12990(NY).
Guayaramerin,
Exceptfor SE Brazil,Campylopussurinamensis is a lowland species. Recordsfrom the Andes

of Peru concerneither C. capitulatusor C. pauper, which resemble C. surinamensis morphologicallywith appressedfoliate stems and comal
tufts. Recordsfrom Uruguaybelong to C. introflexus, in which female plants have a similarappearancewith comal-tuftedperichaetia.
Young plants, which consist only of a basal
rosette and do not show this characteristichabit
with slender stems and pencil-like comal tufts
can be difficultto identify.
al. 52265 (INPA);km 60 EstradaManaus-Caracarai,

Lisboa 50936 (INPA); Estrada Maua, Prance et al.
30045 (INPA);Rio Negro, boca do Rio Apuai,Lisboa 58. Campylopus tallulensis Sullivant & Les51749 (INPA);Rio Cueiras,Lisboa54937 (INPA);Rio
quereux, Musci bor.-amer. 17. 1865. Type.
Negro, Cachoeira, Prance 15802 (INPA); Manaus,
U.S.A. Georgia:TallulahFalls, Small s. n. (hoPonteNegra,Frahm1369 (hb.Frahm);Rio Negronear
lotype,
NY).
Figs. 128, 129.
Manaus,Armitage 153 (NY); km 10 along road Manaus-BoaVista, Frahm 1524 (hb. Frahm);Rio Negro,
Campylopustequendamensis
Th6riot,Rev. Bryol.
Cucuhy, v. Liitzelburg22879 (B, JE); Rio Icana, v.
Lich6nol.11: 58. 1939.Type.Colombia.Tequen22974
Liitzelburg
(JE);Ducke forest reserve26 km N
dama,Troll2036(holotype,PC).
of Manaus, Griffinet al. 171 (FLAS).BAHIA:15 km
NW of Leucois along BR 242, Boom & Mori 1087
Plants in yellowish green to brownish green
(NY); Mun. de Una, 30 km S of Olivenca, 15?12'S, tufts. Stems to 5 cm tall,
slender,equallyfoliate,
39?3'W,Lewis & Carvalho723 (hb. Frahm).ESPIRITO
SANTO:
Along BR 262 near Viana, Schafer-Verwimp without tomentum. Leaves about 5 mm long,
8865 (hb. Frahm).GOIAS:
Mun. de Cristallina,17?8'S, narrowlylanceolate,ending in a narrow,serrate
Mun. de tip. Costa filling 12-/3of the leaf base, ridged at
47?44'W, Vital 8284 (SP). MATOGROSSO:
Bataguacu,Vital2192 (SP);Mun. de Barrado Garcas, back, excurrent,in transversesection with large
Vital1374 (SP);alongBR 364 betweenAlto Garcaand
Alto do Aragua,Schafer-Verwimp8652 (hb. Frahm); ventralhyalocystsoccupying1/2the height of the
Chapadados Guimaraes,Schdfer-Verwimp8564 (hb. section and dorsalgroupsof small stereids.Alar
Frahm). MINAS GERAIS:Along road MG 55 between
Morro do Pilar and Sao Sebastiao,Frahm 1592 (hb.

Frahm). PARA:Mun. de Braganca,13 km N of Braganca, 0?59'S, 46?15'W,Davidse et al. 18038 (NY);
Serrado Cachimbo,km 780-820 on Cuiabi-Santarem
highway, 9?22'S, 54054'W,Reese 16122 (NY); Cataractson Rio Curua8?45'S,54?57'W,Reese 16486 (NY);
763 km N of Cuiaba, 9035'S,54?55'W,Reese 16059
(NY); km 774 on Cuiaba-Santaremhighway,9030'S,
54?55'W,Reese 16039 (NY); Mun. de Oriximina,rio
Trombetas,Ramos & Rosas 1868, 1946 (NY); Mun.
de Vigia, ca. 9 km SE of Vigia along PA 140, 0?55'S,
cells hardly developed. Basal laminal cells hyaline, thin-walled,rectangular,4-10 x 13-38 ,um,
not differentiatedat margins.Upperlaminalcells
shortrectangular,incrassate,3-6 x 6-13 gm, ca.
1:4.
Distribution(Fig. 129). On soil and treetrunks
in montane forests in Mexico, Guatemala,Nicaragua,Venezuela,Colombia,Ecuador,Peruand
Bolivia, also disjunctin SE United States,in the
183
3.5mm
.'m /
D'"'
FIG. 128. Campylopus tallulensis (Delgadillo 3055, MEXU). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
50 Mm.
Andes in evergreen rain forests between 2500

and 3000 m, in Mexico in pine-oak forests between 2000 and 2500 m and in the Appalachians
between 100 and 1200 m.
et al. 4240b (TENN); 67 km N of Ixtlan de Juarez,

Norris& Taranto16199 (TENN);alrededoresdel Cerro de San Felipe NE de Oaxaca, Delgadillo 247b
(TENN). SANLUISPorosi: 44 miles W of Antiguo
20 miles
Specimensexamined.MEXICO.DURANGO:
N of Las Nieves on border to Chihuahua,Bowerset
al. 6298b (TENN). JAuSCO:Hwy. 80 above Autlan,
del Cielo above Gomez Farias,Iwatsuki& Sharp4864

Cima del Cofre de Perote, Del(TENN). VERACRUZ:
gadillo 3055 (MEXU); between Perote and Jalapa,
Frahm, Camp. Exsicc. 13 (ALTA, B, BM, C, FLAS,
H, NICH, S, U).
Sharp et al. 3187 (TENN). MICHOACAN: E of Las
Perason Hwy. 15, Norris& Taranto15646a (TENN).

NAYARM:
Mun.de Compostela,Norris& Taranto15201
N of Oaxaca on Hwy. 175, Sharp
(MICH).OAXACA:
Morelos, Reese 5250 (TENN). TAMAUUPAS:Rancho
GUATEMALA.
Fuentes GeorQUEZALTENANGO:
ginas, W slope of Volcan Zunil, Standley 67371 (F,

FH);Chiquimula,VolcanIpala,Steyermark30608 (F):
184
Flora Neotropica
rO
90
a(1
*0
20
...,_A
a0*;l
P~~~.d ~
70
80
60
80,06
so0
o*3
..............<{
Distnb~ito o
tallni
@ n
rcylpao
20
NICARAGUA. MATAGALPA:
Hacienda Santa Maria
de Ostuma,Almeda 1453d (MO).

San Pedro, Onraedt 10442
(hb. Frahm);vicinity of Medellin,Charetier149 (NY).
NW de Arcabuco, Cleef et al. 3506 (U).

SANTANDER:
VENEZUELA. AMAZONAS:Cerro Marahuaca,
3?37'N, 65?21 'W, Maguire et al. 65676 (NY). MERIDA:
ParamoLa Negraabove Bailadores,Grifin et al. 2175

(FLAS).
ECUADOR.NAPO:Km 40-46 from El CarmelotowardsLaBonita,77030'W,0?34'N,Lojtnantet al. 12218
(AAU).
PERU. AREQUIPA:Lomos de Atiquipa, Hegewald
8574 (hb. Frahm).HuANuco:Above Huallancaalong

road Huanuco-Huaraz,Frahm, Camp. Peruv.Exsicc.
25 (ALTA, B, BM, FLAS, H, NY, S, U).
59. Campylopus trachyblepharon (C. Muller)
Mitten, J. Linn. Soc. Bot. 12: 80. 1869. Dicranum trachyblepharon C. Muller, Syn. musc.
frond. 1: 389. 1848. Type. Brazil. Beyrich s.n.

(holotype, destroyed at B; lectotype, NY).
Figs. 129, 130.
Campylopusarenicola(C. Miller) Mitten,J. Linn.Soc.
Bot. 12: 77. 1869. Dicranum arenicola C. Muller,
Bot. Zeit. 13: 762. 1855. Type. Brazil.Itajahi,Pabst
s.n. (holotype,destroyedat B; lectotypusnov., NY;
isolectotypes,GOET, H-SOL, JE, S).
CampylopusbermudianusWilliamsin Britton,J. New
YorkBot. Gard.13:193. 1913. Type. Bermuda.Paget Marsh, Brown 651 (holotype, NY; isotypes,
CANM, DUKE).
Campylopusdetonsus(Hampe)Kindberg,Enum.Bryin.
exot. 88. 1889. DicranumdetonsumHampe, Flora
64: 341. 1881. Type. Brazil.Rio de Janeiro,Glaziou
11746 (isotypes, H-BR, JE, S).
CampylopusschwaegricheniiDuby, Mem. Soc. Phys.
Geneve 20: 360. 1870. Type. Brazil. Porto de Estrelle, Beyrichs.n. (holotype,G).
CampylopussubarenicolaC. Miller ex Brotherus,Bih.
185
2mmn
I-l^^Bt~r
lH
:u
1f
6.5mm
?o?
FIG. 130. Campylopustrachyblepharon

(CampylopodesBrasiliaeExsiccatae 16). A. Plant. B. Leaf.C. Leaftip. D. Transversesection of leaf. E. Basal laminal cells. F. Upper laminal cells. G. Capsule.H. Operculum.
Unlabeled scale bar for microscopicdetails = 50 Am.
Kongl.SvenskVetens.Akad.Forh.21,3(3): 7. 1895.
Type. Brazil.Santos,Mostn 94 (holotype,destroyed
at B; lectotypusnov., H-BR).
Campylopusvillicaulis(Hampe) Jaeger,Ber. Thitigk.
St. GallischenNaturwiss.ges. 1877-1878:381. 1880.
DicranumvillicauleHampe, Vidensk. Meddel. Naturhist.Foren.Kjoebenhavner. 3,4:44. 1872. Type.
Brazil.Rio de Janeiro,Glaziou4680 (isotype,H-BR).
Plants in dark green loose tufts, lighter at tips,
to 7(-16) cm high. Stems tomentose below, sev-
eral times interruptedlyfoliate, ending in a comose tip. Leaves 6-7 mm long, serrate in the
upperthird, narrowedinto a serratepoint. Costa
taking 1/3of the leaf base, ending in the leaf tip
or shortly excurrent,in transversesection with
ventral and dorsal stereids, lamellose at back in
the upperpart of the leaf with lamellae 2-3 cells
high. Alar cells large,hyaline or brownish.Basal
laminalcells incrassate,elongaterectangular,10-
186
Flora Neotropica
13 x 12-32 ,m, subquadrateat margins.Upper

laminalcells incrassate,subquadrate,6-16 x 1319 ,um,reachingnearly the apex.
Seta to 1 cm long, lightto darkbrown.Capsule
asymmetric,furrowedwhen empty, light brown,
dark brown in age. Operculumobliquely and
longly rostrate,darkeras the capsule. Calyptra
ciliate at base.
Distribution(Fig. 129). On bareopen wet sand
on coastal areas near sea level, very frequentin
restingasof SE Brazil,rarelyalong the Brazilian
coast to the Guianasand collectedin one locality
on Bermudabetween 1908 and 1916.
Specimensexamined.GUYANA. WithoutLocality:
soil. C. trachyblepharonis, however, distinguished by quadrateupper laminal cells, a lamellose costa and the lack of an excurrentsubhyaline point in the comal leaves. In the field,
all possibleintergradationscan be foundbetween
such small forms on open places and tall forms
up to 10 cm high under restingabushes.
Campylopustrachyblepharonoccurs also in E
Africa, also in coastal areas on sand and at the
same latitude, as a closely related,geographical
vicariant subspecies comatus (Renauld & Cardot) J.-P. Frahm.This subspeciesdiffersonly by
ventral hyalocysts in transverse section of the
costa and shorterdorsal lamellae.
Abraham 145 (NY). EASTDEMERARA:

Timehri, Daka-
raCreek,6?29'N,58?15'W,Gradstein3734 (U);Demerara-MahaicaRegion, 1 km E of Soesdyke, 6?30'N,

58?11'W,Pipoly 9290, 9296, 9305, 9287, 9289 (NY);
16 mi S of Georgetown,6?20'N,58?15'W,Pipoly9211
(NY).
BRAZIL. BAHIA:Mun. Entre Rios, 2-5 km W of

Subauma,Boom & Mori 990 (NY); Mun. de Ilheus,
road from Olivenca to Una, Boom & Mori 707 (NY);
Mun.de Salvador,vic. airport,Boom &Mori913 (NY);
Mun. de EntreRios, 2-5 km W of Subauma,Boom &
Mori 989 (NY);km 20 alongBR 101 betweenEsplanada and Jandeira,Schafer-Verwimp8717 (hb. Frahm).
Mun. de Picuna,Lagoada Piabanha,
ESPRITO
SANTO:
Vital2849 (SP);Mun. de Aracruz,2 km E of Barrado
Maua, GlaRiacho, Vitals. n. (FLAS). Rio DEJANEIRO:
ziou 7474 (PC). SANTACATARINA:
Jaguarone, Ule 709
(H-BR). Sio PAULO:Campo Grande, Loefgrens.n.

(JE);St. Vicent,nearSantos,Horeaus.n. (H-BR);Concepciao,Krypt.Exsicc. 2675 (B, F, H, HBG);Palmeiro
de Sao Lorenzo,Schiffner1440 (H-BR);Bertioga,along
road to Guaratuba,Frahm 1483 (hb. Frahm);between
GuaratubaandSaoSebastiao,Frahm1481 (hb.Frahm);
between Peruibe and Itanha6m, Frahm 1480 (hb.
Frahm);Cananeia,Ilha do Cardoso, Frahm, Camp.
Bras. Exsicc. 16 (ALTA, B, C, DUIS, FLAS, GOET,
GZU, H, HBG, INPA, JE,KR, M, MO,NFLD, NICH,
PC, S, SP, TENN, U); 2 km N of Peruibe, 24?13'S,
46?53'W, Vitt20687 (ALTA).
The localitiesin the Guianasand Bermudaare

very isolated from the main rangein SE Brazil.
The specimens are not as well developed and
especially do not show the verticillate foliation
as distinctlyas in SE Braziland, as shown by the
occurrencein Bermuda,the recordsseem not to
be persistent.Thus these northernrecordsmay
be causedby sporeor fragmentdispersalby birds
along migrationroutes.
Young unproliferatedplants have been described as Campylopus arenicola. They were
placed into synonymy with C. surinamensis,

which occurs on similar habitats on bare sandy
60. Campylopus trichophylloides Theriot in

Herzog, Repert. Spec. Novi Regni Veg. 95.
1938. Type. Peru.CordilleraBlanca,Kinzls.n.
(holotype,JE; isotype, B).
Figs. 131, 132.
Plants in dense tufts, light green to brownish
green,to 3.5 cm high. Stems equallyfoliate with
appressedleaves, slender, filiform. Leaves only
2.5 mm long, lanceolate,endingin a canaliculate
leaf tip. Marginsentire.Costafilling2/3of the leaf
base, excurrent,in transversesection with ventral hyalocysts,slightlyridgedat back. Alar cells
lacking.Basal laminal cells hyaline, thin-walled,
rectangular,5-10 x 45-60 ,m, towardsthe margins in 5-7 rows narrower.Upper laminal cells
short rectangular,ca. 2:1, 10-16 x 16-32 ,m,
incrassate.
Distribution (Fig. 132). On rocks in alpine
regionsof the Andes at high elevations between
3500 and 4900 m in Venezuela, Colombia, Ecuador, Peru and Bolivia.
Specimensexamined.COLOMBIA.CALDAS: Neva-
do de Ruiz,Cleef2440a(U).
VENEZUELA. MERIDA:
Paramode Piedras Blan-
cas, Griffinet al. 1423 (FLAS);Sierrade SantoDoetal. 1182(FLAS).

mingo,ParamodeMucubaji,
Griffin
ECUADOR. IMBABURA:
Lago de San Marcos Ca-
63 (hb.Frahm).NAPO:
yambe,Cazalet&Pennington
78?21'W,
0?47'S,Lojtnant&MoLagunaYuragcocha,
lau11584(AAU);Paramo
deGuamani,
PifoCarretera
Jaramillo4118(NY).
Papallacta,
PERU. ANCASH:
ParqueNacional Huascaran,Laguna Llaganuco,Frahm 823900 (hb. Frahm).JUNIN:
La RayabeAcopalea,Kunkel1131(MICH).PUNO:
tweenSantaRosaand Sicuani,Hegewald5490 (hb.
Frahm).
Cordillera de Tunari,
17?17'S,66?23'W,Lewis 79-2573 (F).
187
IE
FIG. 131. Campylopustrichophylloides(Kinzls.n., B). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Am.
61. Campylopus trivialis C. Miller ex Britton,

Bull. Torrey Bot. Club 23: 476. 1896. Type.
Bolivia. Mapiri, Rusby s.n. (holotype, NY).
Figs. 133, 134.
CampylopuscuatrecasiiRobinson, Bryologist70: 16.
1967. Type. Colombia. Cuatrecasas19090a (holotype, US; isotype, COLO).
CampylopustenerBrotherus,Rev. Bryol.47: 2. 1920.
Type. Ecuador.Azuay:Rio Gualaquiza,Allioni s.n.
Plants small, usuallyonly to 1.5 cm high, rarely 2 cm, in soft, light green or brownish green
tufts. Stems radiculose below, erect, the lower
stem leaves smaller,the upperstem leaves longer
pointed, pencil-like. Leaves 4-5 mm long, narrow lanceolate,from ovate base contractedinto
a long, fine point. Costa long-excurrent,finely
serrateat tips, filling 1/2of the leaf base, in transverse section with ventral hyalocysts, slightly
ridgedat back. Alar cells lacking. Basal laminal
Flora Neotropica
188
70
80
60
50
140
0
A" ~b
_---o
-...-'
----..
- ....II- -
I?
400
600
800
100km
30
ml.I
--- _. ..- ..
........
~~~oo
200
100 200 300 400 500 600
- - "_--------
I'
01
3b--to-------------3
F_I
FIG. 132.
20
Distribution of Campylopus trichophylloides (*) and C. uleanus (0).
cells short rectangular,hyaline, ca. 3-4:1, 6-16

x 13-38 um. Upper laminal cells rectangular,
ca. 3-4:1, 3-10 x 10-16 um. Lamina vanishing
below midleaf.
Sporophytefoundonly once (Laegaard52906).
Seta to 1 cm long, yellowish. Capsule 1.5 mm
long, erect, symmetric, olive-brown, furrowed.
Operculumobliquelyrostrate,0.5 mm long, reddish. Calyptrafimbriateat base.
Distribution(Fig. 134). On soil and rocks in
high alpine regions of the Andes in elevations
between 3000 and 4900 m in Bolivia, rarely on

open banks at lower altitudes in Costa Rica, Venezuela, Colombia, Ecuador, Peru and Bolivia.
Specimens examined. COSTA RICA. Cerro de la
Muerte,Schultes 12050 (FLAS).
COLOMBIA. CUNDINAMARCA: Sabana de Bogoti,
Schultes 11058 (FLAS);Mun. de Supata,CuchillaEl

Tablazo, Linares& Bulla 462 (COLO).
VENEZUELA.MERIDA:Sierrade Santo Domingo,
Paramode Mucubaji,Griffinet al. 934 (FLAS);above
Pico El Aquila, Griffinet al. 1330 (FLAS);road to La
Azulita and Jaji, Fransen 1218 (GB).
189
'
tmm
FIG. 133. Campylopustrivialis(Griffinet al. 1330, FLAS). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
Mm.
50 /am.,
ECUADOR. AzuAY:Cuenca-Ofiaroad, Laegaard is 79-2130(F);headof Rio Zongo,1617'S, 6806'W,
52906 (NY); Zamora,4S, 79?W,Ortega531 (MO).
Lewis79-1792(F).
PERU. ANCASH:
PuyaraimondiiNat. ParkE of ChiAlongroad
quian,Philippis.n. (hb. Frahm).AYACUCHO:
Huanta-SanFrancisco,Frahm 823975 (hb. Frahm). 62. Campylopusuleanus(C. Miiller)Brotherus,
Cordillera de Tunari,
17?17'S,66?23'W,Lewis 79-2571 (F);base of Serranias

Tarucani, 17?14'S,66?24'W,Lewis 79-2474A (F); LA
PAZ:1 km below Escoma-Charasaniroad NNW of
Chuma, 15?16'S,6902'W,Lewis 79-816 (F);Aguastermalesde Rio JunthumaKhuchu,18?6'S,6902'W,Lew-
Nat. Pflanzenfam.1(3): 333. 1901. Dicranum

uleanum C. Miiller, Hedwigia 39: 258. 1900.
Type. Brazil.S. Catarina:Flaggenberg,Ule 148
(holotype, destroyed at B; lectotypus nov.,
H-BR).
Figs. 2C, 132, 135.
Flora Neotropica
190
80
.-
'-
ct<
/*
60
70
<
!
~~~J~--~
.....
/,
2i,,.'o
/'
40
50
10
.0
20
100
60
40
200
300
400
800 1000k
500
600
"
FIG. 134.
20
m
_mile
Distribution of Campylopus trivialis (0) and C. viridatus (0).
Plants in loose tufts, brownishgreen with yellowish stem tips. Stems to 5 cm high, loosely
foliate.Leaveserectpatent,5-7 mm long, narrow
lanceolate, entire, with only a few teeth in the
apex. Costa filling 1/ of the leaf base, excurrent,
in transversesection with ventralhyalocystsand
dorsal groups of stereids, ridged at back. Alar
cells reddish brown, thick-walled, large. Basal
laminal cells longly rectangular,incrassateand
pitted, 13-29 x 6-16 .m, narrower at margins.

Upper laminal cells rhombic, incrassate, pitted,
6-10 x 10-19 tm.
Distribution (Fig. 132). On wet rocks, found
only a few times in SE Brazil.
Serrados Orgaos,MorroAssu, v. Liitzelburg7040 p.p.,
6928b (JE).
191
FIG. 135. Campylopusuleanus (Ule 148, H-BR). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of
leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 ,um.
63. Campylopusviridatus(C. Muller)Brotherus

in Engler & Prantl, Nat. Pflanzenfam. 1(3):
333. 1901. Dicranum viridatum C. Miiller,
Hedwigia 39: 257. 1900. Type. Brazil. Sao

Francisco,Rio Passucao,Ule 31 (holotype,destroyed at B; lectotypus nov., H-BR).
Figs. 134, 136.
25: 54. 1954,hom. illeg.non Dixon, 1922.Type.

Brazil.EstaoaoBiologicaAltodaSerra,Hoehne528
(holotype,JE;isotypes,B, S).
Campylopus pseudodichrostisBrotherus, Denkschr.
Akad.Wiss.Wienmath.-nat.
KI.83:263.1926.Type.
Brazil.SaoPaulo,Schiffner
402 (lectotype,H-BR).
Plants in loose tufts, slender, yellowish green

above, blackishbelow. Stems 5-10(-15) cm high,
not tomentose, loosely appressedfoliate. Leaves
CampylopusangusticostaJ.-P. Frahm, Bryoph. Bibl.
5: 17. 1975, nom. nov. pro Campylopusangusti- 6-8 mm long, lanceolate,canaliculate,the marnervisHerzog,Memoranda
Soc.FaunaFl. Fennica gins serratein the upper third. Costa relatively
Flora Neotropica
192
:c
a 0u
FI.
3.
apyousvriaus(Vtl
leaf.; E.Bsllmnlcls
.Uprlmnlcls
63,S).A
Pat. B. L af.C eftp

naee
cl
a o
irsoi
.Trnvrescino
etis=5Mm
FIG. 136. Campylopusviridatus(Vital 7663, SP). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails = 50 um.
narrow,fillingonly 1/4-1/ of the leaf base, ending

in the leaf tip or shortly excurrent,in transverse
section with ventral and dorsal stereids, ridged
at back. Alar cells numerous,inflated,brownish,
thickened. Basal laminal cells rectangular,incrassate, pitted, 13-67 x 6-10 um, at margins
in several rows narrower.Upper laminal cells
rectangularto oblique,incrassate,10-38 x 3-10
Am.
Distribution (Fig. 134). Known only from a
few localities on wet rocks and swampy places

in SE Brazil.
Specimens examined. BRAZIL. BAHIA: Mun. de
Leucois, 5 km N of L., Boom & Mori 1187 (NY). SAo
PAULO:Mun. Sao Bernardodo Campo, along road
Caminho do Mar, Vital5385 (SP).
64. Campylopus widgrenii (C. Miiller) Mitten, J.
Linn. Soc. Bot. 12: 88. 1869. Figs. 137, 138.
DicranumwidgreniiC. Miiller,Bot.Zeit. 14:418. 1856.
Type. Brazil. Minas Gerais: Caldas, Widgrens.n.
193
'
/i
ki
4mm
^jY~f~c
.00..
FIG.137. Ca
'00
D.
Transvee
FIG. 137. Caampylopus widgreni (Widgren

H-BR). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section
srn.,
of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details = 50 Mm.
(holotype,destroyedat B; lectotypusnov., NY; isotypes, H-BR, H-SOL).

Plants 3-5 cm high, dirty green to brownish,
in dense tufts. Stems equally foliate with loosely
patent leaves. Leaves 4-5 mm long, lanceolate,
margins entire, slightly canaliculate at tips. Costa
filling 1/3 of leaf base, excurrent, the excurrent part
slightly serrate, in transverse section with ventral
hyalocysts and dorsal groups of stereids. Alar
cells large, inflated, hyaline or reddish. Basal
laminal cells rectangular, incrassate, slightly pitted, 10-35 x 6-10 Am, at margins shorter and
narrower. Upper laminal cells oval, 1:4, 3-6 x
10-26 ,m.
Distribution (Fig. 138). On rocks in montane
forests in SE Brazil.
Falls of Rio
Specimensexamined.BRAZIL.BAHIA:
Ferro Doido, 18 km E of Morro do Chapeu,Irwin et
Km 392 along BR
al. 30748 (NY). MINASGERAIS:
Flora Neotropica
194
1o
90
/~~~~~~~~30
./I~/SC~=
00 O mile
^~~~~~~~~~~
\,0 r
X'
I20o
.2
100 200 300 40 0
'~.
50
60
70
80o
- --
1/
Prepared by Hendrik R. Rypkena
FIG. 138.
Distribution of Campylopus 2wdgreni (0) and C zygodonticarpus ().
135, Mun. de Ouro Preto, Frahm s.n. (hb. Frahm).

SANTA CATARINA:Nova Friburgo, Sehnem 7133b
(LBLC).
65. Campylopus zygodonticarpus (C. Miiller)
Paris, Ind. bryol. 265. 1894. Figs. 138, 139.
Dicranumzygodonticarpum
C. Muller,Linnaea42:471.
1879. Type.Venezuela.Tovar,Fendler35 (holotype,
destroyedat B; lectotypusnov., NY).
Campylopodiumtuerckheimii(C. Miiller)Brotherusin
DicranellatuerckheimiiC. Miiller,Bull. Herb. Boissier 5: 186. 1897. Type. Guatemala.Alta Vera Paz:
Coban, Tuerckheim6652 (holotype,destroyedat B;
lectotypusnov., NY; isotype, BM).
Plants to 2.5 cm high, yellowishgreen,in loose

tufts.Stemsequallyfoliatewith appressedor erect
patent leaves, the fertile plants comose at tips,
giving a pencil-like appearance.Leaves 5 mm
long, narrowlanceolate,graduallycontractedto
a long acumen, serrateat tips. Costa filling /3of
the leaf base,excurrent,in transversesectionwith
ventral hyalocystsand dorsal groupsof stereids,
smooth at back.Alarcells reddish,inflated.Basal
laminal cells shortlyrectangular,ca. 1:1.5-3, incrassate, 6-16 x 13-35 ,m, smaller and
subquadrate at margins. Upper laminal cells
quadrateto short rectangular,6-13 x 8-20 ,m,
ca. 2:1.
iA
195
FIG. 139. Campylopuszygodonticarpus(Fendler 35, NY). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse

50 Mm.
Seta 5-7 mm long. Capsuleupright,symmetric, 1.5 mm long, yellowish to dark brown, furrowed when emptied. Calyptraciliate at base.
Distribution(Fig. 138). On open bare soil and
rotten stumps, on palm trunksand log, in Mexico, Honduras, Guatemala, Costa Rica, El Salvador, Venezuela,Colombia, Ecuador(only GalapagosIslands),Peru and Bolivia, in elevations
between 1200 and 2500 m, in Peru and Bolivia

up to 3200 m.
3 km
Specimens examined. MEXICO. VERACRUZ:
NE Coatepec, Vivenos76c (TENN).
GUATEMALA.ALTAVERAPAZ:
Above SantaCruz,
Standley71051 (F);vic. of Lobin, Standley92068 (F);
along road to Tamalin, Standley 90784 (F).
vic. Fuentes Georginas, Standley
QUEZALTENANGO:
86013 (F).
196
Flora Neotropica
El Achote near SignaHONDURAS. COMAYAGUA:
There is also no isotype in the herbariumof

Brotherus(H-BR) and, surprisingly,Brotherus
EL SALVADOR. Los Planes del Monte Christo,
did not include this speciesin the second edition
Watson 77-05 (MO).
COSTA RICA. La Fuente, Falda del Turrialba,Al- of the NatiirlichePflanzenfamilien,even though
he had newly combined this species in the genus
faro I (F); Tierra Morenas, Alfaro 24 (F); Las Concavas,
Lankaster s.n. (F); Rio Jesus y el Picacho del MonCampylopusin the first edition.
tepec, Standley 56117 (F).
dango de San Ram6n, Branes 21861 (F); Juan Vinas,
Roll 1260 (JE).
San Cayetano,HaCOLOMBIA.CUNDINAMARCA:
cienda Portugal, Cleef 6045 (U).

VENEZUELA. MERIDA: Carretera Merida-La
Azulita, Lopez & Keogh 13741 (FLAS).
PERU. Cuzco: Urubamba, Aguas Calientes, Hegewald 8784 (hb. Frahm).LA LIBERTAD:
LagunaSausa-
CampylopuspercurvatusC. Muller,Flora83:331.
1897. Type. Venezuela.Tovar, 1800 m, Fendler s.n.
Campylopus subfalcatus (Horschuch) Jaeger,

1870-1871: 442. 1872. Dicranumsubfalcatum
cocha near Huamachuco,Hegewald6015 (hb. HegeRoad Chachapoyas-Moyobamba
Horsch., Fl. bras. 13. 1844. Type. Brazil."In
wald).SANMARTIN:
km 387, Frahm et al. 337 (B); km 403, Frahm et al.
montibussilvestribuspropeSincoraprov. Ba308 (B).
Martiuss.n.
hiensis,"
s.n.
near
BOLIVIA.Near Pata, Williams
Williams1758 (F).
(F);
Apolo,
A species which is usually found with sporophytes. It is closely related to C. pauper, as seen
by the pencil-like appearance of the comose stems,
but differs by the symmetric capsules. Microscopically it is easily recognized by the short,
incrassate, basal laminal cells.
Campylopus zygodonticarpus has been included in the species concept of C. flexuosus by Bartram. Herbarium revisions have revealed, however, that this is a widespread species and
probably more frequent than realized hitherto.
DoubtfulSpecies
The following species are known only from the
type localities. However, type material is not
available. Most of the species were described by
Carl Miller, whose herbarium at the Botanical
Museum in Berlin was destroyed, and in contrast
to many other species described by him, isotypes
could not be located in other herbaria.
Campylopus cruegeri (C. Miller) Paris, Ind. bryol.
Suppl. 91. 1900. Dicranum cruegeri C. Miller,
Hedwigia 37: 226. 1898. Type. Trinidad. St.
Anne, Criiger s.n.
Campylopus exfimbriatus C. Muller, Flora 83:
331. 1897. Type. Venezuela. Sierra Nevada de
Merida, Goebel s.n.
Campylopus itacolumitis (C. Miller) Brotherus
in Engler & Prantl, Nat. Pflanzenfam. 1(3):
334. 1901. Dicranum itacolumitis C. Miller,
Gen. musc. frond. 263. 1900. Type. Brazil.
Serra de Ouro Preto, Ule s.n.
Campylopusterebrifolius(C. Muller)Jaeger,Ber.
Thatigk.St. GallischenNaturwiss.Ges. 18771878: 385. 1880. Dicranum terebrifoliumC.
Miiller, Linnaea 38: 593. 1874. Type. Ecuador. "Andes Quitenses,monte Pinchincha,in
summis declivibus,"Karstens.n.
According to the description, with spirally
twisted leaf tips; perhaps identical with CampylopusspirifoliusHerzog (cf. Robinson, 1967),
which is synonymouswith Ditrichumbogotense
Hampe.
5. DicranodontiumBruch,Schimper& Gimbel,
Bryol. Eur. 1: 159. 1847.2
Type species D. longirostre(Weber & Mohr)
Bruch,Schimper& Gimbel.
Plantsin soft tufts.Leavesnarrowlylanceolate,
chanelled,endingin a long, slender,often serrate
subula. Costa filling about 1/3 of the leaf base,
excurrent,fillingthe subula,in transversesection
with a median band of deuter cells and multicellularlayersof ventraland dorsalstereids.Alar
cells inflated, hyaline or reddish. Basal laminal
cells rectangular,narrower at margins. Upper
laminal cells elongaterectangular,narrow,often
porose.
Seta cygneous in immature sporophytes,
straightand twistedin maturesporophytes.Capsule ovoid to shortcylindrical,symmetric.Operculumlonglyrostrate,as longas the capsule.Peristome teetn 16, split, vertically striate below,
papillose above. Annuluslacking.Spores 13-15
2
In collaboration
withMonikaGiese-Stral3er.
197
um in diam., finely papillose. Calyptraentire or
fringedat base.
This genusis most closely relatedto Atractylocarpus,fromwhichit differsby longerupperlam-
inal cells, more incrassatebasallaminalcells and

especiallya cygneous seta with shortercapsule.
Dicranodontiumis confined to montane forests
whereasAtractylocarpusis alpine.
Key to the Neotropical Species of Dicranodontium

1 Inner basal laminal cells incrassate,except for size not much differentiatedfrom the outer basal and
upperlaminal cells.
4. D. pulchro-alare.
2 Laminalcells stronglypitted. ...........................................
2. D. denudatum.
.............................................
2 Laminalcells smooth. ......
1 Innerbasal laminal cells hyaline,pellucid,distinct from outer basal and upperlaminal cells.
3. D. meridionale.
3 Leaves not falcate;leaf tips stronglyserrate .....................................
D. brasiliense.
3 Leaves falcate;leaf tips indistinctlyserrate. .......................................1.
1. DicranodontiumbrasilienseHerzog,Hedwig- Campylopuschionophilus(C. Muller)Mitten,J. Linn.

Soc. Bot. 12: 81. 1869. DicranumchionophilumC.
ia 67: 254. 1927. Type. Brazil. Serrados OrMiller,Syn.musc.frond.1: 398. 1848.Type.Vengaos, v. Liitzelburg6519 (holotype,JE).
ezuela.Merida,Funck& Schlims.n. (holotype,deFigs. 140, 141.
stroyedat B;lectotypusnov.,NY).
andfurthersynonymsfromthe Holarctic.
foliate
Plants to 4 cm high, robust, densely
Plantsyellowishto brownishgreen,slender,in
with stronglyfalcate,homomallousleaves.Leaves
4-5 mm long, from ovate base contractedinto a dense mats, to 3(-5) cm high. Stems often partly
long, slendertip. Costa filling /3 of leaf base, in defoliate, sometimes with homomallous leaves
transversesection with bands of median deuter at tips. Leaves erect patent, 4-8 mm long, cancells and dorsal and ventral stereids, excurrent aliculate.Costafilling1/3of the leaf base, in transin a long, only slightly serratesubula.Alar cells verse sectionwith ventraland dorsalstereidsand
inconspicuous.Inner basal laminal cells rectan- a medianbandofdeutercells, excurrentin a long,
gular,thin-walled,pellucid, 15-18 x 30-45 Am. narrow,faintly serratesubula.Alar cells hyaline
Outerbasal laminal cells in 10 rows narrowand or reddish, often auriculate.Basal laminal cells
elongate, 2-4 x 25-35 Mm,chlorophyllose.Up- short rectangular,8-15 x 25-40 gm, narrower
per laminal cells rectangular,incrassateand pit- at margins.Upperlaminalcells elongate,slightly
lm.
ted, 10-13 x 40-70 Mm.Lamina vanishingbe- pitted, 5-8 x 35-100
low midleaf.
Seta 10 mm. Capsule1.2 mm long,cylindrical,
symmetric.Calyptraentire at base.
Distribution(Fig. 141). In damp montaneforests in elevations between 2000 and 3000 m on
type locality in SE Brazil.
The only speciesof Dicranodontiumoccurring soil, rocks and rotten wood, from the boreal
in the southernhemisphereand, as in Paraleu- regions of North America through the Rocky
cobryum longifolium ssp. brasiliense, the only Mountains to Mexico, Guatemala, Honduras,
representative of a primarily boreal northern Nicaragua,Costa Rica, Venezuela and Bolivia,
hemisphericgenus in SE Brazil.It resemblesD. also in Europeand Asia.
meridionalewith respect to the pellucid inner
48 km NE
examined.MEXICO.OAXACA:
basal laminal cells and differsonly in the falcate deSpecimens
Llanode las Flores,Delgadillo830 (H). VERACRUZ:
leaves and the less serrateleaf tips. Thus, it may Orizaba,97?18'W,18?43'W,
AndrewB 543 (F).
GUATEMALA. Argueta, Bernoulli & Cario s.n.
be also regardedas a disjunct race of D. meridionale in SE Braziland recognizedas a subspe- (GOET). East of Totonicapan, Sharp 2600 (FLAS).
Above San Mateo,Sharp4951 (F).
HUEHUETENANGO:
cies of the latter.
de las Minas,SteyerSEof Concepcion
Chiquimula,
mark 31000 (F).
HONDURAS.MORAZAN:
Slopesof CerroZynca,
2. Dicranodontiumdenudatum(Bridel) Britton
in Williams, N. Amer. Fl. 15: 151. 1913. Di- Standley& Williams760 (F).
NICARAGUA. GRANADA:Upper slopes of Volcan
cranum denudatum Bridel, Muscol. recent. Mambacho
Croat
alongW shoreof LakeNicaragua,
39150 (MO).
Suppl. 1: 184. 1806. Type not seen.
COSTA RICA. HEREDIA:Vara Blanca,Bishops.n.
Figs. 141, 142.
Flora Neotropica
198
"
FIG. 140. Dicranodontiumbrasiliense(v. Liitzelburg6519, JE).A. Plant. B. Leaf.C. Leaf-tip.D. Transverse

50 Mm.
(FLAS);vic. LagunaBarba, 10?5'N,83?55'W,Crosby
9854 (MO).
VENEZUELA.BOLVAR:Mt. Roraima,Steyermark
58924b (F). MIRANDA:
Pico de Naiguara,Steyermark
62968 (F).
BOLIVIA.LAPAZ:Nevado JankhoUma, 15?51'S,
68?34'W,Lewis 79-1430A (F).
3. Dicranodontium meridionale Bartram, Contr.
U.S. Natl. Herb. 26: 65. 1928. Type. Costa
Rica. Heredia: Cerro de las Caricias, Standley
52296 (holotype, FH; isotype, NY).
Figs. 143, 144.
Plants slender,to 5 cm high, brownishgreen,

in loose tufts. Leaves 4-7 mm long, narrowly
lanceolate,canaliculate,endingin a long, slender,
serrate tip. Vegetative propagation by whole
leaves, whicheasily fall off. Costa /3of leaf width,
excurrent,in transversesection with ventraland
dorsal bands of stereids and a median band of
deuter cells. Alar cells hyaline or reddish. Basal
laminalcells shortrectangular,hyaline, 16-20 x
35-40 lm, narrowerat margins,forminga border of 7-8 rows. Upper laminal cells elongate,
2-4 x 25-50 Am.
199
10C
80
90
60
7C
40
5C
73~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~3
h,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~2
O~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'
- - - -I -----
20~~~~~~~
i o
'"~
20 40
Go
80010
...!
.........
'i",
k,1
1 2, 4 ..00'"-..
......
0...
. ~
--~
. . . . ,~~-"-,------~
IL
_"0
Prepared by Hendrik R. Rypkerna
FIG. 141.
Distribution of Dicranodontium brasiliense (0) and D. denudatum (0).
Seta 9 mm long. Capsule oval. Peristome teeth

split to the base. Spores 13-15 Am in diam.
Distribution (Fig. 144). Epiphytic in montane
rainforests in Guatemala, Costa Rica and Venezuela.
Specimens examined. GUATEMALA. HUEHUETENANGO:Above San Mateo, Sharp 4954 (DUKE).
Orilla de Riachuela, GrifCOSTA RICA. ALAJUELA:
fin 181 (FLAS);ParqueNacionalVolcande Pods,Griffin & Araya 53 (FLAS). HEREDIA:Cerros de Zurgui,
Standley 50492 (NY); Cerro de las Lajas N of San

Isidro, Standley& Valerio51647 (NY); Barba,Alfaro
1219 (F);Rio Patria12 km above San Rafael,Richards
R5847 (MICH).
VENEZUELA. AMAZONAS:Cerro Duida, 3022'N,
65?36'W,Buck & Brewer15535 (NY).
This species resembles much Dicranodontium

denudatum in appearance but is distinct by its
inner basal laminal cells, which are enlarged, hy-
aline and conspicuously pellucid, a character

found also in the SE Asian D. fleischerianumW.
Schultze-Moteland the EurasianD. uncinatum.
4. Dicranodontium pulchro-alare Brotherus,
Trans. Linn. Soc. London Bot. ser. 2, 6: 89.
1901. Type. Venezuela. Mt. Roraima, McConnell & Quelch 345 (lectotypus nov.,
Figs. 144, 145.
H-BR).
Plants very robust and tall, to 12 cm high,
yellowish to brownishgreen.Stems with reddish
tomentum. Leaves 8-12 mm long, canaliculate,
narrowly lanceolate, ending in a long, serrate
subula. Costa filling 13 of the leaf base, in transverse section with multilayeredbands of ventral
and dorsal stereids, excurrent.Alar cells large,
reddish, inflated and prominent. Basal laminal
cells rectangular,20-25 x 50-100 ,tm, incrassate
Flora Neotropica
200
and pitted, narrowerat margins.Upper laminal
cells incrassate,stronglypitted, narrowlylanceolate to elongate, 2-5 x 13-25 rm.
Distribution(Fig. 144). In moist montane forests, known only from the type locality at Mt.
Roraima and one additional record from Costa
Rica.
Specimenexamined.COSTA RICA. HEREDIA:Vic.
LagunaBarba,10?5'N,83?55'W,Crosby9854(MO).
In the stronglypitted basal laminal cells and
the inflatedreddishalarcells this speciesstrongly
resembles Dicranodontium subporodictyon
Brotherus,a holarcticspecies, occurringin China, westernNorth Americaand westernEurope.
The relationshipbetweenthesespeciesis not clear
and needs furtherstudy.
ExcludedSpecies
Dicranodontiumsetosum Williams, Bull. Torrey Bot. Cl. 34: 570. 1908. Type. Colombia.
Paramode BuenaVista, Pittier2060 (holotype,
NY; isotype, PC) = Atractylocarpuslongisetus
(Hooker)Bartram.
Dicranodontiumschwabei Herzog & Theriot,
Beih. Bot. Centralbl. 60B: 21. 1939. Type.
Chile. Aysen: Istmo de Ofqui, Grosses.n. (holotype, JE) = Chorisodontiumsp.
6. Microcampylopus(C. Miller) Fleischer,Musci Fl. Buitenzorg1: 59. 1904.
C. Muller,Hedsubg.Microcampylopus
Campylopus
wigia38:77. 1899.
With the characteristicsof M. curvisetus.
Type species:M. subnanus(C. Miller) Fleischer (=Microcampylopuskhasianus (Griff.)Giese
& J.-P. Frahm).
A genus, in which 27 species have been included world-wide.A recent monograph(Giese
& Frahm, 1985b)revealedthat most specieshad
to be placed in Campylopus,Dicranella or had
been confused with species of Campylopodium.
In conclusion, three species remainedin this genus, of which one (M. curvisetus)is neotropic,
another (M. khasianus)is SE Asian in distribution, and the third (M. laevigatus)occurs in SE
Asia and tropicalAfrica.
1. Microcampylopus curvisetus (Hampe) Giese

& J.-P. Frahm, Lindbergia 11: 116. 1985.
Figs. 1F, 4C, 6B, 8D, 146, 147.
Campylopodiumcurvisetum(Hampe)Paris,Ind. bryol.
237.1894. AongstroemiacurvisetaHampe,Ann. Sci.
Nat. Bot. ser. 5, 3: 355. 1865. Dicranellacurviseta
(Hampe) Mitten, J. Linn. Soc. Bot. 12: 38. 1869.
Type. Colombia. Bogota, Tequendama,Lindig s.n.
(holotype,BM; isotype, S).
Campylopodium
fendleri(C. Miiller)Paris,Ind. bryol.
237. 1894. Dicranellafendleri Jaegerex Paris, Ind.
bryol. 295. 1894, nom. invalid. Aongstroemiafendleri C. Miiller,Linnaea42: 470. 1879. Type. Venezuela. T6var,Fendler32 (holotype,B; isotypes,BM,
H-BR, NY, S).
itatiaiense(C.Miiller)Paris,Ind.bryol.
Campylopodium
Suppl.88. 1900.AongstroemiaitatiaienseC. Muller,
Bull. Herb. Boissier6: 83. 1898. Type. Brazil.Serra
do Itatiaia, Ule 102 (holotype, destroyedat B, lectotypusnov., NY; isolectotypes,BM, H-BR, JE, PC,
S).
(C.M'iller)Paris,Ind.bryol.
Campylopodiumpilopogon
238. 1894. Dicranellapilopogon(C. Miiller)Jaeger,
Ber. Thatigk.St. GallischenNaturwiss.Ges. 18771878:373. 1880.AongstroemiapilopogonC. Muller,
Linnaea38: 630. 1874. Type.Mexico.Mirador,Santoriuss.n. (holotype,destroyedat B; lectotypusnov.,
NY; isolectotypes,BM, S).
Campylopodiumpusillum (Schimper) Williams, N.
Amer. Fl. 15: 94. 1913. CampylopuspusillusSchimperin Bescherelle,Mim. Sc. Nat. Cherbourg16:165.
1872. Type. Mexico. Orizaba,Miillers.n. (holotype,
not located at BM; isotypes, NY, PC).
CampylopusexustusMitten, J. Linn. Soc. Bot. 12: 83.
1869. Type. Periu.Pingullu, Spruce 57 (holotype,
NY).
Campylopusleucogaster(C. Miller) Mitten, J. Linn.
Soc. Bot. 12: 75. 1869. Type.Mexico.Xalapa,Deppe
&Schiede1072 (holotype,destroyedin B; lectotypus
nov., H-BR; isotype S).
Campylopodiumsartorii(C. Miller) Paris, Ind. bryol.
238.1894. Dicranellasartorii(C.Miiller)Jaeger,Ber.
373. 1880.AongstroemiasartoriiC. Miiller,Linnaea
38:629.1874. Type.Mexico.Orizaba:Mirador,Sartoriuss.n. (holotype,destroyedat B;lectotypusnov.,
BM; isotype, S).
Plants 5-10 mm tall, yellowish green to dark
green, in loose tufts. Stems short, reddish, comose foliate at tips. Leaves 2.5-4 mm long, the
lower ones from an ovate base gradually narrowed, lanceolate, the upper ones with broad base,
contracted into a long subula. Costa filling 1/3 of
the leaf base, excurrent, filling the subula, in
transverse section with a median band of deuter
cells and several rows of stereids dorsally and
ventrally. Basal laminal cells short rectangular,
40-60 x 8-10 ,um. Upper laminal cells shorter,
201
,J r
0.~~~~~~~~~~~~~~~~;iu~i;
031~
0
a0~~~~~l~~
B 0~~~~.
0
00
0 D ~~~~~,
ieiIrni;I
sectin
ofleaf
E. asallamial
clls.
3cm
. Uper
lmina
/o
cels.
Ulabeed
scle
br
fo
micoscoic
dtail
aU0
FIG. 142. Dicranodontiumdenudatum(Millspaugh11, U). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse

50 sm.
20-26 x 4 tm, rectangularto rhomboid. Alar

cells lacking.
Seta 3-5(-7) mm long, curved or flexuose in
age, yellowish green to brownish. Capsuleabout
1 mm long, ovate to short cylindric. Annulus
present. Stomata lacking. Exothecial cells incrassate, rhomboideal. Peristome teeth 16, divided to 2/3 or totally into two prongs, reddish
and striate at base, yellowish to hyaline and pa-
pillose in the upperpart.Operculum/2-2/3as long

as the capsule,rostrate.Spores18-24 ,m in diam.,
coarsely papillose with 6-8 papillae per diam.
Distribution(Fig. 147). On humic soil in open
habitats in forests, preferablyin disturbedareas
as roadside banks, in Mexico, Guatemala,Panama, Dominican Republic,Jamaica,Venezuela,
Colombia,Peruand Brazil.Recordsfrom Puerto
Rico belong to Campylopodiummedium.
202
Flora Neotropica
\^ô
r^00-'0
?0
<9<:>^0
FIG. 143. Dicranodontium meridionale (Richards R5847, MICH). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details
= 50 Am.
Specimens examined. MEXICO. CHIAPAS:Finca

Liquidambar, Sharp 4480 (NY); San Crist6bal, Muench
7391 (BM, H-BR, NY). Orizaba, Liebmann s.n. (NY).
Cordova, Mohr s.n. (NY). HIDALGO:Between Varios
and Chapulhuacan, Sharp 1584 (MO, NY).
GUATEMALA. El Quiche below Nebaj, Bartram
2356b (MO).
PANAMA. NW del Campamento, 8?15'N, 82?15'W,
Salazar 538 (NY).
JAMAICA. Cinchona, Nichols 164b (NY). Flamstead, Port Royal Mtns., Maxon 8720 (NY). Blue
Mountain Peak, Hegewald 8119 (BM).
DOMINICAN REPUBLIC. INDEPENDENCIA: 5 km
S of El Aquacate on road to Pedernales, 18?18'N,
7 142'W, Buck 4712 (NY). LA VEGA:22 km S of Constanza, 18?49'N,70?37'W,Reese 15894 (NY).

COLOMBIA. CAUCA:Near Popayan,Alston 8033
(BM, S).
VENEZUELA. MERIDA:
Along Merida-LaAzulita
road, Griffin2004 (B, MO, NY).
BRAZIL. AMAZONAS:
226 km N of Boa Vista just
S of Venezuelanborder,Frahm 1936 (NY); Taruma
Grandealong Rio Taruma 15 km E of Manaus,Keel
et al. 1792 (NY). MINASGERMS:Passo Quatro,Zikan
337 (JE); Serra de Caparao Nat. Park, Schdfer-VerVic. Itatiaya,
wimp 8901 (hb. Frahm). Rio DEJANEIRO:
Rose & Russell20545a (NY). SAO PAULO:Camposde

Jordao,Schafer-Verwimp8518 (hb. Frahm).
90
203
^^r^
^f~'~~~~~~-
...
\S
'G
X
14.Dsjbto
30
'o
FG
200
.. 600 800 o100 k
400
100 200 300 400 500 600.....
0 ......
by Hendrik R.Rypkema
/?~~0
jh,~~~~~
so0
60
70
80
fd.caootu
eiinl
.plhoae()
FIG. 144. Distributionof Dicranodontiummeridionale(0) and D. pulchro-alare(0).
As seen from the list of synonyms,this species

has usually been placed in Campylopodiumand
the genus Microcampylopushas never been reported from the Neotropics but was confined,
accordingto the literature,to Africaand SEAsia.
A re-evaluation of both genera revealed, however, that Microcampylopusis characterizedby
the absence of stomata in the capsules and
coarselypapillose spores. Therefore,the records
from the neotropicshad to be transferredto Microcampylopus.Both genera are closely related
and may also be regardedas subgeneraof one
genus. Gametophytically,they much resemble
Dicranella subg. Anisotheciumin the leaf morphology, with the broad sheathing base and
transversesection of the costa. Therefore,sterile
plants cannot be distinguished.Dicranellais differentiatedby erect setae and largerspores.
7. PilopogonBridel, Bryol. univ. 1: 519. 1826.
Plants to 5 cm, rarely to 8 cm high, in loose
tufts,brownish,blackish,greento yellowishgreen.
Stems mostly conspicuouslyequallyfoliatewhen
dry,giving the plantsa filiformappearance,erect
spreadingwhen moist, sparselyradiculoseat base.
Leavesstraight,3-7 mm long, from a contracted
base, lanceolate, longly subulate. Perichaetial

leaves with broad sheathingbase, suddenlycontracted to long fine subula, enclosing the seta
from /4 to nearly to the capsule. Costa at base
-1/2 of the leaf width, excurrentin a short awn,
smooth, denticulate at the extreme tip, or excurrentin a long serratehyaline awn, in transverse section with ventral and dorsal stereids,
smooth, ribbed or lamellose at back. Alar cells
not differentiated,rarely inflated and reddish.
Lowerlaminalcells thin-walled,hyaline,rectangular, narrowerat margins, extending upwards
along leaf margins,rarelyincrassate.Upper laminal cells thick-walled,oblique,rhomboidal,oval,
elongate-ovalor short rectangular.
Seta 1-2 cm long, erect. Capsule2-4 mm long,
light brown, cylindrical, sometimes broader at
base, straight,without stomata. Peristometeeth
16, on a basal membrane,undivided or split to
the base, densely papillose.Annuluspresent,dehiscent. Operculumlongly rostrate,erect. Calyptra cucullate,fringedat base with long cilia, rarely smooth. Spores ca. 13 gtm in diameter,
yellowish to brownish, smooth to minutely papillose.
Type specimen:P. gracilis(Hooker)Bridel(=P.
guadeloupensis).
Flora Neotropica
204
i
rx%~i
mm W
O
n10c'
C:):
,
1-.
CD
ii
FIG. 145. Dicranodontiumpulchro-alare(Quelch345, H-BR). A. Plant. B. Leaf.C. Leaf-tip.D. Transverse

50 ,m.
For a worldwide monograph see Frahm only two additional species occur: P. schilleri
(1983a). This genus has a center of radiationin Herzog in Chile and P. africanus Brotherusin
the northernAndes. Six of the 8 species known CentralAfrica.
worldwide occur there. Outside the neotropics,
Key to the Neotropical Species
5. P. peruvianus.
1 Abaxial surfaceof the leaf with lamellae 2-4 cells high ...............................
1 Abaxial surfaceof the costa smooth, ribbedor lamellose with lamellae 1(-2) cells high.
2 Basallaminalcells incrassate;alarcells differentiated,inflated;operculumand peristometeeth as long
3. P. longirostratus.
or longerthan the urn .......................................................
205
!8.
u
i :', fi A
,\1,
I,
'//i
^/
F/
'
:II" <
I
~Y'~1i
-~"\".
j.
1
~;:
'q/''
nL
-:lo'
'J'
teethshorter
thanthe urn.
peristome
^^y^,
iL
/
icj~
.5mm'
O';
Xr
I,
-/
4 Upper
laminal cells oval to elongate oval.
FIG. 146. Microcampylopus

curvisetus(Lindigs.n., BM). A. Plant. B. Leaf.C. Leaf-tip.D. Transversesection
of leaf. E. Basal laminal cells. F. Upper laminal cells. G(H). Capsule(s).Unlabeled scale bar for microscopic
details
details = 50
50 ~m.
Am.
2 Basal laminal cells thin-walled, hyaline; alar cells not conspicuouslydifferentiated;operculumand
peristometeeth shorterthan the urn.
3 Lower stem leaves contractedto a long fine subula, excurrentpart of the nerve longer than the
4. P. macrocarpus.
lamina;calyptrasmooth at base ............................................
3 Lowerstem leaves graduallynarrowed,nerve shortlyexcurrent;calyptraciliate at base.
P. tiquipayae.
4 Upper laminal cells subquadrate.............................................6.
4 Upper laminal cells oval to elongateoval.
2. P. laevis.
5 Leaves with hairpoints;peristometeeth split ...................................
5 Leaves without hairpoints(rarelyin perichaetialleaves);peristometeeth entire ...........
1. P. guadeloupensis.
....................................................................
ted. Alar cells developed, inflated, reddish.

The species can be arrangedin three subgenera:
species:P. longirostratus
ParapilopogonJ.-P. Frahm, Lindbergia9: 114.
1983.
Pilopogonella (Bartram) Theriot, Rev. Bryol.
Lichenol. 9: 13. 1936.
Basal laminal cells incrassate,sometimes pit-
206
Flora Neotropica
e09C
XC2070
60
50
0..
Prepard by Hendk
R.Rypkema
FIG. 147. Distributionof Microcampylopuscurvisetus.

Pilopogongracilis(Hooker)Bridel,Bryol.univ. 1: 519.
1826.
Didymodongracilis Hooker, Musci exot. 1: 5. 1818.
Type.Colombia."In monte Quindiuin regionetemperata Andes Cundinamarcae,"Humboldt24 (holotype, BM; isotype, B).
Pilopogon bernoullii (C. Muller) C. Muller, Genera
muse. frond. 256. 1900. Pilopogongracilisvar. bernoulliiC. Miller, Bull. Herb. Boissier5: 185. 1897.
Type. Guatemala.Toyahay,Bernoulli& Cario 1870
(holotype,destroyedat B; lectotypusnov., NY).
PilopogoncalycinusSchimperin C. Miller, Syn. muse.
frond.2: 256. 1851. Type. Mexico. Lagunade Tales,
Liebmanns.n. (holotype,destroyedat B; lectotypus
1. Pilopogonguadeloupensis(Bridel)J.-P. Frahm,
nov., NY).
comb. nov. Campylopusguadeloupensis(Bri- PilopogoncapitiflorusAngstroemex C. Muller, Gen.
muse. frond. 256. 1900. Type. Guadeloupe.Forsdel) Mitten, J. Linn. Soc. Bot. 12: 77. 1869.
stroems.n. (holotype,n.v.; isotype, BM).
DicranumguadeloupenseBridel, Spec. musc.
glabrisetusC. Miller, Bull. Herb. Boissier
1: 213. 1806. Type. Guadeloupe. Balby s.n. Pilopogon
2: 551. 1897. Type. Jamaica.Cinchona,Harriss.n.
Figs. 148, 149.
(holotype, B).
(holotype,destroyedat B; lectotypusnov., NY).
Peristome teeth split to the base. Costa lamellose at back.

species:P. laevis
P. peruvianus
Pilopogon
Peristometeeth entire. Costa not lamellose at
back.
species:P. guadeloupensis
P. macrocarpus
P. tiquipayae
207
i\
II
Ill/
5cm
BE
FIG. 148. Pilopogon guadeloupensis (Rehder 5, GOET). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse section
of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopic details = 50 gm.
Campylopusliliputanus(C. Miiller) Brotherus,Rev.

Bryol.47: 3. 1920. PilopogonliliputanusC. Miiller,
Nuov. Giorn. Bot. Ital. n. ser. 4: 36. 1897. Type:
Bolivia. Cochabamba:prope Choquecamata,Germain 1107 (holotype,destroyedat B;lectotypusnov.,
NY; isolectotypes,H-BR, JE, PC).
Pilopogonlongefimbriatus
Schimperin C. Muller,Nuov.
Giorn. Bot. Ital. n. ser. 4: 164. 1897, nom. nud.
Material.Andes Boliviae, Mandons.n. (BM).
Pilopogon microcarpusGeheeb & Hampe, Flora 64:
340. 1881. Type. Brazil.Apiahy, Puiggari200 (holotype, BM).
PilopogonsubjulaceusHampe, Vidensk. Meddel. Na-
ser. 3, 6: 136. 1875.

turhist.Foren.Kjoebenhavn
Type.Brazil.Itatiaia,Glaziou7065(holotype,BM).
Plants very variablein size, from 0.5 to 15 cm
high, generally 2-5 cm, yellowish to dirty or
brownishgreen. Stems very slender, equally foliate with appressedleaves, only the perichaetia
comose, tomentose in wet habitats. Leaves narrowly lanceolate, 3-5 mm long, with excurrent
costa, smooth at margins.Nerve occupying1/2 of
the leaf base, ribbed at back, excurrentin a ser-
208
Flora Neotropica
1(g
90
200 300 o so60
so
o0
20
X-
ml.
Prepared by Hendrik R Rypkema
FIG 149.
'
---
Distribution of Pilopogon guadeloupensis.
rate awn. Excurrentpart of the nerve short in

lower leaves to as long as the lamina in upper
leaves. Perichaetialleaves from broad sheathing
base abruptly narrowed to the long excurrent
nerve, the excurrentpartof the nerve sometimes
hyalineor subhyaline.Alar cells indistinctor absent. Lowerlaminalcells rectangular,hyalineand
thin-walled, 6-9 x 40-60 im. Upper laminal
cells elongate oval, thick-walled, 4-8 x 20-28
Jtm.
60
1
o0lo
70
80
Operculum rostrate, half as long as the urn. Calyptra ciliate at base.

Distribution (Fig. 149). On open bare gravelly
soil from the lowlands up to 4900 m, preferably
in altitudes between 1500 and 3500 m in Mexico,
Guatemala, Costa Rica, Dominican Republic,
Jamaica, Guadeloupe, Venezuela, Colombia, Ecuador, Peru, Bolivia and SE Brazil.
Specimens examined. MEXICO. CHIAPAS:Tapa-
chula, Benito Juirez, Frahm 792124 (hb. Frahm);5

Seta lengthvariable, 10-20 mm, nearlytotally km S of Tapilula on road to Tuxtla Gutierrez,Maye
surroundedby the perichaetialleaves. Urn 2-3 & Madison 337 (hb. Frahm);Crist6bal,Muench 7424
Cerro de Tutotepec near Acapulco,
mm long, 4-6 times longer than broad, cylin- (PC). HIDALGO:
4142 (FH, NY); Mineraldel Chico, Orcutt6753
Sharp
drical, erect or sometimes slightly asymmetric (FH); Honey Station, Barnes & Land 533 (JE, NY,
and curvedor slightlybroaderat base. Peristome PC); 3 miles S of Santa Ana, McGregor16362 (NY).
teeth undivided, filiform, densely papillose. OAXACA:
La Cumbre25 km above Oaxaca,Sharp2563
209
(FH, NY); San PabloAyutla70 km E of Oaxaca,Sharp (NY); 3 km W of Constanza,Jones & Norris 1267B
et al. 4580 (FH, NY); Llano de las Flores between (NY); La Lagunita,Norriset al. 5611 (NY). PERAVIA:
Tuxtepec and Oaxaca, Sharp 73 (FH); SierraJuarez La Nevera area 47 km S of Constanza,Reese 15708
above Valle Nacional,Sharp1885 (B, FH);SierraJua- (NY).
rez 29 miles N of Ixtlan, Hermann26201 (B, H, NY,
GUADELOUPE.Withoutlocalities,Duss 184 (NY);
PC, S);57 miles fromIuntepectowardOaxaca,Manuel Husnot 131 (BM, FH, H-BR, NY, PC); Parker 1107
590 (B, H); 26 km NE de Oaxaca hacia Ixtlan, Del- (BM). La Soufriere,Crosby4861 (MO).
Cerro del Padre Amaya
gadillo 680 (H); 50 km S of Tuxtepec, Conrad3216
(H);65 km N of Ixtlan,Mickel& HellwigB3700 (NY); 19 km W of Medellin, Luteyn 7094 (NY). ARAUCA:
22 mi N of Pochutlaon Hwy. 175, Norris& Taranto Sierra Nevada del Cocuy, Cleef 8864 (U). BOYACA:
15950 (TENN);33 km N of Ixtlan, Norris& Taranto Sierra Nevada del Cocuy, Cleef 5849 (U); carretera
16379 (TENN);betweenTuxtepecand Oaxaca,Sharp Hondo-Labranzagrande,Cleef 9281 (U); Paramo de
2304 (TENN);22.4 km N on Hwy. 175, Horton7589 Pisva, carreteraSocha-La Punta, Cleef 4550 (U).
Near Zotalapabridge W of Huau- CUNDINAMARCA:Paramo de Chipaque above Bogota,
(ALTA). PUEBLA:
chinango, Sharp 954 (MEXU); near Honey Station, Schultes 11407 (FLAS);Monserrate,Sabana de BoPringle10656 (B, BM, FH, H, NY, PC, S);nearOcos- gota, Schultes 12298 (FLAS);Choachi, Schultes s.n.
toc below Tezmitlan,Sharp3976 (TENN).VERACRUZ: (FLAS);Paramode CruzVerde(Bogota),Onraedt6392
Alrededoresde Chiconquiaco,Delgadillo 3975 (NY, (hb. Frahm);Guadelupe,Onraedt6229 (hb. Frahm);
MEXU); Camino a Ixhuacan, 12 km SW Teocelo, 10 km N of Guasca, King et al. C-691 (S); La Pena,
Juarez & Vazquez461 (NY); 5 km al N de Coatepec, Manzanos,Lindig 2009 (BM, GOET, H-BR, PC);La
Gil 156 (TENN);Los HuerfanosnearYecuautla,Sharp Vega,Apollinaires.n. (H-BR, NY, PC). San Crist6bal,
et al. 3035a (MEXU).
Apollinaires.n. (S); 18 km NE of Fusagasuga,King &
BetweenSanMarcos Guevara C-716 (S). CAUCA:Near Popayan, Alston 8040
GUATEMALA.SANMARCOS:
and San Rafael, Standley 86444 etc. (FH, NY, S). (BM,FH);Paramode Purac6,Killip&Lehmann38608
QUEZALTENANGO:Between San Martin and Colomba,
(PC);San Antonio, Pennell & Killip 7422 (FH, NY);
Mt. Purac6,Killip 6775 (FH, NY). NARINO:
Volcan
Civija, Sharp 5235
Standley s.n. (S). BAJA VERAPAZ:
Sta. Rosa
(MEXU).
Galeras,Plowman 1984 (FH). RISARALDA:
de Cabal, van Reenen et al. 3429 (U). SANTANDER:
COSTA RICA. ALAJUELA:
Sarchi, Alfaro 79 (NY).
Volcan de P6as, Pittier 5516 (NY); along road from Paramode las Vegas,Killip& Smith 15646 (FH, NY);
Vara Blancato La Concordia,Maxon & Harvey8426 Pamplona, Paramo de Fontibon, Alston 7154 (BM);
Km 73 on Pan American Picacho, carretera Bucaramanga-Cucuta,Hermann
(BM, FH, H, NY). CARTAGO:
Nevado de Tolima, van
Hwy. SE of El Empalme, Crosby5769 (MO); 10 km 25067 (B, H, NY, S). TOLIMA:
NW of summit at La Ascensi6nalong InterAmerican der Hammen & Jaramillo3316 (U).
VENEZUELA.MERIDA:
EntreElMorroy Aricagua,
Hwy., Crosby 6140 (MO). Cordillera Talamanca,
McDaniel6746 (NY); Irazu,Pittier5515 (NY, PC, S); ParamoDon Pedro,Ruiz-Terdn&Lopez9543 (FLAS);
Buena Vista-massif,CerroParamo,Kuhbier337 (B). Pico El Aguila, Distr. Miranda, Griffinet al. 1332
HEREDIA:Volcan Barba(S); Cerrosde ZurquiNE of (FLAS);ParamoLa Negraabove Bailadores,Griffinet
San Isidro, Standley 50581 (B, H, JE); Cerro de las al. 17461 (B, FH, FLAS,NY, S); Rio Capazarribade
Caricias,Standley 52137 (FH); 3.3 mi N of San Jose La Azulita,Steyermark&Rabe 97080 (B);Libertador,
de la Montana, Crosby3873 (MO). SANJOSt: Near betweenLaAguadaandLaMontania,Griffinet al. 1408
Turrialba,Schultes 11865 (FLAS);along InterAmer- (FLAS);SierraNevada,Teleferico,Cleef&Huber4812
ican Hwy. 13 km SE of El Empalme,Crosby6156 (H, (U); Mucuy, Onraedt5607 (hb. Frahm);La Aguada
MO); 67 km S of San Jose on Pan AmericanHwy., betweenMeridaand Pico Espejo,Steyermark&KoyaKoch 5078 (NY); San Jose, Schultes 11919A (FLAS); ma 102380 (NY). TRUJILLO:10 km al SE de Bocon6,
Volcan Irazu, Schultes 11839 (FLAS); Las Nubes, Steyermark104867 (FH); entre Trujillo y Bocon6,
Steyermark& Rabe 97302 (NY); Teta de Niquitao,
Standley38378 (NY).
JAMAICA.BlueMts.,Harris309 (BM,NY);Morces Paramode Cabimbu,Lopez 12014 (FLAS).
ECUADOR. Quito, Jameson 17 (BM, NY, PC).
Gap, Maxon & Killip 1327 (BM, NY); Cinchona,Underwood177 (NY); New Haven gap. Britton70 (NY); AZUAIU:Prope Chunchi,Allioni 8066 (H-BR). Cerro
St. Helen'sGap,Maxon 649 (S);SirJohn'sPeak,Cros- Antisana,Grubbet al. 2508b (BM).BetweenBanosand
Rio Verde, Bell 892 (BM). CARCHI:Tulcan-Maldoby 3045 (MO).
HAITI. Along road from La Descubiertoto Hondo nado road 32-36 km W ofTufino, Luteyn7898 (NY).
CHIMBORAZO: Humboldt s.n. (JE). NAPO: Road beValle, Reese 15358 (NY).
DOMINICAN REPUBLIC. LA VEGA:Cienaga de
tween Tena and Baeza, SteereE-148 (NY). PICHINCHA:
Benoist4602 (PC,S). Tunguragua,Spruce46 (BM,FH,

H-BR, NY, PC, S). Cutan supra Gualaquiza,Allioni
8057 (H-BR).
Liogier 24917 (NY). INDEPENDENCIA:Sierra de Neiba
alongHaitianBorder,Norriset al. 5960 (NY).LAVEGA: PERU. Pancarlambo,Jay s.n. (NY). Miradornear
Above La Siberia,road from Constanzato Valle Nue- Chinchao,Mexia 7769 (BM, FH). Matucana,Bryan
vo, Smith 10309 (NY); 13 km from Valle Nuevo to 346 (NY). Carabaya,Weddell1848 (PC). AMAZONAS:
San Jose de Ocoa, Norriset al. 7232 (H, NY); between StrasseCajamarca-Chachapoyas
zwischenBalsasund
La Constanza and La Nuez, Jones & Norris 1292A Leimebamba,Frahm823990 (hb. Frahm);21 km von
la Culata,Liogier5 (FH);road from San Jos6 de Ocoa
to Constanza,GasbarroB 10932 (NY, S);ValleNuevo,
Flora Neotropica
210
5.5ra
:II
77.E
FIG. 150. Pilopogonlaevis(Lindigs.n., H-BR). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of leaf.

E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 mm.
Ingenioin RichtungPomacocha,Hegewald7093 (H).
ANCASH:
Laguna Querococha, Hegewald 7673 (hb.
Frahm);Tunnel CahuishSW Huaraz,Hegewald7700
37 km NE Abancay an der
(hb. Frahm).APURMAC:
Strasse nach Cuzco, Frahm 823946 (hb. Frahm).
AYACUCHO:
An der StrasseHuanta-SanFrancisco 10
km S Manchete,Frahm 823901 (hb. Frahm).Chachapoyas, Calla-Callanear Leimebamba, Wurdack1325
(S). HuANuco: Carpish, Cerrate 6679 (MO). PUNo:
PC);bei Incacorral,Herzogs.n. (JE,PC);bei der Abra

de San Benito, Herzog s.n. (B, JE, H-BR, NY, S); bei
Samapata,Herzog4128 (JE);zwischenSan Mateound
Sunchal,Herzog 4436 (JE, S); bei Tablas, Herzogs.n.
(S).NearParadiso,SanJos6-Apolotrail, Williams1762
(BM, FH, NY, S). LAPaz: Along road from Soratato
Achacachi, Lewis 79-1283 (F); near Yungas, Rusby
3159 (NY); nearUnduavi, Eyerdam25140 (BM);Tolapampa, Williams 1761 (NY, S); Illampi, Troll s.n.
Valle Ollaches, Soukup 9455 (PC). SAN MARTIN: Km (S). SANTACRUZ:Comarapa,Hermann 24697 (NY).
387 an der StrasseMoyobamba-Chachapoyas,Frahm Antahnacana,7 miles SSWof Locotal,Hermann25242
823883 (hb. Frahm).
(FH, NY). Lago Colomi, along road to Corani, HerBOLIVIA. COCHABAMBA:
mann 24657 (FH).
Along carretera Villa
BRAZIL.MINASGERAIS:
Serrado Piaui, Schwecker
Tunari,Hermann25181 (B, BM, FH, H, NY, PC, S);
prope Choquecamata,Germain1208 (JE,H-BR, NY, 5277 (H-BR); Ouro Preto, Vital 5537 (SP). PARNA:
211
Ponte Grossa2 km suedl. ParqueNacionalVila Velha,
Frahm 1631 (hb. Frahm);NE of Curitiba53 km SW
of Parana-SaoPaulostateborderon BR 116, Vitt21467
(ALTA). Rio DE JANEIRO:Serra do Itatiaia, Ule 214
(BM, FH, HBG, GOET, JE, NY, PC, S); vic. Rio de
Janeiro,Glaziou 7065 (BM, NY, S); Petr6polis,Bandeira 181 (NY, S);Serrados Orgaos,Campodas Antas,
Rizzini 1047 (FH); Serrado Mar zwischen Paratiund
30
w Î
I)
2|0
All varieties of P. gracilis described (var. bernoullii C. Miiller, var. comigera C. Miiller, var.
divaricatus Herzog, var. parvus C. Miiller and
var. pittieri Renauld & Cardot) are expressions
of the broad variability in this widespread species
and are not of taxonomic value. This species
varies much regarding size and color. Even
blackish forms have been found. Occasionally,
the excurrent parts of the costaes may be subhyaline or even hyaline in perichaetial leaves,
which may cause confusion with normally hairpointed species as P. laevis and P. peruvianus.
The Brazilian population, which is disjunct
from the main range of this species, has even
been separated as P. subjulaceus. It differs by the
length of the setae, which is about 10 mm in P.
subjulaceus and about 18 mm in the Andean
population of P. guadeloupensis, but this seems
to be the only differing character and not sufficient to split P. subjulaceus from P. guadeloupensis.
Pilopogon guadeloupensis is a relatively common species in the neotropics. The list of specimens examined gives an insufficient impression
of the herbarium material, which has accumulated during the past hundred years and which
was collected mostly at the same localities. Thus,
about a hundred specimens have been collected
on the Blue Mountains of Jamaica alone, and
several dozens in the Itatiaia Mountains in Brazil.
2. Pilopogon laevis (Taylor) Theriot, Rev. Bryol.
Lichenol. N.S. 9: 12. 1936.
Figs. 6A, 150, 151.
CampylopuslaevisTaylor,LondonJ. Bot. 5:47. 1846.
Pilopogonellalaevis (Taylor) Bartram,Rev. Bryol.
Lichenol.6: 10. 1934. Type. Ecuador.Mt. Pinchincha, Jameson s.n. (holotype, FH).
PilopogonpiliferusHampe, Ann. Sc. Nat. Bot. ser. 5,
3:362.1865. Type.Colombia.Boquer6n,Lindigs.n.
(holotype, BM; isotypes, FH, GOET, H-BR, PC).
PilopogonnanusHampe,Linnaea32:137. 1863.Type:
0
O00
400
600
800
1000km
200300
500
600
miles
-.-- -I-
.H
--
--
Cunha, Frahm 1604 (NY). Rio GRANDESO SUL: Far-
roupilha,Vital9312 (SP).SAOPAULO:
Camposdo Jordao, Froehlich30 (S);propeApiahy,Puiggari22 (BM).
0i
FIG. 151.
g~~~~~.__
~
I-
Distribution of Pilopogon laevis.
Colombia. Bogota, Laches, Lindig 2010 (lectotype,

BM; isolectotypes, FH, GOET, H-BR, NY, PC, S).
Plantsto 4 cm high, darkbrownish,red brown

or almost blackish. Stems appressed foliate.
Leaves 5-6 mm long, narrow lanceolate. Costa
shortly excurrent in the lower stem leaves, in
upperstem leaves excurrentin a hyaline, serrate
awn;in transversesection with ventralsubstereids, ridgedat backwith distinctlyprotrudingcells.
Alar cells lacking. Basal laminal cells hyaline,
thin-walled, 6-9 x 35-45 ,tm. Upper laminal
cells oval to elongate-ovalor rhomboid, 7-11 x
18-22 um, incrassate.
Seta to 1 cm high. Capsule 2 mm long, dark
brown,long-rostrate.Peristometeeth 16, split to
the base into filiform prongs. Sporesca. 13 ,tm
in diam., smooth. Calyptracucullate, ciliate at
base.
Distribution(Fig. 151). On wet soil and soilcoveredrocks.A speciestypicalforparamovegein Costa Rica, Vento 4400m
tation from 2800
and
Peru.
Ecuador
ezuela,
Colombia,
Peru.
and
Ecuador
Colombia,
ezuela,
212
Flora Neotropica
::I . ?
~.,
'E
weX'?
FIG.152. Pilopogonlongirostratus
section
(Spruce47, H-BR).A. Plant.B. Leaf.C. Leaf-tip.D. Transverse
of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 inm.
Specimensexamined.COSTA RICA. Cordillerade
Talamanca,CerroParamo,Kuhbier334 (BR).
COLOMBIA. ARAUCA:
Cleef 8888 (U). BOYAC:Peiia de Amical N de Vado
Hondo, Cleef 9488 (U); Paramode Pisva, Cleef 4476
(U); Piramos al NW de Belen, Cleef2125 (U); Siberia
entrePefiaArical y Alto de Mogotes,Cleef9559a (U);
entreSogamosoy Vado Hondo, Cleefet al. 9218b (U).
CUNDINAMARCA: Paramo de Cruz Verde, Cleef 3560
(U); Paramo de Sumapiz, Cleef 10.396 (U); Paramo
de Palacio, Cleef3673 (U); ParamoentreCoguay San
Cayetano,Cleef 701 (U): Piramo del Boquer6n,Troll
AmericanHwy.to SantoDomingo,Cosby10815 (MO);

Cotopaxi,Gradsteinet al. GSV107 (U); Valleede Lloa,
Benoist 2461 (PC).
PERU. ANCASH:
LagoSafunaNE of Alpamayo,Bunin B-49878 (COLO).LagunaLlaca 27 km NE von
Paramo
ta, Rangelet al. 892 (COLO,FLAS,U). META:
der Strasse Huanta-San Francisco oberhalb Quinoa,
Sierra Nevada de Santa Mar2162 (JE). MAGDALENA:
de Sumapaz, Cleef7594 (U). RISARALDA:Sta. Rosa de
Cabal,Volcan de Sta. Rosa, van Reenen 733 (U).

VENEZUELA.
Merida: Paramo La Negra above
Bailadores,Griffinet al. 2151 (FLAS);SierraNevada

de Merida,between Aguadaand La Montafia,Griffin
et al. 257 (FLAS).
ECUADOR. PIcmNCHA:Along road 71 from Inter
An
Huaraz, Frahm 823882 (hb. Frahm). AYACUCHO:
213
Frahm823929(hb.Frahm);oberhalbTambo,Frahm
824009(hb.Frahm);PasszwischenTamboundQui- -~0
noa, Frahm823938 (hb. Frahm).HuANuco:Pass
zwischenTingoMariaundHuanuco,Frahm825113
(hb. Frahm). LA LIBERTAD:
Laguna Sausacochabei
6021(hb.Frahm);
Huamachuco,
Pampasde
Hegewald
la Juliabei Quiruvilca,
Hegewald5979(hb.Frahm).
Pilopogon laevis is closely related to P. peruvianus,from which it is distinguishedmainly by

the ridged,not lamellose, back of the costa. Both
species can occur (as in Peru) within the same
range,but are significantlydifferentin their ecology. Pilopogonperuvianus,with long dorsal lamellae at the costa, has a better possibility to
storewaterin these lamellaeand can growin dry
habitats, even in the coastal desert, whereas P.
laevis with a nearly smooth dorsal side of the
costa does not have the ability to storewaterand
depends on wet habitats such as paramos and
wet punas.
Sterile specimens of Pilopogonare nearly impossible to distinguish from species of Campylopus without experience.The species with protrudingcells or lamellae at the dorsal side of the
costa as P. laevis and P. peruvianuscan be distinguished from Campylopusby conspicuously
thickerapical cell walls of the outermost cells.
SterilespecimensofP. laevisare differentiated
from sterile specimens of P. guadeloupensisby
the reddish brown color and the ribbed abaxial
surface of the costa. These charactersand the
differentperistomecharactersindicateclearlythat
P. laevisis a good speciesand not merelya piliferous formofP. guadeloupensisas it has been commonly misinterpreted.
3. PilopogonlongirostratusMitten, J. Linn. Soc.
Bot. 12: 70. 1869. Type. Ecuador. "In Valle
Patasaad pedem montis Tunguragua,"Spruce
47 (holotype, NY; isotypes, BM, H-BR,
H-SOL, PC, S).
Figs. 152, 153.
Plants to 4 cm high, blackishbelow, yellowish
green above. Stems tomentose. Leaves 5-7 mm
long, narrowlanceolate;not appressedas in other
species of Pilopogon but erect spreadingwhen
dry. Costa half as broad as the leaf base, excurrent in a denticulatepoint, smooth at back. Alar
cells well differentiated,inflated,reddishbrown.
Basal laminal cells elongated, thick-walledand
pitted, 7-10 x 40-60 jm, narrowerand shorter
at margins.Upper laminal cells elongatedoval,
J)
-----------
.o
0 00,.
200oo00 600
0oo
0.oo
Vt
BIr
o *x2X
"b
^
500
Prep"redbyHendr k R.Rypk.ema
---------
--
-,*
"'----
20
r'
/
7Oi
FIG. 153. DistributionofPilopogon longirostratus

()) and Sphaerotheciumphascoideum(0).
incrassate, 3-6 x 16-24 ,m. Perichaetial leaves
short, reaching half the length of the seta.
Capsule 2 mm long, cylindric, (3-)4 times long
or longer as broad. Operculum longly rostrate, as
long as or longer as the urn. Peristome teeth 16,
filiform, split to the base, densely papillose. Calyptra cucullate, longly fringed at base. Spores ca.
13 ,um in diam., yellowish, smooth.
Distribution (Fig. 153). On soil in subalpine
rain forests, known only from Eucador and Colombia.
Specimensexamined.COLOMBIA.CUNDINAMARCA:Subachoque,Paramo El Tablazo, Frahm 885242
(hb. Frahm).
ECUADOR. Chonchin, Spruce s.n. (NY, S). Rio
Verde, Spruce s.n. (NY). Quito, Jameson s.n. (NY).
"Andes Quitenses," Spruce 46 (BM, NY). ZamoraChinchipe, road Loja-Zamorakm 17, Holm-Nielsen
et al. 3582 (S). PICHINCHA:
Vic. Rio Alambi, 0?5'S,
78?30'W,Crosby10739 (MO).
4. Pilopogon macrocarpus Brotherus, Rev. Bryol.
47: 3. 1921. Type. Ecuador. Azuay: Granadilla, Allioni 8061 (lectotype, H-BR).
Figs. 1C, 4A, 154, 155.
Plants to 4 cm high, slender, light to dirty green,
appressed foliate. Perichaetia bud-like, clustered
at top of the plants. Leaves 5-8 mm long, narrow
lanceolate, with excurrent costa. Excurrent part
of the costa nearly as long as the lamina or longer,
214
Flora Neotropica
"N
FIG. 154. Pilopogonmacrocarpus(Allioni8056, H-BR).A. Plant.B. Leaf.C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Am.
nearly smooth, the uppermost part sometimes

hyaline. Base of leaf longly decurrent.Alar cells
not differentiated.Basal laminal cells hyaline,
thin-walled, 6-12 x 60-100 ,um, narrowerat
margins.Upper laminal cells oval, rhomboid or
short rectangular,5-10 x 20-35 ,jm, incrassate.
Seta to 1 cm long, 2/3 or nearly totally surroundedby the perichaetialleaves. Capsules2.53.5 times longerthan broad, dark brown. Operculumlonglyrostrate.Calyptracucullate,smooth
at base.
Distribution(Fig. 155). On peaty soil in paramos and subalpine forests, 2700-3100 m, accordingto one ancient herbariumlabel also epiphyticin forests.This wouldbe the only epiphytic
occurrenceof the genus. Known only from Ecuador and Colombia.
Specimensexamined.COLOMBIA.CUNDINAMARCA:Cordillera
de LaLeonora60 kmNNEde Bogota,
Represadel Neusa, vanderHammen&Jaramillo3205
Paramode Cho(U);Sabanade Bogota,Villapinzon,
215
quas, Schultes12263 (FLAS);Subachoque,ParamoEl
Tablazo,Frahm885220, 885222, 885237 (hb.Frahm).
ECUADOR. AZUAY:Gualaquiza,Allioni 8056 (HBR). Without locality, Spruces.n. as Pilopogonspec.
nov. (FH). Quito, Jameson s.n. as P. inclinatus nom.
nud. (BM).
According to the type description this species
differs from P. guadeloupensis by a shorter capsule, absence of a peristome and a non-ciliate
calyptra. A study of the type specimen revealed
that some capsules have a normally developed
peristome, whereas it has fallen off with the lid
in others. Non-ciliate calyptras do also occur
rarely in P. guadeloupensis. This species, at least,
is known from too few specimens to decide
whether the capsule length is definitely shorter
than in P. guadeloupensis. It is separated here by
the very long-pointed stem leaves, which seem
to differentiate this species from P. guadeloupensis, which has only long-pointed perichaetial
leaves.
I200. .
0
CampylopusperuvianusWilliams,Bull.Torr.Bot. Club
42: 393. 1915. Type. Peru. Near Mollendo, Rose &
Rose 18997 (holotype,NY).
Plants to 5 cm high, strongly resembling the
common P. gracilis, yellowish or dirty brownish
green to blackish. Leaves 4-6 mm long. Costa
ending in a long, hyaline hairpoint, short in lower
leaves but forming a long, serrate awn in upper,
especially in perichaetial leaves, on dorsal side
with lamellae 2-4 cells high; end cells of lamellae
with thicker outer cell walls. Basal laminal cells
thin-walled, hyaline. Upper laminal cells oblique
to short oval, incrassate, 6 x 6-9 Am.
Seta to 1 cm long. Capsule ca. 2 mm long,
cylindric, broader at base. Peristome teeth filiform, split to the base.
Distribution (Fig. 157). On earth-covered rocks
and gravelly soil from sea level to more than
3000 m in Ecuador and Peru.
Specimensexamined.ECUADOR. "Andes Quitenses," Spruce48 (FH, PC).
PERU. ANCASH:Lago Safuna NE of Alpamayo, Bunin B 49878 (COLO). HUANUCO: Strasse HuarazHuanuco bei La Uni6n, Frahm 823890 (hb. Frahm).
LA LIBERTAD:
Huancamarca, Hegewald 5134 (H).
This species is easily distinguished from all

other species of Pilopogon by the dorsal lamellae
60
210
5. Pilopogon peruvianus (Williams) J.-P. Frahm,

Figs. 156, 157.
Lindbergia9: 104. 1983.
70
80
200 400 800 800 1OO

100 200 300 4400
0...
'
rr'J
600
omet
X )
:
?20
."
1
\.
'
(1
__
of Pilopogonmacrocarpus
FIG.155. Distribution
(@)and P. tiquipayae(0).
of the costa. Because of the absence of sporophytes in the type materialit had been described
as Campylopus.It is indeed, vegetativelyhardto
distinguishfrom Campylopus,much resembling
Campylopuspiliferssp. galapagensis,from which
it can be differentiatedin the sterile state only
by the thickerouter cell walls of the end cells of
the costal lamellae.
6. Pilopogon tiquipayae Herzog, Bibl. Bot. 87:
24. 1916. Type. Bolivia. Cumbre de Tiquipaya, Herzog 2655 (holotype, JE; isotypes, B,
PC, S).
Figs. 155, 158, 159.
Plants to 2.5 cm high, very slender, not
branched,in dense tufts,golden yellowish-green,
reddishat tips. Leaveserect,appressed,to 3 mm
long,narrowlanceolate.Costa 1/ of the leafwidth,
excurrent, in the upper part with a few teeth,
dorsallywith protuberantcells. Alarcells lacking.
Basal laminal cells thin-walled,lax, 8-12 x 4070 um. Upper laminal cells shortly rectangular,
8-12 x 12-16 um, slightly incrassate.
Flora Neotropica
216
5cm
FIG. 156. Pilopogonperuvianus(Hegewald5134, H). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection

of leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 ,tm.
Seta ca. 5 mm long. Capsule 1.5 mm long,

short cylindric, 2.5-3 times longer than broad,
pale brown,smooth. Operculum1 mm long, rostrate. Peristome teeth 16, undivided, filiform,
densely papillose. Spores ca. 13 um in diam.,
smooth to finelypapillose,yellowish brown. Calyptraciliate at base.
Distribution(Fig. 155).On banksand soil-covered rocks in subalpine to alpine regions from
2700 to 4100 m, found only in Bolivia.
In size, Pilopogon tiquipayaeresembles small

forms of P. guadeloupensis, which were describedas P. liliputanus.It is differentiated,however, from all other species of Pilopogonby relatively thin-walled, short rectangularor oval
upper laminal cells, by the costa with protuberant cells at the dorsalside, and short,coarseteeth
in the leaf apex. It is furtherconspicuousby the
reddishtingeof the stem tips and shortsetaewith
short capsules.
Mt.
Specimensexamined.BOLIVIA.COCHABAMBA:
Ann. Sci. Nat. Bot.
Tunari,Hermann25133 (NY). TARJA:Caste6n,Lewis 8. SphaerotheciumHampe,
79-538(F).
ser. 5, 3: 361. 1865. With the charactersof S.
217
phascoideum. Type species. S. phascoideum
Hampe.
A genuswith threespeciesworldwide,one each
in the neotropics,in southernAfricaand Sri Lanka. All species are known from only a few localities in a very small range.This and the disjunctoccurrencesindicatean old ageforthe genus.
The presentrangesseem to be relictsfrom larger
ranges in former climatic periods. For a short
review of the genus see Frahm (1986b).
70
80
0
^f
O?
J
~'f _e
200
>
100
400
200
600
300
400
800
1000km
500
600 miles
1. Sphaerothecium phascoideum (Hampe)

Hampe, Bot. Zeitschr.27: 867. 1869.
Figs. 153, 160.
.0.0
32:136.
HampeLinnaea,
phascoideum
Thysanomitrion
comosumHampe,Ann.Sci.
1863.Sphaerothecium
Nat.Bot.ser.5, 3: 361. 1865,nom.illeg.Type.Colombia.Bogota,LosLaches,2500m alt.,Lindigs.n.
(holotype,BM;isotype,NY).
Th6riotRev. Bryol.Lichenol.
Campylopus
bogotensis
11:60. 1939.Type.Colombia.Bogota,ElBoquer6n,
Troll2248 (holotype,PC).
Plants in very low tufts, less than 7 mm high.
Stems 3-5 mm high,radiculosebelow.Leavs2.54 mm long, fromoblongbase graduallynarrowed
into a narrow,chanelledtip, marginsentire.Costa filling l/2 of the leaf base, excurrent,in transverse section with ventral and dorsal stereid
bands. Alar cells differentiated,incrassate.Basal
laminal cells short rectangular,incrassate,ca. 45:1, in 3-5 rowsnarrowerand shorterat margins.
Upper laminal cells rectangular,incrassate,ca.
3-4:1.
Seta 3-4 mm long, curved. Capsule rounded
to ovoid, immersed amongst the perichaetial
leaves. Peristome small, on a low basal membrane, hardly exceeding the mouth of the capsule. Annulus huge, 150 gm high, dehiscent.
Spores about 21 ,m in diam. Calyptra small,
cucullate,not ciliate at base.
Distribution.Known only from the two type
specimens near Bogota, Colombia, at elevations
of 2500 and 3400 m.
Campylopusbogotensisis placedhere as a synonym of S. comosumwith some doubt, because
the type materialis sterile. Sphaerotheciumcan
ultimatelynot be distinguishedvegetativelyfrom
Campylopus,but is very distinct in the sporophytic characterswith extremelyshort setae and
globose capsules immersed in the comal tufted
upper leaves. Immersed capsules also occur in
011
....
U.
~i20
FIG. 157. Distributionof Pilopogonperuvianus.
Campylopus subg. Campylopidulum,which is

representedin the neotropicsby C. carolinae.It
differs,however, by smaller spores of about 13
am in diameter,no dehiscentannulusand a larger, dicranoidperistome.
PARALEUCOBRYOIDEAE
Brotherus, 1924
Plants variablein size, 3-80 mm high, in loose
tufts or tight cushions. Stems radiculose,erect.
Leaves erecto-patent or homomallous, lanceolate from ovate on base, endingin a narrowoften
218
Flora Neotropica
~--'..Y;?
...
:~~~~~~~~~~~~~~~~~~~~~?-
~~~~~~~~~~~~~~~~~~~~~~~~~~~~I
v' i
FIG. 158,
Pilopogon tiquipayae (Hermann 251331, NY).
219
1f
3mm
2.5cm
:;
:2O
EE
FIG. 159. Pilopogontiquipayae(Herzog2655, JE). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of

leaf. E. Basal laminal cells. F. Upper laminal cells. Unlabeledscale bar for microscopicdetails = 50 ,im.
canaliculatepoint.Leafmarginsentire,rarelywith longly rostrate. Peristome teeth entire or split

a few teeth at leaf tips. Perichaetialleaves with into two prongs. Calyptracucullate. Spores 10broaderbase and narrowercosta. Costa filling '/ 35 stm in diam.
to %of the leaf base, fillingthe subula, in transverse section with a median band of chlorocysts
and dorsal and ventralbands of hyalocysts.Alar
Key to the Genera of
cells weaklydifferentiatedor distinct. BasallamParaleucobryoideae
ina cells longly rectangular,narrow, relatively
1 Costa in transversesection without stereids;capthin-walled.Upper lamina cells elongate.
sules with stomata. .........
3. Paraleucobryum.
Seta straightor flexuose and twisted. Capsule 1 Costa in transversesection with median
groups
of stereids;capsuleswithout stomata.
erect, symmetric,ovate to longlycylindrical.Lid
220
Flora Neotropica
91
": __ .
rAq
FIG. 160. Sphaerotheciumphascoideum(Lindig s.n., NY). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse

50 gm.
2 Peristome teeth split; costa in transversesection with 2-4 ventral stereids; brood leaves
1. Campylopodiella.
lacking. ...............
2 Peristometeeth entire;costa in transversesection without ventralstereids;brood leaves frequent in the axils of the upperleaves. ......
...............................
.2. Brothera.
1. Campylopodiella Cardot, Bull. Herb. Boissier
ser. 2, 8: 90. 1908. Type species: Campylopodiella tenella Cardot.
Plants in loose to compact tufts, 5-30 mm

high. Stems radiculose,often with flagelliferous
branches. Leaves erecto-patent, rarely homomallous, lanceolate,ending in a long narrowsubula, entire or with a few teeth at tips. Perichaetial leaves frombroaderbase suddenlycontracted
into a long, slender subula. Costa filling
1/2-2/3
of
the leafbase,excurrent,in transversesectionwith

large ventral and dorsal hyalocysts, a median
band of groups of stereid chlorocysts and 2-4
221
'*I
i''
\I
'!
* 0.c \
Anr
FIG 11.
setinoflaf
\I
II/
'-'i
/'
amylpodela fagllcea(Hrzo
. asllaialcll.F.Upe
yX
457
lmna
E~~ '
JE. . Pan. . Laf C.Lef-ip.D.Trasvrs

els Ulaeedsal
arfr
icocoi
dtil
^'
FIG. 161. Campylopodiella flagellacea (Herzog 4357, JE). A. Plant. B. Leaf. C. Leaf-tip. D. Transverse
50 um.
ventral stereids. Alar cells weakly developed.

Laminalcells elongate rectangular.
Seta yellowish or brownish in age, erect and
twisted or flexuose. Capsule erect, elliptical to
cylindrical, yellowish, without stomata. Operculum longly rostrate. Peristome teeth divided
nearly to the base. Spores 11-19 um in diam.
Calyptracucullate, fimbriateor entire at base.
The ventral stereids are visible without dissecting the costa, in view upon the ventral side
of the leaf as a darker band. The leaf is thus
differentiatedinto the lamina, the transparent
marginal part of the costa and the darker central

part of the costa. This makes it relatively easy
to distinguish Campylopodiella from related genera such as Brothera, or certain species of Campylopus.
Key to Species of Campylopodiella

1 Lamina cells incrassate,becoming quadratetowardsthe leaf base; alar cells distinct, inflated..
.............................
1. C.flagellacea.
1 Laminalcells thin-walled,becoming rectangular
Flora Neotropica
222
110
30
100
80
90
70
60
I.
300
--
L__R
i~
00
L-r lO-, 30 40 50
7 .6
..(------ -:---
..
FIG. 162. Distributionof Campylopodiellaflagellacea(0) and C. stenocarpa(0).

towardsthe base;alar cells weaklydifferentiated,
not inflated ..................
2. C. stenocarpa.
1. Campylopodiella flagellacea (C. Muller)
Frahm & Isoviita, Taxon 37: 968. 1988.
Figs. 161, 162.
Dicranumflagellaceum C. Miller, Syn. musc. frond.
2: 597. 1851. Campylopusflagellaceus (C. Miller)
Mitten, J. Linn. Soc. Bot. 12: 77. 1869. Dicrano(C. Muller)Williams,N. Amer.
dontiumflagellaceum
Fl. 15: 153. 1913. Metzlerellaflagellacea(C. Muller)
Brotherusin Engler& Prantl,Nat. Pflanzenfam.2,
10: 191. 1924. Type. Mexico. Michoacan: Cerro
Andres,Chrismars.n. (holotypedestroyedat B; lectotypus nov., PC).
Campylopodiellamalagensis (Herzog) J.-P. Frahm,
Bryologist87: 250. 1984. Campylopusmalagensis
Herzog,Biblioth. Bot. 87: 23. 1916. Type: Bolivia.
Cerrode Malaga,Herzog4367 (holotype,JE;isotype,B).

Plants erect, yellowish green, in tufts. Stems
3-15 mm high, radiculose,equally foliate with
erectpatentleaves when wet or appressedleaves
when dry, often with flagelliferousbranches in
the axils of the upperleaves. Leaves 1.8-2.3 mm
long, lanceolate, graduallycontracted, margins
entire, only slightly denticulate at the extreme
apex. Costa filling 1/22/3 of the leaf base, 155 fum
wide, excurrent,in transversesection with large
ventraland dorsal hyalocysts,a median band of
stereidsand 2-4 stereidsventrally.Alar cells hyaline or brownish, often bistratose, slightly inflated.Basal laminal cells in about six rows, rec-
223
t .
S
, E
*vH
~~~~~~
}
ofE
t~~~~~~~''~~~
FIG. 163. Campylopodiellastenocarpa(Richardset al 3009, hb. Frahm).A. Plant. B. Leaf. C. Leaf-tip.D.

= 50 ,um.
details =
Am.
tangular to subquadrate, 15-40 x 10-16 ,tm,

narrowerat margins.Upper laminalcells rectangular, 13-30 x 4-12 zm.
Seta ca. 10 mm long, light brown, sinuose.
Capsule1.5 mm long,lightbrown,slightlyribbed.
Operculumas long as the capsule,obliquelyrostrate. Calyptracucullate,entire at base.
Distribution(Fig. 162). On peaty soil, earthcovered rocks, on roofs, at base of trees in elevations between 2700 and 4000 m in Mexico,
Guatemala,Venezuela, Colombia and Bolivia.
W of La
Specimens examined: MEXICO. DURANGO:
Ciudad,Gillet 17053 (hb. Frahm).

GUATEMALA.
HUEHUETENANGO:
Above Todos
Santos, Sharp 4794 (MO); San Marcos, slope of Tajumulco, Sharp 5462 (F).
COLOMBIA."AndesBogotenses,"Sebate,Weir117
(H-BR, MO, NY).
VENEZUELA.MERIDA:Paramode Don Pedroentre
El Morroand Aricagua,Ruiz-Terdn9559 (FLAS).
This species is usually found sterile. A sporophyte has been found only in the specimen
from Venezuela.
Flora Neotropica
224
2. Campylopodiella stenocarpa (Wilson) P.
Muller& J.-P. Frahm,Nova Hedwigia45: 290.
1987. TrichostomumstenocarpumWilson in
Seemann,Bot. voy. Herald:344. 1857. Atractylocarpus stenocarpus (Wilson) Zander,
Bryologist85: 128. 1982. Type:Mexico. Sierra
Madre,Seemann 1925 (holotype, BM). Figs.
1A, 4E, 162, 163.
GUATEMALA. QUICHE:Los Cinquentos, Richards
et al. 3009 (hb. Frahm);SE of Chichicastenango,Richards s.n. (hb. Frahm).

COSTA RICA. ALAJUELA. Near San Jose, Crosby
6270 (hb. Frahm).
Campylopodiellaflagellaceagives the impression of a depauperateform of C. stenocarpa,because the shorterleaves of C.flagellacea resemble the leaves of flagelliferous shoots of C.
Plants yellowish green, erect, in tufts. Stems stenocarpain shape. However, in C. flagellacea
8-15 mm high, radiculosebelow, equallyfoliate, the alar cells are always more well developed,
erecto-patentwhen wet, appressedfoliate when the basal laminal cells are more incrassateand
dry, sometimes with flagelliferousbranches in the upper laminal cells shorter.
the axils of the upper leaves. Leaves 3.5-5 mm
Homomallous leaves are rarelyfound in this
long, lanceolate,from ovate base graduallynar- genus, as in a specimen collected by Maslin s.n.
rowed into a long acumen, entire at marginsex- from Mexico (NY).
cept for few teeth at the leaf tip. Perichaetial
leaves suddenlycontractedfrom a broadlyovate
2. BrotheraC. Muller,Generamusc. frond.258.
base. Costa filling /3-2/5 of the leaf base, 175-200
1901. Type: B. leana (Sullivant)C. Muller.
,um wide, excurrent,in transversesection with
ventraland dorsal hyalocysts,a median band of
Monotypicgenus with only the followingspestereidsand 2-3 ventralstereids.Alarcells weak- cies.
ly differentiated.Basal laminal cells in 15-18
rows, hyaline,thin-walled,rectangular,45-95 x 1. Brotheraleana (Sullivant)C. Muller,Genera
4-20 im, narrowerat margins. Upper laminal
musc. frond. 258. 1901.
Fig. 164.
cells elongate, 74-140 x 4-7.5 um.
leanus (Sullivant) Sullivant & LesqueSeta erect,8-19 mm high, sinistrorselytwisted Campylopus
reux, Musci bor.-amer.18. 1856. Leucophanesleain the upper part. Capsule(without operculum)
num Sullivant,Musci allegh. 41. 1846.
long-cylindric,0.4 x 2.5-3 mm, yellowishgreen Syrrhopodonleanus (Sullivant)Sullivant, Lesquereux
& James,Man.N. Amer.Moss.78. 1884.Leucoor brownish in age. Exothecial cells incrassate,
leanum(Sullivant)
Eur.N. Amer.
bryum
Kindberg,
rectangular.Stomata lacking. Operculumlongly
Bryin.2: 176. 1897. Type:U.S.A. MusciAllegh. 41
rostrate,reddishbrown, 1-1.2 mm long.Annulus
(holotype,NY; isotypes BM, G).
present. Peristome teeth divided nearly to the andmoresynonymsfromAsia.
base, yellowish brown, lighter at tips, striate at
Plants small, only 3-6 mm high, in dense, yelbase and papillose at tips, about 290 ,umlong.
lowish to yellowish-greentufts. Stems radiculose
Sporesyellowish green, 11-13 ,tm. Calyptracu- at
base, frequentlywith clustersof brood leaves
cullate.
in the comal tufts. Leaves 2-3 mm long, erectoDistribution(Fig. 162).On rottenlogs, stumps,
when wet, curled when dry, lanceolate,
burnedwood, on soil and the bases of trees, at patent
contractedto a short,canaliculateapex;
gradually
1100-2900 m, in Mexico,Guatemala,CostaRica,
entire. Perichaetialleaves with broader
margins
Hondurasand Panama, also found once in the
base, abruptlycontracted to a narrow leaf tip.
USA.
Costafilling2/3of the leaf base, 155-180 ,m wide,
not sharply differentiatedfrom the lamina, exSpecimens examined. MEXICO. CHIAPAS: San
Christ6bal de Las Casas, Hermann 26419, 26417; current,in transversesection with lax dorsaland
Maslin s.n. (NY); PUEBLA:Popocatepetl, Eggers &
ventralhyalocysts(ventrallyoften with two rows
Frahm 792235 (hb. Frahm). MICHOACAN:
Las Peras,
Norriset al. 15583 (NY); Lagunade Vaca, McGregor of hyalocysts), and a median band of stereids
16543 (NY). MEXIco:Lagunasde Zempoala,Eggers (becoming substereidal towards the margins).
& Frahm 792474 (hb. Frahm).MORELOS:
Zempoala, Alar cells weakly differentiated,hyaline. Basal
Diill. s.n. (hb. Frahm).OAXACA:
BeyondIxtlande Jua- laminal cells hyaline, consisting of 8-10 rows,
rez,SharpM 59179b(hb.Frahm);SierraJuarez,Smith bistratose
towardsthe costa, rectangular,19-55
et al. 418A (hb. Frahm);Ixtlan, Norris et al. 16375;
Tellez et al. 4105 (hb. Frahm). NAYARIT:La Cienaga,
Norriset al. 14408 (hb. Frahm).
x 4-19 Am, narrower at margins. Upper laminal
cells rectangular,24-55 x 3-7 im.
225
FIG. 164. Brotheraleana (Sullivant172, BM). A. Plant. B. Leaf. C. Leaf-tip.D. Transversesection of leaf.
E. Basal laminal cells. F. Upper laminal cells. Unlabeled scale bar for microscopicdetails = 50 Tm.
Seta 5-6 mm long, yellowish, sinuose and

twisted sinistrorselyin the upper part. Capsule
erect, yellowish green, 0.5 x 1.2 mm, smooth,
contracted towards the peristome. Exothecial
walls incrassate,variable, without stomata. Annulus present. Peristome teeth entire (or rarely
perforated("windowed")in fully developedcapsules), yellowish, striateat base and papillose at
tips, 210-240 um long. Spores yellowish green,
smooth, 10-13 Am. Calyptracucullate, fringed
at base.
Distribution.On rottenwood, on barkat bases
of trees, on humic or peaty soil and on rocks

from 200 to 2500 m in Central America (Mexico,
Guatemala), also in North America (Arkansas to
Virginia) and in Asia (Himalaya, East Asia).
Specimensexamined.MEXICO.JALISCO:
Nazatlan,
Pringle 15269 (JE).
GUATEMALA.
HUEHUETENANGO:
Sharp 4939a
(TENN).
3. Paraleucobryum (Lindberg ex Limpricht)
Loeske, Allg. Bot. Z. Syst. 13: 167. 1907. Dicranum subg. Paraleucobryum Lindberg ex
226
Flora Neotropica
cells.
section of leaf. E. Basal laminal
am.
50 ,um.
F. Upper laminal cells. Unlabeled scale bar for microscopic details =
Limpricht, Laubm. Deutschl. 1: 373. 1886.

Type species. P. enerve(Thedenius)Loeske.
Plants dioicous, in dense tufts, whitish to dark
green.Stems to 8 cm high, erect,often branched,
radiculose.Leaves 4-8 mm long, erect or homomallous, lanceolate, ending in a long, canaliculate apex. Leaf tips serrateor smooth. Perichaetial leaves broaderat base, suddenly contracted
to a narrow subula. Costa filling
1/43-
of the leaf
width, ridged at back or smooth, in transverse

section with ventral and dorsal hyalocysts and
median or median and dorsal chlorocysts.Alar

cells differentiated,inflated, lax. Basal laminal
cells elongate, hyaline, thin-walled, slightly pitted. Upper laminal cells elongate.
Seta erect, 10-13 mm long, brownish,twisted
dextrorselyin the upper part. Capsuleerect, cylindrical,symmetric.Operculumlongly rostrate.
Annulus lacking. Peristome teeth divided into
two prongs. Exothecial cells incrassate, rectangular.Stomata in two rows near the base of the
capsule. Spores 13-34 ,jm. Calyptrayellowish,
cucullate,entire at base.
:, I
. -o
227
..
-- --.
"DO
30\ 200
Heondb
Prepared
by
FIG. 166.
40050-0 60
..
. -----..
k R Rypkema
1 Costa in transversesection without dorsal chlorocysts,smooth at back, filling3/4 of the base leaf
or more; leaf tips nearlyentire ..... 1. P. enerve.
1 Costa in the median and upperpartin transverse
section with dorsal chlorocysts, ridged at back,
1/3-2/3 of
...............
,1
--
Distributionof Paraleucobryumenerve(-) and P longifoliumssp. brasiliense(0).
Key to the Neotropical Species of

Paraleucobryum
filling
the leaf base; leaf tip serrate ....
2. P. longifoliumssp. brasiliense.
1. Paraleucobryumenerve (Thedenius) Loeske,

Allg. Bot. Z. Syst. 13: 167. 1907. Dicranum
enerveThedeniusin Hartman,Handb. Skand.
fl. ed. 5: 393. 1849. Type:Sweden.Herjedalen,
Skarfjellet,Thedeniuss.n. (holotype,H-SOL).
Figs. 165, 166.
Plants in dense whitish green tufts. Stems to
8 cm, rarely12 cm high, often radiculoseat base.
Leaves erect appressed or erect patent, rarely
homomallous,4-5 mm long, lanceolate,with entire margins and entire leaf tips. Costa at leaf
base 475-525 Am broad, filling
3/4-4/
of the leaf
width, excurrent,smooth at back, in transverse

section with a single median band of chlorocysts
and ventraland dorsal hyalocysts.Alar cells inflated,large,lax, protrudinginto the costa. Basal
laminal cells thin-walled,weakly pitted, in 8-10
rows elongate,47-72 x 7.5-13 Am.Upper laminal cells elongate, 47-90 x 5.5-7 Am.
Seta golden yellow to brownish, to 16 mm
high, erect, twisted dextrorselyin the upperpart
(as seen from above). Capsule yellowish to
brownish,2.5 mm long without operculum,cylindrical, smooth. Operculum longly rostrate.
Annulus present. Peristome teeth divided, dark
red to brown,lighterat tips, 300 Amlong. Spores
verrucose, 19-26 ,m in diam. Exothecial walls
rectangular,incrassate.Stomatapresentnearbase
of capsule. Calyptracucullate, yellowish, entire
at base.
Distribution(Fig. 166).Acidophilouson earthcovered rocks, in fissuresof rocks, on humic or
Flora Neotropica
228
..
' I.
fit:,
:X
FIG. 167. Paraleucobryumlongifoliumvar. brasiliense(Dusen s.n., H-BR). A. Plant. B. Leaf. C. Leaf-tip.

= 50
details
details=~
50,u~m.
,m.
peaty soil in subalpine to alpine regions, most

frequentlyabove the tree line in Europe, Asia,
Northand CentralAmerica(fromAlaskathrough
the Rocky Mountainsto Mexico), also in Greenland.
This specieshas been reportedalso from Brazil
(Yano, 1981, as P. albicans(Schwaegr.)Loeske)
and Peru (Hegewald& Hegewald, 1975). These
records are referred,however, to literaturereferencesof Leucobryummegalophyllumand thus
it seems these records concern the type of Di-
cranum albicans Schwaegr., which consists of a

species of Leucobryum.
Specimens examined. MEXICO.
DURANGO:
El Salto,
Crum 1851 (DUKE, NY); Nevado de Toluca, Delgadillo 1866 pp. (DUKE, NY). HIDALGO: Mineralel Chico, Sharp et al. 1768 (hb. Frahm). PUEBLA:Pico del
Orizaba, Delgadillo 1334, 1312 (B, NY, U). VERACRUZ:
Cofre de Perote,De Luna 297 (DUKE).
2. Paraleucobryum longifolium (Hedwig) Loeske

ssp. brasiliense (Brotherus) P. Muller & J.-P.
LiteratureCited229
Frahm,Nova Hedwigia45: 303. 1987. Para- and gave critical comments and suggestionson
leucobryumbrasilienseBrotherus,Denkschr. the taxonomy of some species. The illustrations
Akad.Wiss.Wienmath.-nat.K1.83:259.1926. were prepared by Doris Wamper and Birgit
Type:Brazil.Serrado Itatiaia,Dusen s.n. (ho- K6chling (Campylopus),Monika Giese (Camlotype, H-BR).
Figs. 1B, 167. pylopodium, Dicranodontium, Microcampyloand Pepus), Monika Padberg(Atractylocarpus)
Plants greyish to dark green, to 4.6 cm high, tra Muller(Paraleucobryoideae).
Thanksaredue
in compacttufts.Stemsradiculoseat base.Leaves to the curatorsof the herbariaB,
BM, G, GOET,
usually stronglyfalcate, 8-13 mm long, lanceo- F, FH, H(-BR, -SOL),JE, KIEL, L, MO, PC, S
late, ending in a long, canaliculatesubula. Leaf and U for the loan of specimens. The herbaria
tips and marginsserratein the upperthirdof the BM, F, FH, H, MO, PC, and S providedworking
leaf. Costa3/4of the leaf width, exurrent,in trans- facilitiesduringherbariumvisits. The New York
verse section with ventral hyalocysts, median BotanicalGarden(NY) made it possibleto work
chlorocystsand dorsalhyalocystsalternatingwith for six weeks in that herbarium.Other herbaria
chlorocysts, ridged at back. Alar cells inflated, (ALTA, COL, COLO, FLAS, INPA, MEXU,
hyaline.Basallaminalcells hyaline,thin-walled, PRC, QCA, SP, TENN and the private herbarslightlypitted, rectangularto elongate, 32-68 x ium E. & P. Hegewald)contributedspecimens
7.5-11 um. Upper laminal cells elongate, nar- for identification.The German ResearchFounrower at margins, 38-85 x 3.8-9.5 ium.
dation (DFG) supported fieldwork in Brazil,
Mexico and Peru and visits to the above menDistribution(Fig. 166). At the bases of sub- tioned herbariaby travel grants.
alpine shrubs(Drimyssp.), on soil and in small
niches of rocks("sapieiras"),describedfrom the
LITERATURECITED
ItatiaiaMountainsin SE Braziland found there
only at elevations from 2500-2800 m.
Arts, T. 1987. The occurrenceof tubersin Campy-
lopuspyriformis.Lindbergia12: 125-128.
Specimens examined. BRAZIL. Rio DE JANEIRO: Bartlett,J. K. & J.-P. Frahm. 1983. Notes on CamParque Nacional Serra do Itatiaia, Agulhas Negras,
pylopusand Chorisodontiumfrom New Zealand.
Frahm 1420 (hb. Frahm);Vital 7245 (SP).
J. Bryol. 12: 337-341.
Bartram,E. B. 1949. Mossesof Guatemala.Fieldiana
This species resembles ParaleucobryumlonBot. 25. 442 pp.
gifoliumssp. longifoliumin the leaf morphology, Biebl, R. 1964. Austrocknungsresistenztropischer
UrwaldmooseaufPuerto-Rico.Protoplasma59(2):
the strongly hamate leaves and especially the
277-297.
transversesection of the costa. SubspeciesbrasiM., R. B. Pierrot& T. Pocs. 1974. Troisgenres
liensediffers,however,by the largerleaf size, the Bizot,
nouveauxdes muscinees.Rev. Bryol.lichenol.N.S.
extremely hamate leaves, the wider costas and
40: 25-31.
the somewhat larger laminal cells (a character Bridel, S. E. 1819. MuscologiaeRecentiorumSupplementum.Gotha.
correlatedwith the largerleaf size). Subspecies
V. F. 1901. Musci. In A. Engler& K.
Brotherus,
next
occurs
and
a
boreal
is
species
longifolium
Die natiirlichen Pflanzenfamilien 1(3).
Prantl,
in the Rocky Mountains(Colorado).Thereis no
Leipzig.
1924. Musci. In A. Engler& K. Prantl,Die
explanationat present for this unusual disjuncnaturlichenPflanzenfamiliened. 2, 10, Berlin.
tion of ssp. brasiliense,which is the only known
Catcheside,D. G. & J.-P. Frahm. 1985. Additions
occurrenceof this species in the tropics.
to the Campylopusfloraof Australia.J. Bryol.13:
Brotherus
wide
of
the
because
costa,
Perhaps
359-367.
(in the type description)had comparedthis taxon Crum, H. A. & L. E. Anderson. 1981. Mosses of
eastern North America. Vol. 1. Columbia Uniwith Paraleucobryumenerve.This species, howversity Press,New York.
ever, has a totallydifferentstructureof the costa.
Crundwell,A. C. 1979. Rhizoids and moss taxonomy. Pages 347-363 in G. C. S. Clarke& J. G.
Duckett,Bryophytesystematics.Chapman& Hall,
ACKNOWLEDGMENTS
London.
A. & J. Florschiitz-deWaard. 1974.
I wish to thank Drs. J. L. Luteyn, S. A. Mori Florschiitz,P.
Studieson ColombiancryptogamsI. Variationof
and S. R. Gradstein for their care during the
charactersin South Americanspecies of Campypreparationof the manuscript.Drs. B. Allen and
lopus.J. Hattori Bot. Lab. 38: 111-114.
W. R. Buck kindly corrected the English text, Florschiitz-deWaard, I. & P. A. Florschiitz. 1979.
230
Flora Neotropica
EstudiossobreCript6gamasColombianasIII. Lis. 1982c. New CampylopusrecordsfromSouth

ta comentadade los Musgosde Colombia.Bryoland CentralAmericaII. Bryologist85: 322-323.
. 1982d. TaxonomischeNotizen zur Gattung
ogist 82(2): 215-259.
& M. Worrell-Schets. 1980. Studies on CoCampylopusXII. CryptogamieBryol.Lich6nol.3:
lombian cryptogamsVII. Culturestudies on the
59-66.
taxonomic relevance of costal anatomy in the
. 1982e. Problems of infragenericclassification in Campylopus.Nova HedwigiaBeih.71:219complexand
Campylopusleucognodes-subconcolor
in Campylopuspittieri. Proc. Koninkl. Nederl.
224.
Akad. Wetensch.Series C, 83(1): 37-45.
. 1983a. A monograph of Pilopogon Brid.
Frahm, J.-P. 1974. Zur Unterscheidungund VerLindbergia9: 99-116.
. 1983b. A new infragenericclassificationof
breitungvon Campylopusintroflexus(Hedw.)Brid.
und C. polytrichoidesDe Not. Revue Bryol. Lithe genus CampylopusBrid. J. Hattori Bot. Lab.
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. 1975a. Conspectus der mittel- und siid. 1984a. CampylopusBrid. 1. SubgenusThyamerikanischen Campylopus-Arten (Dicranasanomitrion (Schwaegr.) Kindb. emend. J.-P.
Frahm.Nova Hedwigia39: 585-621.
ceae). Bryoph.Biblioth. 5. 144 pp. Vaduz.
. 1975b. Revision der Gattung Thysanomi1984b. A survey of the genus Campylopus
triumin Sudamerika.J. Bryol.8: 255-264.
in Sri Lanka.J. Bryol. 13: 163-192.
. 1975c. TaxonomischeNotizen zur Gattung
. 1984c. Dicranaceenaus Peru. Nova HedCampylopus.Rev. Bryol. Lichenol.41: 321-332.
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1976a. TaxonomischeNotizen zur Gattung
1984d. A reviewof Campylopodiella.BryolCampylopusII. Rev. Bryol. Lich6nol. 42: 603ogist 87: 249-250.
616.
. 1985a. Afrikanische Campylopus-Arten.
.1976b. TaxonomischeNotizen zur Gattung
Bryophyt.Biblioth. 31. 216 pp.
CampylopusIII. Rev. Bryol. Lich6nol. 42: 8911985b. TaxonomischeNotizen zur Gattung
908.
CampylopusXIII. CryptogamieBryol. Lich6nol.
1976c. TaxonomischeNotizen zur Gattung
6: 205-208.
. 1985c. TaxonomischeNotizen zur Gattung
CampylopusIV. Nova Hedwigia28: 131-136.
1977. Zur Campylopus-Floravon Mexico.
CampylopusXIV. Nova Hedwigia41: 273-277.
. 1986a. Ecosystematicsand geotaxonomyof
Bryologist80: 119-124.
- . 1978a. Ubersichtder Campylopus-Arten
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INDEX OF SCIENTIFICNAMES
Anisothecium3, 22
Aongstroemia
curviseta200
euphoroclada37
fendleri 200
itatiaiense200
pilopogon200
sartorii200
Atractylocarpus3, 4, 5, 9, 10, 11, 12, 14, 18, 20, 21,
23, 25, 197
alpinus 18, 25
brasiliensis20, 25, 26, 27*, 28*
costaricensis26
flagellaceus26
longisetus7, 11, 15, 16, 17, 20, 26, 29*, 30*
madagascariensis18, 20
mexicanus25, 26
nanus 20, 26, 28*, 31, 32*
stenocarpus26, 224
strictulus26
Bartleya3
Bizotia 160
densifolia 159
Brothera3, 4, 5, 8, 10, 12, 18, 23, 220, 221, 224
leana 16, 19, 20, 224, 225*
Bryohumbertia3, 4, 10, 12, 14, 15, 17, 18, 20, 21,
25, 31
filifolia 15, 17, 19, 20, 22, 31, 35, 80
var. filifolia32, 36
var. humilis 31, 35
var. longifolia32, 36
flavicoma35
metzlerelloides20, 31
Bryopteris19
Bryotestua3
Calymperaceae9
Campylopidulum3
Campylopodaceae21
Campylopodiella2, 3, 4, 5, 16, 18, 19, 23, 25, 26, 220,
221
crenulata19
flagellacea19, 23, 221*, 222*, 224
himalayana19
malagensis222
stenocarpa7, 11, 12, 19, 23, 222*, 223*, 224
tenella 220
Campylopodioideae2, 3, 6, 8, 10, 12, 14, 22, 23, 24
Campylopodium2, 3, 4, 5, 9, 10, 12, 14, 16, 18, 22,
23, 24, 36, 200, 203
curvisetum200
euphorocladum36, 37
fendleri 200
itatiaiense200
lineare 37
medium 7, 11, 16, 22, 37, 201
pilopogon200
pusillum 37, 200
sartorii200
Campylopus2, 3, 4, 5, 6, 8, 9, 10, 12, 14, 15, 16, 17,
18, 19, 20, 21, 22, 23, 25, 37, 39, 200, 213, 215,
221
sect. Campylopus13, 39, 47

sect. Filifolii 31
sect. Homalocarpus13, 39, 160
subg. Campylopidulum10, 12, 39, 67, 105, 217
subg. Campylopus39
subg. Thysanomitrion5, 9, 10, 12, 21, 38, 47
acervatus143, 146
aemulans5, 20, 39, 41, 42*, 43*, 44
albidovirens 16, 20, 40, 43*, 44, 45*, 46, 47, 58,
118, 160
alopecurus112, 142
var. bolivianus142
altissimus 118
alto-filifolium32
amboroensis20, 39, 46*, 47, 48*
anderssonii20, 40, 47, 49*, 50*
andicola91
angusti-alatus44
angusticosta191
angustifolius152
angustiretis19, 48*, 50, 51*, 67
annotinus 112
apollinairei67
arachnoideus106
araucarieti35
arctocarpus13, 19, 20, 41, 52, 54*, 80, 96, 97
ssp. caldensis55
ssp. madecassus20, 55
var. arctocarpus52, 53*, 55, 80
var. caldensis 52, 55, 56*
arenaceum167, 170
arenicola 184, 186
areodictyon13, 20, 23, 39, 46, 47, 56, 57*, 58*, 61,
118
areodictyon59, 61
argyrocaulon13, 20, 40, 58, 59*, 60*, 61, 160, 161
arsenei 135
asperifolius20, 22, 41, 61*, 62, 63*, 118
atratus 162, 166, 176
auribrunneus41, 44
austro-subulatus138
bartlettii23, 167, 169
benedictii161
bermudianus184
beyrichianus145
bogotensis217
bolivarensis81
bolivianus140, 142
brachymitrius143
brachyphyllulus149
brachythysanos143
breweri56
brittonae172, 175
brotherianus176
brunneo-bolax42
bryotropii20, 39, 40, 63, 64*, 65*
cacti 143
cacuminis84
caldensis55
calymperidictyon143
232
Index of ScientificNames
capitulatus20, 39, 63, 66*, 67*, 182
caracassanus162
carassensis81
careiroanus172
carolinae 16, 19, 39, 40, 65*, 67, 68*, 105, 217
catarractilis19, 172
catumbensis179
cavernosus87
cavifolius 9, 20, 23, 40, 61, 67, 69*, 70*
cerradensis67
chilensis 109
chionophilus149
chrismarii135
var. suboblongus138
var. tolucensis137
chrysodictyon13, 148, 151
cinchonae 152
clavatus 5
cleefii 20, 40, 70, 71*, 72*
concolor 13, 20, 41, 71, 73*, 74*
controversus10, 19, 41, 72*, 75, 76*
costaricensis179
crispatus55
crispicoma52
cruegeri196
cryptopodioides20, 41, 76, 77*, 78*
cuatrecasii187
cubensis20, 41, 78*, 79*, 80
var. exaltatus 129
cucullatifolius84
cuspidatus20, 40, 63, 80, 176
var. cuspidatus80, 81*, 82*, 118
var. dicnemioides40, 80, 82*, 83*, 118
cygneus20, 40, 84, 85*, 86*, 87*
delicatulus50
densicoma20, 41, 87, 89*
var. densicoma88
var. yungarum88
dentato-acicularis143
destructilis135
detonsus 184
dichrostis20, 40, 90*, 91*
dicksoniae144
dicnemioides83, 84
discriminatus52
ditrichoides50
var. robustior50
divisus 144
donnellii 179
echerieri 120
edithae 20, 23, 39, 91*, 92*
ekmanii 84
elliottii 84
erectus152
erythrodontius145
exaltatus 129
exfimbriatus196
exustus200
falcatulus 52
fendleri 148
filicaudatus41
filicuspes88
filifolius 32, 80
233
var. humilis 35
var. longifolia36
fimbriatus99
flaccidus75
flagellaceus222
flexuosus 17, 18, 19, 24, 37, 41, 54, 93, 94*, 95*,
97, 113, 114, 196
var. flexuosus93, 97, 118
var. incacorralis41, 93, 96, 97*, 118
fragiliformis101
fragilis5, 18, 19, 24, 39, 58, 66, 98, 101, 102, 148
ssp. fragiliformis20, 98, 100*, 102*
ssp. fragilis20, 98*, 100*
friabilis 135
fulvus 47
fuscatus 144
galapagensis156
gardneri20, 40, 95*, 102, 103*
gastro-alaris20, 41, 104*, 105*
gemmatus20, 41, 106*, 107*
gertrudis88
giganteus 126
glauco-pallidus144
glaziovii 52
gracilicaulis65, 179
var. angustiretis50
griseus6, 20, 39, 109
ssp. griseus 108*, 109, 110*
ssp. ingeniensis 109, 110*, 111*
guatemalensis44
haitensis35
harpophyllus91
harrissii36, 79, 80
helleri 112
hellerianus112
heterophyllus93
heterostachys20, 39, 105*, 112*, 142
hoffmannii81
hondurensis93
huallagensis20, 22, 35, 40, 107*, 114
var. huallagensis114, 115*
var. weberbaueri22, 114, 115, 116*, 129
humifugus144
humilis 35
humoricola75
incacorralis18, 96
incertus20, 41, 116, 117*, 118*
incrassatus19, 172
ingeniensis110
insignis 139
insularis47
introflexus5, 6, 16, 22, 23, 125, 146, 156, 182
var. altecristatus156
itacolumitis 196
japonicus 19, 40, 120*, 121, 122*
jamaicensis 84
jamesonii 9, 20, 39, 118, 119*, 120*
joinvilleanus144
jugorum20, 40, 121, 123*
julaceus 5, 19, 20, 39, 44, 122, 124*, 125*, 126
ssp. arbogastii126
julicaulis 20, 39, 126, 127*, 128*
karstenii165
234
Flora Neotropica
kingii 148
krauseanus136
laevigatus163
laevis 211
lamellatus 156
lamellinervis19, 22, 41, 115, 116, 126, 128
var. exaltatus 126, 129, 130*
var. lamellinervis126, 130*, 131*
lamprodictyon150
lapidicola109
latinervis159, 161
laxiretis26
laxitextus66
laxoventralis109
leanus 224
leptodictyon50
leucobasis139
leucogaster200
leucognodes58, 59, 61
liebmannii 152
liliputanus207
longicellularis20, 40, 128*, 129, 132*
longisubulatus47
luetzelburgii101
luridus152
luteus 20, 41, 130, 133*, 134*
macrogaster75
macrophyllus33
malagensis222
marmellensis179
mexicanus93
microjulaceus123
microtheca52
minarum 180
mosenii 75
muelleri33, 163
multicapsularis139
multisulcatus126
nano-filifolius33
nematophyllus82
nigerrimus163, 166
nitidus56
nivalis 9, 13, 18, 23, 40, 58, 121, 134, 136*, 160,
177
var. multicapsularis40, 134, 139*, 140, 151
var. nivalis 134, 135*, 139, 140
oblongus20, 41, 140, 141*, 142*

occultus 5, 13, 15, 17, 20, 23, 39, 142*, 143*, 146
oerstedianus19, 23, 39, 145*, 146*, 147*
orthopelma75
ouro-pretensis75
paramoensis176
pauper5, 13, 41, 147*, 148, 182
var lamprodictyon40, 148, 150, 151
var. pauper148, 149*, 151
pelichucensis52
penicillatus126
var. exaltatus 129
percurvatus196
perexilis 20, 40, 150*, 151*
perfalcatus55
perpusillus39, 67
peruvianus215
pilifer 5, 6, 8, 9, 18, 19, 23, 39, 44, 146, 147, 148,

152, 154*, 155*, 158
ssp. pilifer 9, 152, 153*, 157
var. pilifer 152, 156
var. lamellatus9, 152, 156
ssp. galapagensis9, 152, 156, 157*, 158, 215
pilosissimus 152
pittieri 8, 16, 20, 40, 46, 61, 70, 118, 158*, 159*,
160
platyneuron144
pleurocarpus75
poasensis 136
polytrichoides156
porphyreocaulis113
33
porphyreodictyon
praealtus 176
procerus120
procerus176
proliferus152
propinquus148
144
pseudobrachymitrius
pseudodichrostis191
pseudodicranum126
pseudofilifolius33
ptychotheca161
puiggarii 163
purpureocaulis19
pusillus200
pyriformis145
ramuliger76
rectipes75
rectisetus52
rectinervis99
recurvifolius109
recurvipilus109
reflexisetus20, 41, 161, 162*, 163*
var. circinatus161
reflexus96
renneri159, 161
var. latelimbata159, 161
restingae144
richardii5, 11, 19, 23, 40, 162, 164*, 165*
rigidiusculus52
rigidus163
roellii 93
roraimae52
rosulatus148, 149
rubricaulis35
rufescens144
saint-pierrei121
sargii 93
savannarum 18, 19, 22, 23, 40, 166, 167*, 168*,
169*, 170
saxatilis 84
scabrellus144, 146
scabrophyllus109
schiffneri167
schimperi 138
schwaegrichenii184
scopelliformis55
sehnemii 19, 39, 170*, 171*, 172
sellowianus52
sellowii 52
setaceo-rigidus41, 44
setifolius 174
sharpii20, 40, 142, 171*, 172, 173*
shawii 10, 20, 41, 172, 173, 174*, 175*, 176
soirifolius196
sphagnicola101, 102
sprucei167, 170
spurio-concolor88
standleyi 118
stenopelma 19, 75
steyermarkii163
stramineolus75
straminifolius93
stricticaulis41
strictifolius53, 55
strictisetus35
strictulus26
strictus152
subannotinus139
subarctocarpus35
subarenicola184
subbrachymitrius144
subconcolor56, 58, 61
subcubitus55
subcuspidatus19, 40, 176
var. damazii 176
var. subcuspidatus176, 177*
subfalcatus196
subfimbriatus136
subfulvus130
subgriseus109
subincacorralis47
subincrassatus41
subjugorum20, 40, 93, 177, 178*, 179*
var. edithae91
subleucogaster180
suboblongus136
subpenicillatus126
subproliferus152
subreconditus145
subturfaceus99
subulatus152
surinamensis8, 10, 18, 19,41,65,179*, 180*, 181*,
182, 186
var. angustiretis50
tablasensis55
tallulensis20, 39, 114, 182, 183*, 184*
tener 187
tenuissimus33
tequendamensis182
terebrifolius196
thysanomitrioides71, 92
tijucae 145
tolucensis137
tortilipilus167, 170
tortilisetus144
tortuosus126
trachyblepharon9, 19, 22, 35, 40, 184*, 185*, 186
ssp. comatus 186
trichophorus62, 63
trichophylloides20, 39, 186, 187*, 188*
trivialis 20, 39, 187, 189*, 190*
trollii 150
235
tuerckheimii194
tunariensis99
uleanus8, 20, 41, 188*, 189, 191*
underwoodii172, 175
ventri-alaris52
verticillatus34
villicaulis185
viridatus20, 41, 190*, 191, 192*
weberbaueri114
weddelii62
widgrenii20, 41, 192, 193*, 194*
wurdackii114
yungarum89
yunqueanus163
zygodonticarpus5,20,40, 114,151,194*, 195*, 196
Campylostelium3
Chorisodontium200
Cynodontium3
Dicranaceae2, 3, 6, 21, 22
Dicranella2, 3, 9, 15, 22, 23, 105, 200, 203
subg. Anisothecium203
cerviculata3
curviseta200
fendleri 200
pilopogon200
sartorii200
Dicranelloideae3, 9
Dicranodontium2, 3, 4, 6, 9, 15, 17, 18, 20, 21, 25,
196, 197
asperifolium62
brasiliense197, 198*, 199*
denudatum20, 149, 197, 199*, 201*
flagellacea 222
fleischerianum199
longirostre196
meridionale20, 197, 198, 202*, 203*
pulchro-alare197, 199, 203*, 204*
pusillum26
schwabei200
setosum26, 200
subporodictyon200
uncinatum199
Dicranum4, 5, 9
subg.Paraleucobryum225
albicans228
alopecurum112
altissimum 118
alto-filifolium32
anderssonii47
angustirete50
araucarieti35
arctocarpum52
arenicola 184
areodictyon56
argyrocaulon59
auribrunneum41
brachymitrium143
brachythysanum143
brasiliense26
cacti 143
cacuminis84
calymperidictyon143
chionophilum197
236
Flora Neotropica
chrismari135
concolor71
controversum75
crispatum55
crispicoma52
cruegeri196
cuspidatum81
cygneum84
densicoma87
denudatum197
destructile135
detonsum184
dichrostis90
dicnemioides83
discriminatum52
divisum 144
donnellii 179
enerve227
erectum152
exaltatum 129
exile 144
fendleri 148
filicaudatum41
filifolium32
flagellaceum222
fragile 99
friabile 135
gardneri102
gastro-alare105
gemmatum 106
glaziovii 52
guadeloupense206
harrisii79
hellerianum112
heterostachys112
hoffmannii81
humifugum144
humoricola75
itacolumitis196
jamesonii 118
joinvilleanum144
julaceum 122
krauseanum136
laevigatum163
lamellinerve126
lapidicola109
laxobasis 180
leucognodes59
liebmannii 152
longisetum26
macrodon26
macrogaster75
macrophyllum32
microjulaceum123
muelleri33
nano-filifolium33
oerstedianum146
orthopelma75
pauper 148
penicillatum126
perexile 151
perfalcatum55
pittieri 26
platyneuron144
pleurocarpum75
porphyreocaule112
33
porphyreodictyon
praealtum 176
proliferum152
propinquum148
144
pseudobrachymitrium
pseudofilifolium33
rabenii35
rectisetum52
recurvipilum109
reflexisetum161
retinerve99
rigidiusculum52
rigidum163
rosulatum148
rufescens144
scabrophyllum109
scopelliforme55
sellowianum52
setaceo-rigidum41
spiripes27
spurio-concolor88
stricticaule41
strictisetum35
strictulum26
strigulosum27
subarctocarpum36
subconcolor56
subfalcatum196
subgriseum109
subincrassatum41
subleucogaster180
sublongisetum27
subpenicillatum126
terebrifolium196
tortuosum126
trachyblepharon184
uleanum 189
ventri-alare52
verticillatum34
villicaule185
viridatum191
widgrenii192
zygodonticarpum194
Didymodon
gracilis206
medius37
Ditrichaceae21
Ditrichum3, 21
bogotense 196
Grimmia 3, 4
Leucobryaceae9
Leucobryum9, 160, 228
megalophyllum228
Maireola3
Metzlera25
Metzlerella2, 25
brasiliensis26
flagellacea 222
longiseta26
Metzleria25
237
brasiliensis26
longiseta26
spiripes27
strigulosa27
sublongiseta27
Metzleriella25
Microcampylopus2, 3, 4, 5, 9, 10, 14, 16, 18, 22, 23,
24, 105, 200, 203
curvisetus7, 11, 14, 16, 19, 37, 200, 205*, 206*
khasianus200
laevigatus200
subnanus200
Microdus2, 9
Mitrobryum3
Paraleucobryoideae2, 3, 6, 8, 9, 12, 22, 23, 217
Paraleucobryum2, 3, 4, 5, 8, 11, 12, 14, 16, 18, 21,
160, 219, 225
albicans228
brasiliense229
densifolium159, 161
var. congestum160
var. latilimbatum160
enerve 18, 20, 23, 160, 226*, 227, 229
longifolium7, 16, 160
ssp. brasiliense18, 197, 227*, 228*
ssp. longifolium229
Phyllogonium19
Pilopogon2, 3, 4, 5, 6, 11, 12, 13, 14, 16, 17, 18, 20,
21, 22, 24, 203,213, 215
subg. Parapilopogon205
subg. Pilopogon206
subg. Pilopogonella205
subg. Thysanomitriopsis12
aemulans41
africanus14
atratus 162
bernoullii206
calycinus206
capitiflorus206
caracasanum162
glabrisetus206
gracilis 19, 20, 44, 156, 203, 206
var. bernoullii206, 211
var. comigera211
var. divaricata211
var. parvus211
var. pittieri211
guadeloupensis203, 205, 206, 207*, 208*, 211,213,
215
laevis 14, 20, 156, 205, 206, 210*, 211*, 213
liebmannii 152
liliputanus207
longefimbriatus207
longirostratus22, 204, 205, 212*, 213*
macrocarpus6, 11, 18, 20, 205,206,213, 214*, 215*
microcarpus207
nanus 211
peruvianus20, 156, 158, 204, 206, 211, 213, 215,
216*, 217*
piliferus211
puiggarii163
schilleri 18
subjulaceus207, 211
tiquipayae18, 20, 205, 206, 215*, 218*, 219*
Pilopogonella
laevis 211
Racomitrium4
Sphaerothecium3, 4, 10, 12, 14, 17, 21, 23, 24, 39,
216
comosum217
phascoideum20, 213*, 217, 220*
Sphagnum9
Syrrhopodon
leanus 224
Thysanomitrion2, 3, 13, 14
aemulans41
arenaceum167
caracasanum162
carassensis81
filifolium 32
flexuosum 93
griseum 109
jamaicense 152
luteum 130
macrophyllum32
miserum163
muelleri163
nigerrimum163
nivale 135
phascoideum217
puiggarii163
rigidum163
scabrisetum52
schiffneri167
yunqueanum163
Trichostomum
stenocarpum224
Weisia
nivalis 125

Flora Neotropica

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Flora Neotropica

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Organization for Flora Neotropica

Dicranaceae: Campylopodioideae, Paraleucobryoideae

Library of Congress Cataloging in Publication Data

' Departmentof Botany, University of Duisburg,Fachbereich6, Postfach 101629, 4100 Duisburg,Federal

conversely, display extreme differentiationof

the presence of alar cells is regardedtoday as

determined by modification, since in all these

FIG. 1. Transverse sections of leaves in different genera of Campylopodioideae and Paraleucobryoideae. A.

b. the change from dorsal groups of stereids

("socii, comites, Begleiter")and function for the

FIG. 2. Transversesectionsof differenttypesof

pend on such early stages of development. The

FIG. 4. Transversesections of stems in differentgeneraof Campylopodioideaeand Paraleucobryoideae.

vapor can pass through them and be absorbed

thencurvesdownwards.In this mannerthe young

shape of the seta causes wide movements of the

as in the common dicranoidtype. In addition to

in the pottiaceous type of peristome). In both

as the most primitive species) have the normal

The spore ornamentation is not yet known by

'i~J ,' 'tf.'/X

SEM pictures oflpenstomes. A, B. Bryvohumbertiafilifolia (Frahm 1555, hb. Fralm). C. Campylopus

revision of the genera (Giese & Frahm, 1985a,

12 arefound in differentpartsof the range.Chromosomenumbersin Campylopusareknownfrom

FIG. 8. Spores ofCampylopodioideae. A Atractylocarpusl

A. Campylopusflexuosus (Hakeliers.n., hb.

triterpenoidhop-22(29)-ene(Huneck, 1983), but

respirationwhich is especially critical for ecological conditions of the tropics.

ferenceis mainly due to differentstructuraland

America. The neotropicalspecies are separated

Disjunctions between Central America and

humbertiafilifolia and Campylopusarctocarpus

Andes from the Equator north and south the

ORIGIN AND EVOLUTION

did not spreadsubstantiallyinto the tropics. In

cobryoideae and Campylopodioideae(as most

The anatomies of Paraleucobryoideaeand

Sexual reproductionplays a differentrole in

capsules.In Brotherathereare numerouscollections of fertile material from its Asian range.

The total lack or rareness of sporophytesin

almost blackish,in loose tufts.Stemserect,rarely

Plants small, from a few mm to 15 cm high,

Key to the Generaof Campylopodioideae

without experience.It is generallynot possible

ArtificialKey for Sterile Campylopodioideae

ric, cylindric,without stomata.Annulusnot differentiated.Operculumlongly rostrate,as long

Autoicousor dioicous.Plants slenderin compact tufts, golden-green,glossy, rarelybrownish

Specimens examined. BRAZIL. Rio DE JANEIRO:

The Liitzelburg specimen is sterile and can be

1. Atractylocarpusbrasiliensis(C. Miller) Williams, Bryologist 31: 110. 1928. Dicranum

Brotherus, Bot. Jahrb. Syst.

Atractylocarpusstrictulus(C. Miiller)J.-P. Frahm, J.

FIG. 10. Atractylocarpus

Belg.31:146. 1893.Type. CostaRica. Foretdu Rancho Flores,Pittier9907 (holotype, PC).

Costa Rica. Forets du Barba, Tonduz 5510 (holotype, PC).

Distribution of Atractylocarpus brasiliensis (O) and A. nanus (0).

one half of the leaf base, excurrent,in transverse

Peristome teeth divided up to 2/3into two prongs.

Monte, Pringle 10603 (NY).

Uyuca, Standley & Molina 4307 (F), 4944 (BM). EL

S of Cartago,Bowers837 (FLAS);Sierrade Salamanca

Prepared by Hendrk R. Rypkema

Distribution of Atractylocarpus longisetus.

as the capsule.Peristometeethsplitto base.Spores