CHAPTER 9

Natural fnemies
James C. Cokendolpher and Plamen G. Mitov

lthough this chapter is titled ''Natural Enemies." it also covers some assnl'i<l -

A tions that arc neutral or potentia lly bcncHcialto Opilioncs. Predation. r; mni-
balism. disease-causing pathogens. and many parasites are dl!ilrly dltrimcntal
to dead or dying Opilioncs. It appears that the imparl of gregarin e p;~ra s i! cs and
phoretic riders is not life threatenin g. but they rnay reduce litn~.:ss. l.ikc wisl'. biiCh.ria .
seeds. and spores that me cmricd by Opilioncs (thro ugh no S[>ediil dTons of the
transmi lled) appear to be neutral except for a reduction in lit ness si mply by thl'l'Xtra
load the anima ls have to carry or waste energy trying to remove. Only inth1. raSl'S of
some miles. cyanobacteria. algae. <md li\'Crwons arc mutualistic rcl<llionshi ps sug
gcsted. In these cases the individuals might re<_.eive some assista nce from till' mit{'S as
they apparently groom the host. or the hosts might <~equ irL' ill'illlHIUIIiiJ.!C from till'
green coloring of the chloroplasts.
We know lill ie about the associations of Opiliones. Al though thlrc art ;q>prnxi-
matcly 6.000 described species in the nrdcr. there are l't11'L'I' than 1()() rcpnrtcd
species of pathogens and pt1rasiles. on ly slightly over 300 species of prLtl;Jtors. <llld
only about two dozen phoretic/endozok/cpizok assodHtions recorded workltl'idt
Part of th e prob lem is that the simple obscrV<llion ;md reporting of sud1 ti SStll'htlitHl S
are not considered of high scholarly vcdue. <IIHI therdnrt m;~n y l'l'l'lllS go llll -
recordcd. Furthermore. most p;~thol og i s ts illHl pantsitologist s canlHit id Lntil'y Opil -
iones even to order. and harvestml'll rl'Se t~rchcr s likcwisl' ran1wt ).!.l' lll'r; dly idLntil~
predators. parasites. or pathogens cxrept in \'cry general terms. Tlw llltlllhlr ul' assu-
ciation s is also underreporte{\ berause nr the dirticuhy in ohsLrving tlll'Sl' <H'Iil'itil'S.
Most harvestmen arc mxturn ttl or cryptic. and thertrure tlll'ir s tru ~g l t ICw lik ;..tn~s
essentially undetected by humans.
The beginnin g poin ts rnr the study or harwsl tllilll l'llCillkS <l l"t'l\\'ll d;~ ss k s tha t
ca me out in the same year (Bristowc. l lJ-!.lJ: Sankey. 19-!.lJbl. Sinn thai linll' nu -
merou s reports have lx:cn publ ished . hut nuuplhlted reviews tl pJX'i ll't'll until thl' mid-
1990s (Cokcndol pher. llJlJ 1: Mi tov. l9lJ'51. In Ihe IC 1IIowin~ disntssi1111S .~ rH.ri l il til' -
340 Natural Enem1es

tail ~ willlw hrid. l.i~t.s 1'1'ablc.s Y. I - Y. ~~arc supplied that detail species records and
tWrlilll'llllitewturt

PARASITES AND PARASITOIDS

l'<~n1 silism is <1 co mmon w;~y ol" Iiii... si nce owr hall" tlw known animals of the planet
<1 rc par;J.sitcs li\ lcdis & Hcnaud. 20011. His no surprise. then. t hat harvestmen arc
<1\Wcktd by lltlllH.'rnus species of par:1sites :md purasitoids (Figure 9 . .1 ). Few para-
sit()logists ;Jntr:lined as urachnologists. ;md thtrcforc many hosts arc not identified
btyond rlass or order in tlw litcmture. Sonll' records of ha rvestmen arc lost in the
more genenJIIists as "f\r<K"hni<l<l. whereas some records for long-legged spiders arc
misidclllilil'd <I S Opilioncs (<..'okcndolphcr. 1Y9 l). Similarly. arachnologists who dis-
cowr parasites often do not identify the agent beyond grega rine"' or nematode."
Cenerc1lly. thL pilr<ISite. if rl'taincd. is improperly preserved (wrong fixative or sta ge of
dewl,lpllll'llt J li1r furthLr idtntilkiltiOJl (for instructions sec Ch:1pter 1 5). Because of
till' h1l'k or gomltaxonumic characters in some groups (e.g.. Microsporida and juve-
niles of i\lcrmithidae). rolkctive groups llilVC been named and are used in Tabl e 9. J .
Sud1 groups or genen1 ol"tcn include species !hat probably arc no! related. This grou p
name is used .simply for "taxonomk conven ience" and includes species not readily
placed in known genera (possibly because a particular life stage is unknown ) and
spel'ics itt!"l'l"lm' scdis.
t\l lhough most purasites arc not fatal in Opilioncs. a ll the nematode parasites
tmd insect parasitoids will kill the harvestman host. In the few cases of pompilid
wusp.s t~llacking Opilioncs. it is not certain if it was parasitis m. predation. or mis-
taktn identity. The wasps were observed attacking harvestmen. but observations of
feeding or nest provision ing were not recorded {Evans. 1948). Most adult pompilids
feed almost cxdu.sivcly on nectar sources (Evans. 19 5 3). Only a l'cw species have also
bt'Cn observed feeding on hemolymph from C<l ptured spiders. but not harvestmen .
Anot her general characteristic of !his group is that all known species feed their
l<lrV<IC exclusively with spiders (Evnns. J 953). This suggests that wasps might mis-
takenly attack harvest men as a spider and later reject the prey. This still might result
in the death of the harvestmen. but such att acks would be very Infrequent.
The two most commonly reported parasites arc the endoparasilic gregarines and
the ectoparasitic mites (Table 9. 1 ). Both of these are large. easily observed parasites.
and for that reason they arc probably more often detected than the smal lcrendopar-
asitic species. The frequency of occurrence of both appears to Increase with the age
ami size of the harvestman host. It is not clear why this increase occurs. Perhaps it is
related to a longer period of exposu re (including the probability of reinfesting one-
self) or the higher biomass ava il ability. or both. Like many topics related to har-
vestman biology. !his is an area that needs detailed study.
In the case of !he gregari nes. adult parasites (Figures 9.1A.B) pass gametocytes
in the harvestmen's feces (Figure 9.1C), which dehisce and release many thousands
of infective oocysts. The oocysts arc ingested by harvestmen and start the cycle over
Figure 9.1. Parasites of Opiliones. (A-C) Undetermined gregarine parasite from the sclerosomatid harvestman
Leiobunum nigripes. (A) Ventral view of dissected harvestman showing large trophozoites/sporonts in the abdomen
(sternites are removed). (B) Two gametocysts in feces of harvestman. (C) Closeup of trophozoite (photos: j. C
Cokendolpher). (D) Immature mermithid parasite emerging from the abdomen of the phalangud Phalangium opmo
(photo: G. Poinar). (E) The nemastomatid Paranemastoma radewi with transparent. jellylike masses flow ing from
the mouth and anus: masses active with variety of saprophage nematodes (photo: P. G. M1tov). (F) Corpse of
Phalangium opi/io (Phalangiidae) that was killed by the nematode Steinemema carpocapsae. Note developmg
nematodes adjacent to the opened body (photo: G. Poinar). (G) leiobwwm politttm (Sderosomatidae) wit h
immature, parasitic leptus mite attached to the abdomen (photo: j.C. Cokendolpher). (H) Adult , free-liv1ng leptus
mite that feeds uj)On the sclerosomatid Leiobunum rownsendi as an immature (photo: j. C. Cokendolpher)
342 Nilttllcll Enem1es
T<~IJ i e 9.1 Paras1tes Cllld parasitoids of Opiliones
Parasite/Parasitoid
PHYLUM MICROSPORA
CLASS MICROSPOREA,
ORDER MICROSPORIOA
M1crosponclwm weisen
PHYLUM APICOMPLEXA:
CLASS SPOROZOAS IDA
Undetennnled greganne
(Frgures9.1A-cJ
CLASS SPOROZOASIDA,
ORDER EUGREGARINORIDA
Actinocepha/us megabuni
Allt/Jorhynchus longispora
Anthorhynclws sophiae
Arachnocystis arachnoidea
Contospora opalniae
cosmetophilus vonones
Doliospora repelini
Doliospora troguli
Echinoocysta phalangii
Sciadiophora sp.
Sciadiophora caudata
Sciadiophora c/aviformis
ScladiophOra fissidens
SciadiophOra gagrel/ula
Sciadiophora geronowitschi
Opiliones host Source
-
Opilio parietirws Cokendolpher. 1993
carinostoma omatum. Dicrano/asma scabrum, Cokendolpher. 1993, pers. obs.;
Equitius doriae. Graecophalangium atticum. Lacinius Gruber, 1993; Mitov. 1993,
dentiger, L. ephippiatus. L. horridus. L. insularis , 2000a,b, 2003, 2004, pers. obs.;
Leiobunwn globosum. L manubriatum, L. nigripes, Mitov & Stoyanov, 2004
L. politum, L. rumelicum. Lophopilio palpinalis,
Metaplatybunus grandissimus. Mitopus mario,
Odie/Ius lendli, 0. pictus. Opilio dinaricvs. 0. ruzickai,
0. saxatilis. Paranemastoma aurigerum ryla. P. radewi,
Phalangium opilio, Pyza bOsnica, Rilaena balcanica.
R. cf. serbica, R. serbica, Trogulus tricarinatus.
zachaeus anatolicus. and z. crista
Megabunus diadema Cokendolpher, 1993
Leiobunum rotundum. Mitopus mario. Opilio parietinus. COkelldOiptler. 1993
and Platybunus bucephalus
Lacinius ephippiatus, Leiobunum b/ackwalli, L. rotundum. Cokendolpher, 1993
Mitopus moria. 0/igolophus tridens, Phalangium opi!io,
and Ri/aena triangularis
Oppalnia sp Cokendolpher, 1993
Oppalnia sp Cokendolpher, 1993
vonones say; Cokendolpher, 1993
Leiobunum rotundum, Megabunus diadema. Mitopus Cokendolpher. 1993
mario, Oligolophus tridens. Opilio parietinus.
Phalangium opilio, and Platybunus bucepha/us
Trogulus tricarinatus Cokendolpher. 1993
Oppalnia sp Cokendolpher, 1993
Leuronychus pacificus 1
Clopton, 2003
Phalangiidae, Mitopus moria, Odie/Ius spinosus, Cokendolpher. 1993
and Phalangium opilio
Mitopus sp Cokendolpher. 1993
Phalangiidae. Lophopilio palpinalis, Phalangium Cokendolpher, 1993
~crassum.~ and P. opilio
Gagrellula saddlana Cokendolpher, 1993
Phalangium opilio Cokendolpher, 1993

SciadiophOia phalangii
PHYLUM PLATYHELMINTHES:
CLASS CESTODA, ORDER
CYCLOPHYLLIDEA
Pseudhymeno/epis redonica
CLASS TREMATODA, ORDER
DIGENEA
Brachylecithum sp
Unidentified larval fluke
PHYLUM NEMATODA
unspecified nematodes
CLASS SECERNENTlA,
ORDER RHABDITIDA
Heterorhabditis bacteriOphOra
HRhatxlitidHnematooes
Steinemema carpocapsae
(Figure9.1F)
CLASS ADENOPHOREA,
ORDER MERMITHIOA
Undetermined/Agamomermis
(Figure 9.1D)
Agamomermis phalangli
Agamomermis truncatula
Agamomermis HincertaH
Natural Enemies 343
Oplllones hOst source
Lacinius dentiger, Leiobunum rotundum. Mitopus Cokendolpher, 1993
mario. Opilio parietinus. Phalangium opilio,
Pha/angium sp., Pfatybunus bucepha/us,
P. pinetorum. and Rilaena triangularis
Phalangium opilio Cokendolpher, 1993
Phalangium opilio Cokendolpl1er, 1993
Phalangium opi/io Cokendolpher. 1993
Synthetonychiidae and/or Triaenonychidae, Rilaena Cokendolpher, 1993; Karaman.
triangularis, and Siro minutus 1993
Phafangium opilio Cokendolpher, 1993
Lacinius horridus. Paranemastoma radewi, and Cokendolpher, 1993; Mitov. 2000b
Phalangium opi/io
Phalangium opilio Cokendolpher. 1993
Phalangiidae, Gonyleptes fragilis. Lacinius denriger. Cokendolpher, 1993; Mitov. 2000b;
L. ephippiatus, L. horridus. Leiobunum globosum. Poinar et al., 2000
L. manubriatum. Lophopilio palpinalis, Mitopus mario,
Opilio parietinus, Paecilaemana quadripunctata,
Paranemastoma radewi, Phalangium opi/io,
Protoloptws sp., Rilaena balcanica, R. cf. serbica.
Thrasychirus sp., Togwoteeus biceps,
and zachaeus crista
Phalangium opilio Cokendolpher. 1993
Opilio sp. and Phalangium opilio Cokendolpller 1993; Poinar et al.
2000
Mitopus mario Cokendolpher, 1993
HoO/IIIJ/Uoo/ 1
Table 9.1 contmued
Parasite/Parasitoid
PHYLUM ARTHROPODA
CLASS ARACHNIDA.
ORDER ACARI
Allotl!rombtwn clumaanense.
A. fultgmoS!/111, and
AI/Otllrombivm sp
A//othrombl!l/11 neapolnum
C/Jarletonia englwffi
C/Jarletoma sourhcottl
Erythrae1dae
Leptus beroni
Leprus btcristatus, L jocquei.
L.polyt/Jrix,and
L.puylaerti
Leptus gagrel/ae
Lepwsgyas
Leptus hidakai
Leptus ignows
Leptus indianensis
Leptus kataallvs
Leptus lomani
Leptus nearcticus
Leptus oudemansi
Leptus Hpha/angii"
Leptus phuketicus
Leptus stieglmayri
Leptus spp. (Figures 9.1G,H)
Oplliones host Source
Oplliones Raykov & RimskiyKorsakov, 1948;
Cokendolpher. 1993
P/Jalangium sp. and zachaeus crista Cokendolpher, 1993
Bvnochel1s canariana Cokendolpher, 1993
Metagagrella tenuipes Cokendolpher, 1993
Leiobunum b/ackwalli, L. rotundum, Mitopus morio, Gabrys, 1991; Cokendolpher. 1993
Ne/ima si/vatica, Oligolophus hanseni, a. tridens,
Opilio parietinus. Paroligolophus agrestis, Phatangium
opilio, and Rilaena triangularis
Lophopilio palpinalis. Megabunus diadema. Mitopus Fain, 1991; Cokendolpher, 1993;
moria. OligoJophus hanseni. Opilio canestrinii, Fain & D'Amico, 1997
0 ruzickai, Opilio sp., Phatangium opilio, and
Rilaena triangularis
Cristina lettowi Cokendolpher, 1993
Gagrella sp Cokendolpher, 1993
Gyas titanus Fain & D'Amico, 1997
Opilio pentaspinu/atus Cokenclolpher, 1993
Opiliones and Phalangium opilio Haitlinger. 1990; Fain & D'Amico,
1997
Leiobunum a/drichi, L calcar, L. formosum, Cokendolpher, 1993; McAloon &
L nigripes. Leiobunum sp., L. speciosum, Durden, 2000
and L ventricosum
Mitopus mario Cokendolpher, 1993
5adocus funestus Cokendolpher, 1993
Leiobunum a/drichi, L nigripes, and L vittatum Cokendolpher, 1993
Cynorta sp Cokendolpher, 1993
Mitopus mario, Oligo/ophus tridens, Dahl et al., 1935; Stadler &
and Platybunus bucephatus Schenkel, 1940; pfeifer, 19S6;
Gabrys,1991
Gagrellula niveata and Gagrel/ula sp southcott, 1994
Opiliones Cokendolpher, 1993
Cynona roeweri, Dicranotasma scabrum, Egaenus MacKay et al., 1992:
convexus. Eumesosoma roeweri, Eumesosoma? Cokendolpher, 1993, pers. obs.;
sp., Eurybunus sp., Krusa sp., Lacinius ephippiatus. Gruber, 1993; Guffey, 1998;
Leiobunum aldrichi, L. aft. depressum, L. flavum, Prieto nueba & Bauza Ferre,
L. monranum montanum, L. nigripes, L rotundum, 1999; Toft, 2004;
 Natural Enemies 345

Oplllones host source

L. politum, Leiobunum spp., L townsendi, L vittatum, 0. Palmer. pers. obs.

Lophopilio palpinalis, Marthana nigerrima. Mitopus
moria, Odie/Ius picrus, 0/igolophus tridens, Opilio
parietinus, o. ruzickai, Paroligolophus agresris,
Pha/angium opilio, Phalangium sp., P/atybunus
bucephalus, Proto/ophus singu/aris, Rilaena
triangulares, Togwoteeus biceps, and Trachyrhinus
marmoratus
New genus near Leptus AcanthOprocta pustulata, Eubalta meridiana/is, Cokendolpher, 1993
Metagyndes pu/chella, Thrasychirus dentichelis.
and T. modesrus
Trombiculidae Gagrella alba, G. curvispina. G. formosae, oankittipakul & sonthichai, 2002
G. sexmacu/ata, and Metamarthana Fusca
Trombidium hungarium Egaenus convexus Cokendolpher, 1993
Trombidium sp. Ne/imasp Cokendolpher, 1993
undetermined mites Synthetonychiidae and/or Triaenonychidae. Equitius Acosta eta!., 1993; Cokendolpher,
doriae, Gagrel/opsis nodulifera. Gtobipes sp., 1993; Gnaspini, 1996; Mitov,
Goniosoma spp., G. spelaeum, Lacinius denriger. 2000b
L. ephippiatus. L. horridus, Leiobunum rownsendi.
L ventrieosum, LOphopilio pa/pinalis, Mitopus mario.
Nelima paessleri, Odie/Ius /endli, Opi/io dinaricus.
0. ruzickai, Pachyloidel/us goliath, Paranemastoma
aurigerum ryla, P. radewi, Phalangium opilio,
Prionostemma panama. Pyza bosnica. Rilaena cf.
serbica. Trachyrhinus rectipalpus. vonones sayi,
Zachaeus anatolicus, Z. crista, and Z. hebraicus

CLASS INSECTA, ORDER

DIPTERA
Ceratopogonidae Gagrellu/a ferruginea and Nelima nigricoxa Cokendolpher, 1993
cyclorrhapha larvae Discocyrtus invalidus Cokendolpher, 1993
Lasiohe/ea opilionivora Sclerosomatidae Lane,1947
Phoridae lporangaia pustulosa G. Machado. pers. cornrn
Phoridae? Goniosoma spp. and Goniosoma spelaeum Gnaspin i, 1996
undetermined larvae Acutisoma longipes Mac11ado et al., 2000

CLASS INSECTA, ORDER

HYMENOPTERA
Anoplius marginarus Odie/Ius pictus Cokendolpher, 1993
Chalcididae Synthetonychiidae and/or Triaenonychidae Cokendotptter, 1993

Original source citations not li'Sted hefe are found in Cokenclolpher (1993)
1. This host genus does not occur in the region reported. There is no doubt that 11 IS m1slflent1hed
346 Naun al Ene1mes

<~)!<~in. 11 is likdy lh<!l h;~rwsllmn rcinli.rtthcmsdvcs by ddcntting <ll or ncar areas

1h:~1 the.\' fnqut'll l for drinkint.:. L'illinl-!. or resting. ;~/though this hus not actua lly
httn uhsl'rwd. 'l'hl' uorysts <ll'l' s nwllt'IHHI).!.h that lwrvcstmcn could pick them up
ll'htn 11111\'ill)! throu~h till' l'IH'irtnlllll'llt ;uul ingest them during grooming. Like
ll'i:-.L'. 1hey nH1Id lw tspuMd to the oocysts if thl'y stop to inspcd feces of other har
w stllll'll. (lilt' crt' us 1/. C. l'crktmhllphcr ) unsurressfully <Jtttmpted to infer! some lnJr-
wst Jmn by pl<~l'ing actin oorysts in tht Wtller ol'fered to captive harvest men
, \l thrnl)!h SOilll' authors tTsurusaki. I YSh: 1\ litov. 20lltlb) have recorded the number
of J,.:rtgarirws in 1\'ild populations mw time. till' lark of good characters to identify
tilt' grtgarims rnlurnlthe anwunt of inli.11nwtion that wu ld be obtained from these
reports. It is not kno\\'ll if the..;e <1ft' ;dlnH:rn!wrs of <1 si ngle species or c han ges in
rlLIIllbl'rs crf 11\'c1 c1r llltli'L' sptrics cJI't'r tillll.'. Thtlark of good idcnti lication characters
;dso keeps us frnrn knu\\'ill )! tht t'Xttnt of {\istribution of these parasites gcographi
l'<JIIy. <IS \\'ell <J S thei r host spet'its. Sumt of the records in the literature arc su spect
htr<Jusc not allthtlifc SI<Jgcs of the pan1sitts arc known tmd it is likely that more
tlwn one spct'ics l'ould be lumptd under a single pan1site's name.
l\1ostnf 1he harwstrn<Jil spLcies found Oll i\ lt. Vitosha (Bulgaria) were highly par-
asitized by greg<J rints in the <m tulllll (~ \it ov. 20t)(Jb ). Ts urusaki ( 1986) found si m-
ihlr infesl<ltion s in IAio/mmun spp. (Sckrosomatidae) in Japan. Some specimens of
tht plwhmgiids Hihrcmr d'. .\'t'l'binr. /,a!'illius /rorridm. and MiiOp iiS moria from MI. Vi-
tosiHI wert p<~rasitizcd by both greg<~rincs and mermithids. The mermilhids were
fou nd during t\pril to Nowmbcr. mostly on the open grassy habitats, which arc in
con \art wi th Willer. Th( placement ncar wattr probably fa cilitates the parasitism by
the tntnnithids. 1\lermithids can emerge from thei r host into damp situations. where
thty mt~turc <llld deposit eggs.
The lik cycle of merrnithids that attack lwncstmen is unknown. It is possible
that most. if not all. han: an indirL'Ctlift cydc like that recorded in spiders (Poin ar
& E;:~rly. 1990) . In this tnsc the infective egg of th e mcrmithid would be ingested
by a parc1tcnk hostthcll would later be C<ltcn by a harves tman. The parasite would
be ingested ;dong with the paratcnic host. In th e known case of spiders, the
para tcnic hosts arc cmldis flies and mayllics-itcms that have a lready been
rc(ordtd in the die t of harvestmen (sec Chapter 8). Mcrmithids with a direct life
q dc enter their hosts via direct penetra ti on of the cu ti cle. The indirect type of de-
ve lopment would bypass cutil'le pen etration a nd would a llow the paras ite access
to the host hcmowcl.
The nematode parasites ;:1rc split between the mermithids that presumably kill
thl'ir host upon emergin g th ro u gh the cuticle to molt to maturity (Figure 9.10) and
thl' rhabditid nematodes (Figure 9. 1 E) that Inject their host with pathogenic bacteria
that kill the harvestman. Other rhabditid ne matodes are saprophagic and can be
seen on harvestman eggs (Mitov. 2000b). as well as on recently deceased individuals
(Figure 9.1 F). These latter nematodes presumably do not harm the eggs.
The literature data concerning other protozoans and helminthes parasitizing
harvestmen a rc very scarce (Table 9.1 ). Only a s ingle bloodborne mic rosporidium
 Natural Enemies 347

(Microsporidium weiseri) has been observed (Silhav:Y. 1960) In Opilioncs. and little
is known about lt. Harvestmen are recorded to be intermediate hosts of a few
Platyhelminthes. but in no case are they the primary host of these flukes or tape-

Mites known to be parasi tic on harvestmen belong to the families Trombiculidae.

Thrombidiidae. and Erythraeidae. Only the larval forms arc parasitic {Figu re 9.1G).
whereas the nymphs and adu lts a re predaceous on sma ll insects. Because the larval
and postlarval stages of these three families are heteromorphic (do not resemble
each other; see Figures 9.1G.H). systematists have long used different scientific
names for each (Southcott. 1991 ). Only after the larval and post larval stages arc as-
sociated by rearing can any meaningful classifications be constructed.
Erythraeid mites deposit a cone of cementing material at the auachment site
that. along with their inserted chelicerae. form s a tight anchomge (Abro. 19!H!). The
attached larva can then suck hemolymph <Hld tissue fluid s over a long period of
time. Parasitic mites appear to luwe a preference for certain regions of the body and
legs of harvestmen (McAloon & Durden. 2000; Mitov. 2000b). It is evident that most
of the mites are a llached to the leg femora. pat ella and tibiae. orularium. and dors<l l
metasoma. The coxae. trochanters. metatarsi. l<lrsi. chelicerae. and ventral meta-
soma bear the fewest larvae. None were found on the pedipalps. The lallt'r fact. as
well as the sparse coverage of metatarsi and tarsi. may be due to tht walking mech-
anism of harvestmen (Kastner. I93la ). or to the possibility that these parts arc
easily reached and readily cle<med by the animals themselves. Ultrastru ctural
studies by McAloon and Durden (2000) a lso suggest that this tl!lachmcn t-sitc sclcc;
lion could be due to integumentary morph ology. Ahro ( 19XH) stated that no vitJlent
defense on the part of the harvestman had been recorded. Observation of living htlr-
vestmen with mites revealed no rubbing or twit ching that would indicatt that the
miles were C<lusi ng any distress (J. C. Cokendolphcr. p(:rs. ohs.). A relati vt.ly small
number of harvestmen in a population will be ptlrasitized by most of tlw mitt'S. This
frequency distribution was observed by rl!l ct\loon ~md Durden ( 21llllll inthtl iSA <md
by Mitov (2000b) in Bulgaria. Multiple attacht.(t mites per h;~n es tman \\"t'rt.not un-
com mon: 709 miles on 500 harvestmen in the lJ$,\ <ltld HI 3 milt'S on 3 3lJ har-
vestmen in Bulgaria. tn scvcr<Jl cHses the ln<lll uf par;1sites per harwstman \\'tiS ex-
traordinary. Ten to 14 mites per /,dohiiiiWHjiJJ"IIIOS/1!11 were recorded on Y out of 500
infested harvestmen in the USi\ study. i\mung the most IH'tl\'i ly aiTectcd sped mtns
in Bulgaria were the phalangiids M. 111orio. with 2 I (only on kgsl. and i'.nduwus
crislll. with the highest number ( 31) of mi le larv<1c found on a si nglt host spedmt.n
(on legs and body). It is uncertain if this n:sults from <I potential hosl hl'ing expostd
to a l;1rge number of host-seeking larvae rteently hatchtd from till egg Ill< ISS or if tl
previously parHsilized harvestman is physically or behmiorodly 1nort' susnptihle to
multiple attacks.
Flies of the family Ceral opogonidac <Jre blood feeders. Thtst tlil.'s hmt occasion-
ally been seen feeding on lu1rvcstmen. This is an t1ssmit1tinn 1lw1 is pruht1hl.v tl\"cr-
looke<l by most observers becau se of the small size of the flit'S. l'horid<ll' Ilk.~ h<~w
been ~L'L'II a tl<Kk ing lkiizili<~n Opiliones !C n<t spini. llJY6 : G. Mac hado. pers. m mm .).
\!em her ~ 11f 1hl' r:~ mily I'IH!rid<n: :~re dinrse in their feedin g habit s. but some species
<lfL' knl!\\'11 111 IK par<~ s itoid s . These puntsitoids lcty an egg on the body of the host.
11 hich h<lldlL' ~ w1d dl'\'elops in terrwl ly in th e host lOr sewra l in stars.

PATHOGEN S

I'Jwre <~re relt~til dy fell' p;nhogens knO\\'n from Opiliones. and most of these are
recunblonly in 11nt: ur limited numbers of incidents IFigure 9.2). The majority of
recurd:-. <~rc ur p;nhogenic fungi - in nwny cases attacking eggs (Goodnight & Good-
night. l i.J7h: .\lonL li.J H/. 1<-.JI.JO: R;unircs & Giarella. 1994: Machado & Oliveira.
l ':J':JH. 21HJ2: Elpinu-C.unpus el <tl .. 21HJ I: .\I<Khado. 2002: Figures 9 .2C.D ). From
1he d<tSS J lyphumyce\l'S. there ilfl' records of H!JIIICIIOSli lllr l'ermcosn in fect ing pha-
l<~ngiitb 1.\ !;tins. llJ51J: Le<llhenla le. I Y70J. AkWrr/1i:iru11 m1isopline in Meynlopsalis
rumir/ol.\leyer- l{ochu\\' & Liddle. llJH X) . .\ 'onwrata tll!Jpico/a in Ll'iobunwn l'illnlllm
l( ;fl'l'll Stone et ;d.. l lJHX I. ;md l:ll.tJ!JOdoml!iwn amntarum infecting several species of
Opiliones ICkca rune & Campadelli. 19'JS). Gonyleptids and manaosbiids a re at-
tacked by pathogenic fung i of the class Pyrenumycetes that may cause the death of
th e individu;t ls (~vi O II c r. 191! I: Koval. IIJ 74: Figure 9.2B). Equally let hal is the zy-
gtunycete t>omlom plwhmyirida. which <II tacks individuals of Plwlm1giwn opilio (Lager-
heim. 1H'Jtl: Elli s. JI.J56: Le<ttherdalc. 195H. 19 70).
One species or fun gus. though. 1:'111omoplwya lmtkoi. is known to cause consid-
erable numbers or dea th s in European harvestmen . An epiwot ic (temporary in-
crease in the incidence of infection s) was observed during late summer by Batazy
l l9 7XJ. Keller I I YX 7) reported that this species of fungus was rather common from
l<ll e Ju ly to the middle of Sept ember and often ca used epizootics along the borders of
rorests and hedges. An infected harvestman will crawl up in the vegetation and die
while hanging onto a blade or grass or ot her plant part (Figure 9.2A). Similar be-
havior has been noted in some insect s. such as nics. grasshoppers, wasps, and cater-
pillars. which become lixed to an aboveground substrate when infected by other en-
topathogenic fungi /Keller, 1987)
No doubt pathogen s arc more numerous and widespread. Because most opilio-
nologists do not recognize diseased harvestmen. it is likely that most cases go unde-
tected. Only two species of pathogenic bacteria (Xenor/wbdus luminesce/Is and X. ne
matop/1ilus) have been recorded for harvestmen. both attacking the pha langiid
Plwhmyiwn opilio /Puinar & Thomas. 1985 ). Additionally. no rickettsial or viral dis-
ease has been diagnosed in harvestmen (Morel. 1978), but this may be more a case
of no one looking than an absence. Viral and rickeltsial diseases have been shown to
occur in as high as 5%-- 10% of a natural population in nonacarine arachnids
studied (Legendre & Morel. 1980: J. Haupt. 2000). Cory et al. (1988) recovered ac~
live Pano/is nuclear polyhcdrosis virus from the feces of harvestmen, but no disease
in the harvestma n was noted.
 Natural Enemies 34 9

Figure 9.2. Pathogens of Opiliones: {A) Diseased Oligolophus /fldell5 {Phalang1idae) hangu\g from a gra$S
blade. Harvestmen infected with Entomophaga bat(oi fungi clasp the upper pans of plants whe1e they d1e
(mod ified from Kell er. 1987). Sca le bar= 5 mm. (B) Corpse of a manaosbiid (?)harvestman from G\1Yill1,l
covered by the pathogenic fungus Torrubiel/a gonylepticida (photo:]. Spatafora). Scale b,u " 5 mm (C)
Corpse of the gonyleptid harvestman Acutisoma diSColor covered by an un,den!lfied fungus (phow G
Machado). Saprophytic and pathogemc fung1 are dtfftcult to separate by s1ght alone Scale bN = 10 mm. (Dl
Egg batch of the gonylept1d harvestman Acut,soma proxmwm att<lcked by fung i (photo BA Buz,ltto)
Sca le bar = 10mm
3 50 N<lt lll <l l Enem1es

PREDATORS

l'rldation of h;~ntstmcn is nunpkx in that the <Jge and reproductive state of both
tile prcd;~tor ;~nd prey have to he considered. lkc;mse of space limitation, we have
only intlic;llctl this in T:~bks Y.2 c1nd lJ. 3 as rega rds Opiliones: eggs versus frce-
nwving individu:~ls. The :~ge of th e harvestman is important primarily because of
the sin. mobilit y. und thickness of the cut ide of the harvest man. Some predators
:tre <~hit to sucrcssl'u lly prtdalt' on ly upon sma ller. softer-bod ied harvestmen.
1\'htre:Js otlll'rs only Cill !;1rgcr. more uclive prey. With some predators. juvenile
h:1rvestnwn escape not ire entirely. while the adults arc easily detected and red
upon. The time am! site of activity of the harvestman is also imporlanl. Predators
th:~t feed prinwrily by sight (e.g .. birds and Ji%ards) luke more prey that is larger. di-
unwl. Hnd more active, whereas predators that feed in the litter or soi l (such as in-
sectivorous llHimmals 1111d some nwrsupials) ol"ten take sma ller. less mobile prey
that they ctlll smell or hear moving. Other small generalist predators, such as ants,
spiders . Hnd assassin bugs (Figure 9.3 ), will take whatever they can catc h and
hold. The age of the predator is importcmt because smaller predators can only take
small prey.
Likewise. the reproductive state of the predator can alter the number of prey
taken. Vertebn1t es have been documented to alter th e number of harvestman prey
they tc1ke during difl"erent seasons. Part of this may be due to the increased per-
CL'ni<lge of harvestman prey available during certai n seasons. but in other cases it
appcc1rs to be related to the reproductive state of the predator or availability of other
foods. Examples are some of the bi rds. which take increased numbers of harvest men
when feeding their young. This is especially true of gra nivorous birds that lack a
food-storage organ. such as a crop. These birds feed large quantities of arthropods
(including harvestmen) to the nestlings (MacMillan & Pollock. 1985. and citations
therein)
Loca lity also appears to be importan t. as demonstrated by the toad Bufo mnrinus
(Bufonidae). In its native habitat in Puerto Rico, this species takes an insignificant
number of ha rvest men {Leonard. 1933). whereas in an introduced population in
Queensland. Australia. it feeds almost exclusively on harvestmen (R. J. Raven. pers.
comm.). Indi vid ual preferences or perh aps opportunity also resu lt in uneven cap-
tures of harvestmen within a population of predators. Examination of guts of 67
Cuban tree frogs, Osll!opilus scplelllrimwlis (Hylidae). from Tortola. British Virgin Is-
lands. West Indies. revealed that 6 frogs had eaten Metncynortoides obscurus (Cos-
mctidac): 1 ate a single harvestman. 3 ate 1 harvestmen. and 1 individual ate 9 har-
vestmen (J. C. Cokendolpher, pers. obs.). These data certainly recommend larger
sample sl%es when conducting experiments with predators/prey.
Some predators will drop a harvestman when exocrine gland chemicals are
first em itted (see Chapter 10). This ini tial response has been recorded in the litera
ture as "none feeding." However. other studies have revea led in the case of some
birds. mammals. and ants that they will repeatedly attack the harvestman until it
 Natural Enemies 35 1

Table 9.2 Vertebrate predators of Opiliones

Oplllones prvy Source

PHYlUM CHORDATA: CLASS
OSTEICHTHYES. ORDER
CYPRINIFORMES
Leuciscus cepha/us Phalangium opilio Bristowe. 1949

CLASS OSTEICHTHYES, ORDER

SALMONIFORMES
SD/mo salar and s. trutta Opiltones, Mitopus moria, and Bristowe, 1949; Thomas. 1962
Phatangium opilio

CLASS USSAMPHIBIA, ORDER ANURA

Bombina bomblna
Bufo achalensis
Bufo asper, B. cognatus. B. compactilis
speciosus. B. marinus, B. terrestris
americanus, B. viridis, B. woodhousei
fowleri, B. woodhousei woodhousei,
andBufospp
Bufo hemiophrys
Bufo ictericus (Figure 9.4A)
Hyla cinerea cinerea
Leptodactylus ace/latus
OSteopilus septentrionalis
Pseudacris triseriata maculata
Rana clomitans
Ranagraeca
Rana pipiens
Rana pipiens sphenocephala
Rana ridibunda and Rana spp.
Rana sytvatica
Rana temporaria and/or Bufo bufo
Opiliones BaruS et al.. 1992a
Pachyloldellus goliath Acosta et al., 1995
Opiliooes Leonard. 1933; Smith & Bragg,
1949: Berry. 1970; R. D. Clarke,
1974; Obrtel. 1976; Tomov. 1990;
Yiyit et al., 1999; R. J. Raven.
pers.comm.
Phalangium opi/io Holmberg, 1970
Acutisoma proximum B. A. Buzatto. pers. comm
Opiliones Kilby,1945
Opiliones (LaniatOfes) A. A. Giarea. pers. comm
Metacynortoides obscurus New data
Phalangium opilio Holmberg, 1970
Opiliones Jenssen & Klims tra. 1966
Lacinius dentiger, L. horridus. Beshkov, 1970
Paranemastoma sp., and Trogu/us sp
Opiliones, Pha/angium opilio, and Moore & Strickland, 1954;
Togwoteeus biceps Holmberg, 1970
Eumesosoma nigrum. Leiobunum Kilby, 1945
aurugineum, Leiobunum sp ..
and Vonones sayi
Opiliones Hristova, 1962; Obrtel, 1976
Leiobunum calcar. Odie/Ius picws. Holmberg, 1970
and Phalangium opitio
Anelasmocephatus cambridgei. Lacinius Bristowe, 1949; Houston. 1973
ephippiatus. Leiobunum blackwalli,
L. rotundum, Mitopus moria.
Nemastoma chrysomelas. Nemastoma
/ugubre, Paro/igoloplws agrestis.
Phalangium opi/io, and Rilae/Ja
triangularis
352 Nalurol Enem1es

Table 9.2 Continued

Predator Opillones prey source

CLASS LISSAMPHIBIA, ORDER
CAUDATA
Ambystoma t1grinwn
Ane1des ferreus
Ewycea /ongicavda
Eurycea/uofuga
Hydromantes italiws gormani
Plethodon cinereus. P. cinereus
cinereus. P. dunni, P. glutinosus.
P. idahoensis, P. jordani,
P. oconatuftee. P. vehicutum , and
P. wehrtei
Sa/amandra atra
Salamandra satamandra
Salamandra s. satamandra
-
Opilio parietinus and Phalangium opilio Holmberg. 1970
Opiliones Whitaker et al.. 1986; Petranka. 1998
Opiliones Anderson & Martino. 1967;
Petranka, 1998
Opiliones. Bishopella laciniosa. Hutchison, 1958; Peck, 1974; Peck &
Leiobunum sp., and Phalangodes Richardson. 1976; Petranka. 1998
armata
ISChyropsalis apuana and Trogulus Bruno, 1973
coriziformis
Opiliones smallwood, 1928; Pope, 1950;
Davidson. 1956; Dumas, 1956;
Brandon. 1965; Edgar. 1971; Powders
& Tietjen. 1974; Hall, 1976; wilson
& Larsen. 1988; Petranka. 1998
Plarybunus pinetorum and Jschyropsalis Klewen. 1988
hellwigii
Nemastomatidae and Trogulidae Beshkov & TSOflchev, 1963
Opilio parietinus Klewen. 1988

CLASS REPT!LUA, ORDER

SQUAMATA
Ablepharus kitaibelii Opiliones BaruS et al., 1992b
Alopoglossus angulatus Opiliones Vitt & Caldwell, 2003
Ameiva ameiva Opiliones Vitt & caldwell. 2003
Basiliscus basiliscus Opiliones Barden, 1943
cnemidophorus sacki and c. tessellatus Trachyrhinus rectipalpus Milstead. 1958
Gonatodes hasemani and G. humeralis Opiliones Vitt & Caldwell, 2003
Lacerta muralis, Lacerta taurica, Opillones Bristowe, 1949; Angelov et al..
andLacerta sp 1972a. c
Lacerta muralis muralis Pyza bosnica Mitov.1995
Lacerta viridis Opiliones and zachaeus crista Angelov et ar 1.972b; P. G. Mitov.
pers.obs.
Lacerta vivipara Opiliones and Mitopus morio oarevskiy, 1953; Avery, 1966; Sirbu.
1977; BaruS et al., 1992b
Lacerta vivipara vivipara Phalangiidae. Lacinius horridus. Mitov, 1995
Phalangium opilio, and Rilaena
triangularis
Ophisaurus apodus Opiliones1 Yadgarov. 1974
Prionodactylus eigenmanni Opitiones Vittetal.,1998
Stellio lehmanni Phalangiidae (?) Petrochenko, 1992
Teratoscincus scincus Opiliones Yadgarov. 1977
 Natural Enemies 353

source
CLASS MAMMALIA, ORDER
CARNIVORA
cerdocyon thOus Opiliones (Laniatores) Facure, 1996
Me/esme/es Mitopus mario, Odie/Ius spinosus. Sankey, 1949b
Paroligo/ophus agrestis, and
Phalangium opilio
Mephites mephites Phalangium opilio Holmberg, 1970
Vu/pes vulpes Phalangium opilio Sankey, 1949b

CLASS MAMMALIA. ORDER

CHIROPTERA
Myotis bechsteini Phalangiidae Wolz, 1992
Myotis emarginatus, M. nattereri, and Opiliones Whitaker et al.. 1977; sauerova, 1986;
M. thysanodes Bauerova & CervenY. 1986

CLASS MAMMALIA, ORDER

DIDELPHIMORPHIA
Didelphis aurita Opiliones C.keres & Monteiro-Filho, 2001
Philander trenata Opiliones Santorietal., 1997
Philander opossum (Figure 9.48) Acutlsoma longipes and Goniosoma Gnaspini-Netto, 1993; Gnaspini, 1996;
spelaeum Pellegatti-Franco & Gnaspini, 1996;
Machado et al., 2000

CLASS MAMMALIA, ORDER

INSECTIVORA
crocidura suaveolens Opiliones Burda & Bauerova, 1985; Bauerova, 1988
Erinaceus europaeus Leiobunum b/ackwal/i, L. rotundum, Sankey. 1949b
Miropus morio, Odie/Ius spinosus,
Oligolophus tridens. Opilio parietinus.
Paroligo/ophus agrestis, and
Pha/angium opi/io
Neomys anoma/us, N. fodiens, Opiliones Churchfield, 1979; Kuvikova, 1985a,b;
and N. fodiens bicolor Churcl1field etal., 1991
5olenodon cubanus Cynorta sp Armas, 1987
sorex a/pinus, 5. bendirii, 5. pacificus. Opiliones Obrtel, 1976; Whitaker & Maser. 1976,
5. sylvaticus, 5. trowbridgii, 5. vagans, Kuvikova, 1986; Churchfield &
and 5orex spp Bmwn, 1987
5orex araneus Opiliones. Carinosroma omatum. Lacinius Bristowe, 1949; Rudge, 1968;
dentiger. Mitostoma chrysomelas, Pernetta, 1976, 1977; Butterfield
Nemastoma lugubre, Paroligolophus et al., 1981; Churct1field, 1982;
agrestis. Paranemastoma radewi, sauerova, 1982, 1984; KuVJkova,
and Pyza bosnica 1985a; Churchfleld et al., 1991 ;
Mitov.1995

,,,.,,,,./,
3 54 Ndlural Encm1es
Table 9.2 Continued
Predator
Sorex mrnuws
CLASS MAMMALIA. ORDER
RODENTIA
ApOdemus flavrcollrs
Apodemus sylvatrcus
Clethrlonomys glareolus
Mus musculus
Onychomys torridus longicaudus
Peromyscus leucopus
CLASS AVES. ORDER
CORACUFORMES
Merops apiaster
CLASS AVES. ORDER
GALLIFORMES
Bonasa umbel/us
Colinus virginianus
cotumix cow rnix
Gallus domesticus
Perdix perdix perdix
CLASS AVES. ORDER
CHARADRIIFORMES
Charadrius mongo/us
Cursorius curso
Laruscanus
Pluvialis apricarius
vane/Ius vane/Ius
CLASS AVES. ORDER
CUCULIFORMES
coccyzus americanus and
c. erythropthalmus
Opiliones prey
Oplliones, Paranemastoma sp., and Pernetta, 1976; Grainger & Fairley,
Paranemastoma radewi 1978; Butterfield et al., 1981;
Bauerova, 1982, 1984; Kuvikova,
1985a; Churchfield et al., 1991;
Mitov, 1995
Opiliones, Lacinius sp., Nemastoma Obrtel. 1973a, 1974, 1976;
sp., Oligo/ophus sp., and Plarybunus sp. Obrtel & Holi~ova. 1974, 1980, 1983
Op1l iones Obrtel & Holi~ova, 1979, 1983
Opiliones Obrtel. 1973b, 1974, 1976; Holi~ova
& Obrtel, 1979; Obrtel & Holi~ova.
1974,1983
Pha/angium opilio Holmberg, 1970
Opiliones Horner et al., 196S
Eumesosoma roeweri and Sclerobunus J. c . Cokendolpher, pers. obs
robustus robustus
Opiliones Crampetal .. 1985
Opiliones Stewart, 1956
Opiliones Bristowe, 1949
Phalangium sp Mihaylov, 1995
Phalangium opilio Holmberg, 1970; Clark & Gage, 1997
Opiliones Glutz von Blotzheim et al., 1973
Oplliones Zlatin, 1968
Opiliones cramp et al., 1983
Opiliones Bakke, 1970; Glutz von Blotzheim &
Bauer, 1982
Opiliones Alldreeva, 1989
Opiliones cramp et al., 1983
Opiliones McAtee, 1926, 1932; Brlstowe, 1949 '
 Natural Enemies 355

Oplltones prey source

CLASS AVES, ORDER
GRUIFDRMES
Porphyrio porphyria melanotus Opiliooes caroll, 1966; Glutz von Blotzheim
etal.,1973

CLASS AVES, ORDER

PASSERIFORMES
Acrocepha/us arundinaceus, Opiliones Cramp et al., 1992
A. dumetorum, A palustris,
A. schOenobaenus, and A. scirpaceus
Alauda arvensis Oplliones Crampetal., 1988
Anthus campestris, A. cervinus, Opiliones Cramp et al.. 1988; GajdoS &
A. spinoletta, and A. trivia/is KriStin,1997
Anrhus pratensis Opiliones and Mitopus morio Skaret al., 1975; Hilgvar & Ostbye,
1976; Cramp et al., 1988
Cenhia brachydacty/a Opi1iones Cramp et al.. 1993
Certhia familiaris Opiliones and Platybunus bucephalus KriStin, 1992; Cramp et al.. 1993;
KriStin & Baumann, 1996; GajdoS
& KriStin,1997;Szentkirillyi &
KriStin,2002
Cettia cetti Opiliones Cramp et al.. 1992
corvus corone, c. frugi/egus. Opiliones and Phalangiidae cramp et al., 1994b
and corvus monedula
Cyanocorax caeruleus Gony!eptidae Reinert & Bornschein, 1998
cyanopica cyanus Opiliones Crarnpet al., 1994b
Dendroica magnolia Opiliones MCAtee. 1926
DolichOnyx oryzivorus Opiliones Oillery. 1961;Wiens. 1969
Emberiza citronella, E. melanocephala, Opi1iooes S. Simeonov, 1981 ; Cramp et al.,
and E. schOeniclus 1994a
Erithacus rubecula Opiliones and Lacinius ephippiaws KriStin, 1992; KriStin&Baumann,
1996; GajdoS & Kn$tin, 1997;
szentkirillyi & KriStin. 2002
Ficedula albicollis Opiliones. Mitopus sp., Platybunus BureS, 1986; KriStin, 1992. 2002;
bucephalus, and Platybunus sp Cramp et al.. 1993; KnStln &
Baumann, 1996; GaJdOS & KnStin,
1997; szentk1rillyr & KrrStnl, 2002
Ficedula hypoleuca Opiliones and Mitopus morio Dornbusch, 1981; sacller &
Dornbuscll. 1990; C1amp
et al., 1993
Ficedula parva Opiliones c ramp et al., 1993
Fringilla coelebs and F. montifringilla Opiliones Prokolyeva. 1961; Darak.tcllrev,
1981; Cramp et al. 1 99<~b. Knstrn
& Baumann. 1996
356 Natural Enern1 es

Table 9.2 c ontmued

Predator Opiliones prey source

Garrulus glandanus Opihones and Phalangi1dae Cramp et al 1994b
Lamus col/uno Opiliones, Phalangiidae, Leiobunum Cramp et al 1993: Nikolov, 2002
rotundum, and Zachaeus crista
Lamus excu/Jitor. L. mmor,and Op1hones Crampetal 1993

Locustella fltiV!atliiS and L. naevia Opihones Crampetal 1992

Lullu/a arborea Qplliones crampetal 1988
MOntiCOia solitanus Opiliones Glutz von Blotzheim & Bauer, 1988a
MOntlfrmglllanivalis Opiliones Cramp et al., 1994b
Motacilla alba, M. cinerea, and M. flava Opiliones Daraktchiev, 1981; Cramp etal., 1988
Muscicapa striata Opiliones crampetal., 1993
Nannus hiema/is Opiliones McAtee, 1926
Oenanthe finsch1. 0. hispanica. Opiliones Belskaja, 1965; wartmann, 1985;
0. isabel/ina, and 0. oenanthe Cramp et al., 1988: Glutz von
oenantt1e Blotzheim & Bauer, 1988a; Szentkirfllyi
&KriS tin, 2002
Parus ater and P. caeru/eus Opiliones and Mitopus mario Sacher & Dornbusch, 1990; KriStin, 1992;
Crampet al., 1993; KriStin & Baumann,
1996; Mattes et al., 1996; GajdoS &
KriStin, 1997; Mihfll, 1998; szentkir.31yi
& KriStin, 2002
Parus major Opiliones, Mitopus mario. Betts, 1955; Dornbusch, 1981; Sacher
Nemastoma lugubre, and Rilaena & Dornbusch, 1990; KriS tin, 1992;
triangularis Cramp et al., 1993; KriStin & Baumann,
1996; GajdoS & KriS tin, 1997;
Szentkir.31yi & KriStin, 2002
Passer domesticus Opiliones, Qpilio parietinus. and s. D. Simeonov, 1964; Holmberg, 1970;
Pha/angium opilio MacMillan & Pollock, 1985; Ivanov,
1990; GajdoS & KriStin, 1997
Passer montanus and Passer sp Opiliones Cirdei, 1958; S.D. Simeonov, 1963;
KriStin, 1988b; TOrOk, 1990;
Cramp et al., 1993; KriStin &
Baumann, 1996; GajdoS & Kri Stin,
1997; szentkir.31yi & KriStin, 2002
Passercutus sandwichensis Op!liones and Pha/angium opilio Dillery, 1961; Wiens, 1969;
Holmberg, 1970
Pastor roseus Opiliones 2 Dementjev et al., 1954
Penthesres atricapillus Opiliones McAtee, 1926
Pheucticus /udovicianus Leiobunum a/drichi, L. vittatum, Edgar, 1971
andL.calcar
Phoenicurus ochruros Opiliones and Platybunus bucepha/us KriStin & Baumann, 1996; GajdoS
& KriStin, 1997
Phoenicurus ochruros gibraltariensis Opiliones Cra mp et al., 1988; Glutz von Blotzheim
and P. phoenicurus & Bauer, 1988a

Oplllones prey
PhOenicurus phoenicurus phoenicurus Opiliones and Mitopus moria
Phylloscopus borealis and P. trochilus Opiliones
Phylloscopus collybita Opiliones and Phalangiidae
Phylloscopus sibilatrix Opiliones. Phalangiidae. and Platybunus
bucephalus
Pica pica Oplliones, Phalangiidae, Phatangium
opilio, and Rilaena triangularis
Pitangus sulphuratus Opiliones
Prunella modularis Opiliones, Phalangiidae, Lacinius
ephippiatus. Mitopus moria.
OligOiophus tridens. Platybunus
bucephalus, and Platybunus spp
Prunella modularis modularis Opiliones, Lacinius ephippiatus,
Leiobunum rupestre. Mltopus mario,
Oligolophus tridens. Paranemastoma
quadripunctatum, and Platybunus
bucephalus
Pyrrhocorax pyrrhocarax Opiliones
Regulus ignicapillus Opiliones and Platybunus sp
Regulus regulus and R. teneriffae Opiliones
Remiz pendulinus Opiliones
SaxicOia torquata Opiliooes
Seiurus aurocapi/lus Opiliones
Sitta europea Opi!iones. Phalangiidae. Lacinius
eohippiatus, Mitopus mario.
Ptatybunus bucephalus, P. pa/lidus.
P. pinetorum, and Platybunus spp.
Stumus roseus, s. unicolor, and Opiliones
s. vulgaris
Sylvia atricapilla, s. borin. s. curruca, Opiliones
s. nisoria. sylvia sp., and Sylvia undata
Sylvia communis Opiliones, Leiobunum sp., Mitopus sp.,
and Phalangium sp
Tarsiger cyanurus cyanurus Opiliones
T1ch0droma muraria Opiliones
Toxostoma rufum Opiliones
Natural Enemies 357
source
Emmrich, 197Sa; Glutz von Blotzheim
& Bauer, 1988a: sacher &
Dornbusch. 1990
Cramp et al.. 1992
KriStin, 1988a, 1991, 1992;KriStin&
Baumann, 1996; GajdoS & KriStin,
1997; szentkirtllyi & KriStin, 2002
KriStin, 1991, 1992; Cramp et al., 1992;
KriStin & Baumann, 1996; GajdoS &
KriStin, 1997; szentkinilyi &
KriStin,2002
Petrusenko & Talposh, 1981; Cramp
etal., 1994b
Argel-de-Oiiveira et al., 1998
Emmrich, 197Sb; cramp et al., 1988;
KriStin, 1993b; KriStin & Baumann,
1996; GajdoS & KriStin, 1997;
Szentkirillyi & KriStin, 2002
Emmrich. 197Sb; Tomek. 1988
Cramp et al., 1994b
Thaler-Kottek, 1973; Cramp et al 1992
Lbhrl & Thaler, 1980; Cramp et al..
1992
szentkirillyi & KriStin, 2002
KriStin & Baumann. 1996
MCAtee, 1926, 1932
Oaraktchiev. 1981 ; KriStin, 1992,
1993a, 1994;Crarnpet al., 1993;
KriStin & Baumann, 1996; GajdoS &
KriStin, 1997; szentkir.illy1&
KriS tin, 2002
cramp et al.. 1994b
Prokofyeva, 1961 : Ganya & Zubkov.
1988;Crampetal.,1992
Emmricll. 1973, 1974; Cramp
etal., 1992
Glutz von Blotzllenn & Bauer. 1988a
cramp et al.. 1993
MCAtee, 1926
358 N<ltllral Enern1es

Table 9.2 continued

\ Predator Opiliones prey source

nogloclytes troglodytes Qp11iones, Lacinius ephippiarus. and Prokofyeva, 1961; Sellin, 1969; Dallman,
M1topus morio 1987; Cramp et al., 1988; Kri~tin,
1992; KriStin & Baumann, 1996;
GajdoS & KriStin, 1997; Szentkirillyi &
KriStin, 2002
Turdus merula and T ph!lomelos Op1!iones Oaraktchiev, 1981; TOrOk, 1985;
Cramp et aL, 1988; Glutz von
Blotzheim & Bauer, 1988b; Kri~tin,
1992; KriStin & Baumann. 1996;
GajdoS & KriStin. 1997; 5zentkiriltyi
& KriStin, 2002
Turdusp1/ans Op1110nes and Phalangiidae Oaraktchiev, 1981; LObcke & Furrer,
1985; cramp et al., 1988; Glutz von
Blotzheim & Bauer, 1988b
vermivora chrysoptera Opiliones Edgar. 1971
Vlfeosylva philadelphica Opiliones MCAtee, 1926
xanrhocephalus xanthocephalus Opiliones Orians. 1966
zoothera sibirica Opiliones Moroschenko, 1986; Glutz von
Blotzheim & Bauer. 1988b

CLASS AVES. ORDER PICIFORMES

Dendrocopos syriacus, D. major. and Opiliones Cramp et al., 1985; 5zentkir31yi &
D. medius Kri~tin,2002

Dryobates pubescens Opiliones MCAtee. 1926

Picoides major Opiliones and Platybunus bucepha/us KriStin & Baumann. 1996; GajdoS
& Kri~tin, 1997
Picoides medius, P. minor, P. syriacus. Opiliones Prokofyeva. 1961; Ruge, 1969;
andP. tridacrylus Glutz von Blotzheim & Bauer. 1980;
Pechtltek & KriStin. 1996; GajdoS &
KriStin, 1997; MiMI, 1998
Picuscanus Opiliones Oaraktchiev, 1981

CLASS AVES. ORDER RALLIFORMES

Crexcrex Opiliones Glutz von Blotzheim et al., 1973;
Cramp et al., 1980
Gallinula ch/oropus Opiliones and Lophopilio pa/pinalis cramp et aL. 1980

CLASS AVES. ORDER STRJGJFORME5

Otus scops and orus scops scops Opiliones Glutz von Blotzheim & Bauer. 1980;
Cramp et al., 1985

original source citations for many of the birds are listed in Cramp et at. (1983-1994) .
t. Yadgar011 (1974) listed arachnids as components of the food of the lizatcl Ophisaurus apodus- "SCOrpionuy," "Falangi." "Paukl," "Kieshi," which translate
to scorpiones, Araneae. ACari" -but it is not clear if the name "Falangi" is Opiliones or SOtifugae. In older Russian merattKe on Arachnida ~Phalangi"
SOli fugae
2. oementjev et at. (1954) also listecl"ptlalang" as food of the bird Pastor roseus . This too could possibly refer to SOiifugae.
 Natural Enemies 359

Table 9.3 Invertebrate predators of Opiliones

!lli~==.._' ____,____Opl
~llonesprey source
PHYLUM MOLLUSCA: CLASS
GASTROPODA. ORDER
STYLOMMATOPHORA
Polygyraalbolabris Leiobunum aldriehi Edgar, 1971

PHYLUM ptATYHELMINTHES:
ORDER TRICLADIDA
Cephaloflexa araucariana Gonyleptidae carbayo et at, 2002; carbayo &
Leai-Zanchet. 2003

PHYLUM ARTHROPODA:
CLASS CHILOPODA, ORDER
UTHOBIOMORPHA
UthObius forficarus Opiliones. Opilio parietinus, and Paroligolophus Sankey, 1949b; Spoek, 1964
agresris
Lithobius sp. Odie/Ius pictus and Phalangium opilio Holmberg, 1970
Lithobius vulgaris Leiobunum aldrich!, L. politum, and L. virrarum Edgar, 1960; Holmberg, 1970

CLASS DIPLOPODA, ORDER

NEMATOPHORA
Undetermined species Opiliones Barnes,1968

CLASS ARACHNIDA, ORDER

ARANEAE
Achaearanea lunata Rilaena cl. serbica P. G. MltOV, pers. ObS
Agelena sp. Paroligolophus agrestis Bristowe. 1941
Agelena naevia Opiliones Bilsing. 1920
Agetenopsis sp. Leiobunum vittatum and Togwoteeus biceps Holmberg, 1970; R. Macias-Ord6iiez.
pers.comm
Amaurobius sp Oicranolasma scabrum Gruber, 1993
Aranea cucurbitina Leiobunum blackwalli Bristowe. 1941
Araneus diadematus Leiobunum rotundum and Paranemastoma Bristowe. 1941; Mitov. 1995
aurrgerum aurtgerum
Argiope argentata Opifiones and Sclerobunus robustus Robinson & Robinson, 1970;
J. c . Cokendolpller. pers. obs
Coelotes sp Dicranolasma scabrum Gruber.1993
coetotes terrestris Opiliones, Mitostoma chrysomelas. and Petto. 1990
Paranemastoma quadripunctawm
Corinnidae (Figure 9.3A) tporangaia pusrulosa B. A. Buzauo. pers. comrn
Ctenus fasciatus Acutisoma longipes (eggs and adults) and Gnaspuli- Netto, 1993; Gnaspuu. 1996.
Goniosoma spelaeum MactladO et al., 2000
ctenus sp. 1 Bourguyia albiomata Machado & 011ve1ra. 2002
360 N.:ltural Enem1es
Table 9.3 Continued
Predator
Cteous sp. 2 {Figure 9.38)
Enoploctenus cyclothOrax
GlildiCOsa gulosa
Hypocllilus gertsc/Ji
Latroclecws geometriws
{Figure9 .3C)
LatrOdectus hesperus
Latrodectus variolus
Linyp/Jiasp
Unyphiidae. Erigonidae
Lycosasp
Meta merianae
Meta segmentata
Metellina merianae
Nenene clathrata
Pardosa fuscula
Pardosa groenlandica
Pardosa mackenziana
Pardosa modica
PhOICUS pha/angioides
Photcus spp
PIJOneutria nigriventer
Physocyclus enautus
Pisaura sp.
SCytodes sp
Steatada bipunctata
Steatoda castanea
Steatoda sp
Steatoda triangutosa
Tidarren sisyphoides
Tegenaria sp. (juvs.)
Tegenaria sp
Tegenaria agrestis {juvs.)
Tegenaria agrestis
Oplliones prey source
LOngipema zonata G. Machado, pers. comm .
Acut/soma /ongipes and Discocyrtus sp Willemart & Kaneto, 2004; G. Machado,
pers. comm
Leiobtmum a/dnc/Ji, L. politum, and L. vitratum Holmberg. 1970
Leiobunum sp Shear, 1969
Opiliones and cosmetidae weed. 1892; J. Warfel, pers. comm
Leiobunum townsendi J. C. Cokendolpher, pers. obs
Nelima elegans w. Reeves, pers. comm
Ritaena batcamca Mitov. 1995
Opiliones Nentwig, 1980. 1983
Opi/io parietinus SilhavY. 1956
Leiobunum blackwalli and Parotigotophus agrestis Bristowe, 1941
Paranemastoma radewi Mitov, 1995
Nelima aladjensis P. G. Mitov. pers. cbs.
Mitostoma gracile P. G. Mitov, pers. obs.
Opilio parietinus a11d Phalangium opilio Holmberg, 1970
Pha/angium opilio Holmberg, 1970
Odie/Ius pictus and Phalangium opilio Holmberg. 1970
Phatangium opilio Holmberg. 1970
Leiobunum blackwalli Bristowe. 1941; Nentwlg, 1983
Mitopus morio and Paranemastoma Uhlenhaut. 2001
quadripunctatum
Bourguyia albiornata Machado & Oliveira. 2002
Leiobunum townsendi J. c. Cokendolpher. pers. obs
Leiobunumblackwalli Bristowe, 1941
Mischonyx cuspidatus Mestre & Pinto-da-Rocha. 2004
Lacinius dentiger and Phalangium opilio Mitov, 1995, 2000b
Lacinius dentiger, Mitopus morio. and Mitov. 1995
Phalangium opilio
Odie/Ius lendli and Opilio ruzickai P. G. Mitov, pers. obs.
Phatangium opilio Mitov, 1995
Opiliones P. H. Pache, pers. comm
Opilio parietinus P. G. Mitov, pers. obs.
tacinius dentiger. L. horridus. Leiobunum Sankey. 1949b; Mitov. 1995, 1997b, 2000b.
rume/icum. Mitopus moria. Ne/ima a/adjensis. pers. obs.
0/igo/ophus tridens. Opilio ruzickai,
Phalangium opilio. and Trogutus tricarinatus
Odie/Jus /endli P. G. Mitov, pers. obS.
Pha/angium opi/io Mitov, 1995, pers. obs.

Tegenaria atrica
Tegenaria derhami
Tegenaria ferruginea
Tegenaria nemorosa
Tegenaria silvestris
Thanatus imbecillius
Theridiidae
Theridiidae. Linyphiidae, and
Erigonidae
Theridion pictum
Theridion sp.
Theridium frondeum
Theridium omatum
Trechalea keyserlingi
Undetermined species
xysticus bifasciatus
xysticus viaticus
Zilla rhorelli
CLASS ARACHNIDA, ORDER
OPILIONES
Acanthopachylus aculeatus
Acutisoma longipes
Acutisoma proximum
Bourguyia a/biornara
Dicranolasma scabrum
Discocyrtus Oliverio/
Erginulus clavoribialis
Gagrellinae
Jschyropsatis hellwigii
Lacinius dentiger
Lacinius ephippiatus
Lacinius horridus
Laniatores
Natural Enemies 361
SOUr<e
Leiobunum blackwalli Bristowe, 1941
Pha/angium opilio Holmberg, 1970
Opiliones. Lacinius dentiger. Opilio ruzickai, Nentwig, 1983; Mitov. 1995. 2000b
Paranemasroma aurigerum aurlgerum, and
Phalangium opi/io
Mitosroma gracile P. G. ~ itov. pers. obs.
Mitopus moria and Paranemastoma Mitov, 1995, 2000b
aurigerumryla
Opiliones Guseinov, 2004
Opiliones and Mischonyx cuspidatus Nentwig, 1980: Mestre & Pintoda
Rocha, 2004
Opiliones Nentwig, 1980
Opiliones { uczak & Dabrowski-Prot. 1970
Phalangium opilio Mitov,1995
Phalangium opilio Holmberg, 1970
Phalangium opilio Holmberg. 1970
Discocyrtus sp G. Machado. pers. comm
Opillones. Bourguyia albiornata, Opilio ruzickai. Savory, 1928: Stipperger. 1928; Bristowe.
Paranemasroma aurigerum aurigerum, 1941, 1949; creme et al.. 1969; Edgar,
Phalangium opilio, Siro sp., and Trogulus 1971: Mitov. 1995, 2000b: Machado
nepaeformis & Ol iveira. 2002
Leiobunum blackwalli Bristowe. 1941
Leiobunum rotundum Bristowe. 1941
Opilio parierinus P. G. Mitov, pers. obs
Acanthopachylus aculeatus (only eggs) Capocasale & Bruna-Trezza. 196<1
Acutlsoma longipes {only eggs) Mac11ado & Oliveira. 1998
Acutisoma proximum (on ly eggs) Ramires&Giaretta,1994
Bourguyia albiornara (only eggs) Machado & Ohve1ra, 2002
Dicranolasma scabrum Gruber, 1993
oiscocyrrus oliverioi (only eggsJ Elpino-Campos et al., 2001
Erginulus clavotibialis (eggs and early nymphs) Goodnight & Goodnight. 1976
Opiliones Star~a.1976a
Leiobunum sp.. Opilio parietirws ver11oeff. 1900; Kiistner. 1926
Lacinius dentiger Mitov,1988
Nemastoma lugubre Todd,1950
Opiliones Star~ga, 1976a
Laniatores Mello-Leltiio. 1932
362 Nauual Enermes

Table 9.3 Contrnuec!

Predator Oplllones prey source

Lerobuninae Lacinius ephippiaws, L. horridus, Leiobunum sp., Pfeifer. 1956
-
Opi/io parietinus, Plralangium opi/io, and
Rilaena triangularis
LetObtmum aldnc/Jt Leiobunum aldrichi Edgar, 1971
Leiobunum blackwalli OligolopiJUS tridens Bristowe, 1949; Parisot. 1962
Leiobumtm limbatum Amilenus aurantiaws and Leioburwm limbarum Stipperger, 1928
Leiobunum po/rlttm Leiobunum politum Edgar,1971
Lerobunum rotundum Leiobunum rotundum, Lacinius ephippiatus, Henking, 1888; Bristowe, 1949; ~il havy, 1956
Oligoloplws tridens. and Phalangium opilio
Lerobunum virtatum Leiobunum vittatum Edgar. 1971
Metaplatybunus carneluWi Metaplatybunus carneluttii Karaman, 1995
Mitopus morio Opiliones. Lacinius ephippiatus, Leiobunum Kastner. 1931a; Bristowe, 1949; Todd, 1949;
rotundum, Mitopus morio, 0/igolophus Phillipson, 1960a: RUffer, 1966; Tischler,
agrestis, Oligolophus sp., Oligo/ophus tridens , 1967; Star~ ga, 1976a; Littlewood &
ParoligOiophus agrestis, and Platybunus Littlewood. 1989
bucephalus
Ne/ima paessleri Nelima paess/eri Holmberg et al., 1984; Angerilli & Holmberg,
1986
Odie/Ius trogu/Oides Odie/Ius troguloides Juberthie, 1964
Oligolophus tridens Opiliones. Leiobunum rotundum, Mitopus Henking, 1888; Bristowe, 1949;
morio. Oligolophus tridens, and Pha/angium Star~ga, 1976a
opilio
Opilio parietinus Opiliones. Leiobunum rotundum, Opilio Henking, 1888; Roters, 1944; RUffer, 1966;
parietinus, Oligotophus tridens, and Sta r~ga, 1976a
Pha/angium opi/io
Opilio saxatilis Opiliones Stan~ga, 1976a
Opiliones Opiliones 0 . Pickard-cambridge, 1890a; Heymons,
1915; Bristowe, 1949; Sankey, 1949a
Pachy/us quinamavidensis Pachylus quinamavidensis (only eggs) Juberthie & Munoz-cuevas, 1971
Paranemastoma Paranemastoma quadripunctatum Immel, 1954
quadripunctatum
Paroligolophus agrestis Paroligolophus agrestis Sankey, 1949b
Phalangiidae Opiliones Star~a. 1976a
Phatangium opilio (nymphs Opiliones. Leiobunum rotundum, Mitopus mario, Henking, 1888; ROhm, 1926; Roters, 1944;
and adults) 0/igolophus tridens, Opi/io parietinus, and Bristowe, 1949; ~ilhayY, 1956; Klee &
Phalangium opi/io (nymphs, eggs) Butcher. 1968; Edgar, 1971; Starega,
1976a; Mitov, 1988
Rilaena triangularis Rilaena triangularis Sankey, 1949b
zachaeus crista Phalangium opilio and Zachaeus crista Mitov, 1988
Zygopachy/us albomarginis Zygopachylus albomarginis (only eggs) Mora, 1990; Cokendolpher, 1993

CLASS ARACHNIDA,
ORDER SCORPIONES
Undetermined species Opiliones Cloudsley-Thompson, 1958

CLASS INSECTA, ORDER
COLEOPTERA
carabiclae
carabus violaceus
Prerostichus oblongopunctatus
Sepedophi/us sp. (larvae)
CLASS INSECTA, ORDER
DIPTERA
Diptera
CLASS INSECTA, ORDER
ENSIFERA
Strinatiasp.
CLASS INSECTA, ORDER
HEMIPTERA
Lygussp
Pentatoma prasina
Reduvius personatus
Zelurus travassosi (nymphs
and adults) (Figure 9.3Dl
Ze/us sp.
CLASS INSECTA, ORDER
HYMENOPTERA
camponotus abdomina/is
Formica balcanina
Formica cinerea
Formica lugubris
Formica pratensis
Formicidae
Pachycondyla villosa
(Figure9.3El
Paravespula germanica
CLASS INSECTA, ORDER
MECOPTERA
Bittacus mastril/ii
Natural Enemies 363
soun:e
-
Opiliones Hillyard & sankey, 1989
Opiliones and Oligolophus sp Sankey, 1949b; Spoek, 1964
Opiliones sergeeva & Grfinthal, 1988
Gonlosoma spelaeum (only eggs) Gnaspini, 1995
Opiliones Obrtel,1976
Acutisoma longipes (only eggs) Machado & Oliveira, 1998; Machado, 2002
Opiliones Wheeler. 1976
Phalangium opilio sankey, 1949b
Qpiliones Sankey, 1949b
Acutisoma longipes (nymphs and eggs) Gnaspini-Netto. 1993; Gnaspini, 1995, 1996;
and Goniosoma spelaeum (nymphs, Machado & Oliveira. 1998; Machado et al.,
adults, eggs) 2000: Machado, 2002
Opiliones Edgar. 1960, 1971
Discocyrtus pectinifemur Matthiesen, 1980
Opilio saxatilis P. G. MIIOV, pers. obs
P/Jalangium opilio Mitov.1995
Mitopus mario, Mitostoma cluysomelas. Cllerix & Bourne. 1980
Oligoloptws tridens, Platybunus
bucepha/us, and Trogulus sp
Lacinius horridus Mitov. 1995
Opiliones. Bourguyia albiomata (only eggs). St1pperger. 1928: Bishop, 1950; Creme et al.,
Gonylepres saprophilus (only eggs). and 1969; Mora. 1987; Mact1ado & Raimundo.
Zygopachy/us a/bomarginis (only eggs) 2001 : Machado & Olive1ra, 2002
Bourguyia a/biomata (only eggs) Machado & Olivera, 2002
Opilio ruzickai P. G. MIIOV, pers. Ob~.
Ooiliones IWi'IS<lkl, 1998
Figure 9.3. Invertebrate predators of Dpiliones. (A-C) Spiders (both wandering and web builders) are the
predominant invertebrate predators of harvestmen. (A) lporangaia pustulosa (Gonyleptidae) being fed upon by a
corinnid spider in southeastern Brazil {photo: B.A. Buzatto). (B)A female Longipema zonata (Gony\eptidae) being
eaten by a Crenus sp. (Ctenidae) in southeastern Brazil (photo: G. Machado). (C) A cosmetid in the web of the
spider Latrodectus geometricus (Theridiidae) in Trinidad (photo: j.G. Warfel). (D) A nymph of the assassin bug
Zelurus rravassosi (Reduviidae) preying on the harvestman Coniosoma spelaeum (Gonyleptidae) inside a cave in
southeastern Brazil (photo: P. Gnaspini). (E) The ponerine ant Pachycondyta villosa preying on eggs of Bourguyia
albiornata (Gonyleptidae) in southeastern Brazil (photo: G. Machado). Scale bars= 10 mm
 Natural enemies 365

presumably expels all the defensive chemicals and then will be eaten (Holmberg.
1970: Machado et al.. 2005 ).
Because of the difficu lty in observing direct predation of harvestmen. most
records in the literature are based primarily on gut con tents or feces of predators.
eaten remains in or below webs of spiders. and feedings done \Vith captive animals.
Indeed. severa l of the predator/ prey records listed in Table 9.3 are based upon cap-
tive feedings. Some of these events would never take place in nature because the
predator and prey do not occur in the same habi tat (e.g .. the araneid spiders of the
genus Argiope and triaenonychid harvestmen of the genus Sdaolnums) or because
they are not a regular food item for the predator. For example. Argel-dc-Oliveira ct a]
( 1998) reported that a great kiskadee. Pilaugus su/plrumlrrs ('l'yrannidm:). offe red a
harvestman to a captive fled gl ing, but the item was refused. Some of the feeding
studies in the laborat ory can be misleading because a well-fed animal will not re-
spond the same as a hungry animal. Some young. well-fed mammals wil l "play"
wit h Opiliones rat her than cCJtthem U-C. Cokcndolphcr. pcrs. obs.)
From Tables 9.2 and 9.3 it uppears that p;;rsscrinc birds <r rc by far the most di-
verse group of predators on harvestmen. Frogs and toads (Figure 9AA). insectivo-
rous mammals. spiders. a nd other harvest men arc the other main documented pred-
a tors. However. the predomimmcc of one predator group over the other may be
misleadi ng because of the differences in the numbers of stud ies n.garding fonrging
habits of each taxonomic group. Preda tion on harvestmen certainly occurs more
often and by more predators than ;:rre reported herein. ~ l os t studies on prLxlators of
Opiliones are not int ensive or recorded over a long period of time. In many other L'X-
amples in Tables 9.2 and 9.3. harvestmen reportedly mad ~.: up tl very minor per-
centage of the prey it ems fou nd . In thcsentses it is often diftkult to ddermint il' tht
harvestmen prey were rare bccmrsc of the season when l<rrgcr mobik h;trwstmtn
a re rare or because they arc not a nonnal prey itern of a par tkul t~r predator. 1\t'caust
larger predatory lish strike at any tll"tivc arthropod on the W<tter"s surl';tce. the rarity
of lish eating harvestmen almost certainly is b~.:causc of the lack or lwrvestmen
falling into water rather th an the refusal of such food by aquatic prtxlators.
In some cases Opiliones were rttorcled in signilkcrnt numbers: 2 ~J:, of foodstuiTs
of the mouse ilpodt'IIIIIS}Javimllis (Mu rida el were Opilioncs (()brtcl. 1"J 7 41: 42':i. of
the prey examined fro m the s~:o rpionll y Billm'm nruslrillii (Bitttll"idm) IVl're Opiliones
(Iwasaki. 199~ ): and at:cording to Hci gvar ctml (')stbye 11 9/(,f. Opilionts were unt ol
the most important (based on fresh weight I food sou rns fo r meadow-pipits :\ullw .~
pmlmsis (Motacillid;:wl. The prevailing food so urces for 1'/wcllii'III"IIS odrnm1s yilmrl-
/arknsis (Turd id;tc) arc Opilinncs. i\ranca~.:. and inserts ll:lutz I'Oll Blutzlll'im &
Bau er. 19HSa ). l'halangiids, mainly P/utnlnuws hun'/llr11/11s. Wt'l"l' <HlHlll)!. the most
J'requent and abundant prty spe~.:ies recorded J'ur tht nut hatch (KriStin. llJ94l and
the collared llyc<llcher (KriStin. 20021. 'l'ht' high tX'fl't'tll<r)!.l' of this httn't'Stm;m
species in the food of these two birds corrcspondetl with its higlll'Sl ahund;ttH'l' ol
hilrvcstman species in fir-beech l'orcst !Kri.';tin. 2002). t\rcordin)!. to Buttlrlicld ct <tl.
( I ~X I J. Son'.\"lllillll/11 .~ (Sorkida ~.:) cHIS arthrupods (lwrvcstmcn. spitl l'rs. hVl'l ks. 11il'S
and CiJtcrpillars) almost exclu sively. Bcrucn,v{r ( llJ X21 <~ l so notl'd thl' importcull'l' 111"
366 N:mual Enem1es

Figure 9.4. {A) The toad Bufo ietericus {Bufonidae) feeding on the gonyleptid
harvestman Acutisoma proximum {photo: B. A. Buzatto). Note tips of two
harvestman legs sticking out of mouth (arrow). (B) The marsupial Philander
opossum {Oidelphidae) is a widespread predator of several species of South
American harvestmen, mainly those that live inside caves (photo: A.A. Giaretta).
Scalebars=Scm.
 Natural Enemies 367

arachnids (espedally Opiliones) in the diet of this same shrew. Pcrnetta {1976)
ranked the foods of this shrew and Sorex amneus. with Opiliones being one of the
dominant components. Whereas the pigmy shrew ate mainly Nemastoma sp.. the
common shrew selected larger species of Opiliones. Examination of Pl!iltmckr
opossum (Didelphidae) feces revealed that a harvestman from caves in Brazil (Gonia-
soma spe/aeum) was an important food item. especially when foods on the surfa ce
(mainly fruits) were scarce (Pellegatti-Franco & Gnaspinl. 1996: Figure 9 .4B ). This
species of marsupial was observed to prey on up to 1 0 harvest men during a si ngle
visit to the cave. One of the major food items was also Opiliones for the opossum
Dide/pl!is aurila (Caceres & Monteiro-Filho. 200 I ).
As reviewed in Chapter 8. cannibalism is common among Opilioncs and their
eggs. Cannibalism occurs more often upon carlier-instar juveniles and during or
after molting. when the cuticle is soft and easi ly penetrated by the chelicerae of other
harvestmen. According to Edgar ( 1971) and Machado and Oliveira ( 1998 . .200.2).
cannibalism upon eggs is more common than upon juven iles an d adults (sec also
Chapter 12).

PHORESY, ENDOZOISM, AND EPIZOISM

Phoretic, endozoic. and epizoic associations arc much marl' common than reported
in the literature. Possibly these arc the most common but unreport ed associations.
The dclinitions of these three interact ions arc often given as anything living/riding
in or on a n anima l. but for the purpose of this du1pter. we an using dl'linitions that
arc more restrictive. A phorctic association is Olll' in which <m org;mism receives n1r-
riage on the outside of a harvcstmcm. It docs not feed while riding on th t htlf-
vestman and only receives a means of transport. Th is tnll1$port may be ;1c ti v~o tsuch
S n insect riding a harvestman) or intJCtivc (s uch as fun!;(a l spores). t\n cndowk
nssodalion is one in which an orga ni:;m passes through the digestive systtm of the
Opiliones unharmed and thereby receives transport to a new location. We do not in-
clude internal parasitt.'S wi th this group beeau Sl' they \\"Cre coverLd earlier under pilf-
asitic associations. An epizoi c assodation is one in whit:h the extlrior of ;1 htl r-
vestman provides an orgcm ism with a substrate for li ving. suc h as <~lg;Jc growing on
the back of harvestmen . Because ex ternal pan1sites arc trc<1t ed undtr par<Jsitic aSSil-
ciations. they arc not treated in the discussion of epizoic tlSSo<."i<J tions.

Phoresy
Some phorctic and parasitic rclcllionships with ht~rws tmen arl' w ry old. Tlw
oldest records tlrc from 40 million years ago. prcserwd in 1\;dtil" amber. Wcitslh;Jt
and Wiclwrd {2002) show a fossi l pscudoscorpion phordk;d]y <lll<ll"ill'd to thllq.tol
a harvestman in ambcr.l.ikewisc. a piece of Baltic amber is known that l"tllltaillS liw
coll cmbohms holding on to the leg of ;1 \wrvestnwn (C. l'oinar. JXrs.rom m. l
Classica l examples or phorctiL: riders of Opilioncs <ll"l' lhl JlSL'UdO.~l"lll"pillllS. This
368 Nmurnl Enemes

~roup of ar;~ehnids is well known to ride <1 v e~rict y of arthropods (Poinar t!l al..
199X l. lntLrcs!ingly. only three: spel'iL~s of chc.:rnctid pseudoscorpions ICiu:/ifer can-
ac>i!lcs. /,mllpmdJ'~"~~'S llot.losus. and l'.~l'laji/Wdwrw.~ tlu/Jius) from Europe have lx.-cn
rcnwtbl fn1111 tlpililllll'S. incl tlding phalangiids and sclerosomatids (W. W. Spicer.
1l'\h/: Smory. I ~ ~X: Vachon . 1(j-J 7: Bdcr. 1()4~: Sankey. 1949b; Cloudsley-
Tillllllpson. 195(,; Spock. ]l)(,-J.: l'oi nar cl ;,I.. I ()9S).lt is dillicult lo understnnd why
thi s is thL case hl'L"ausc L"l\ern ctids and ll<lrvcst mcn arc plentiful in the sa me habitats
i11 tJthcr p;trls tlf till' world. ] >cr he~ps, as sta led by Muchmore ( 1971), "i l is ju s! that
our f<IUIHI h<1 s not been s tudil'd as intensively <IS th<1t of Europe and the <tssociation ,
onurri n ~ spor;~ di c;d ly. h;Js nut yel been ob$Crved by an interesled person ." Gruber
1 I lJlJ ~I <lstl n:ptlrted l"tlll cmbtJians riding on the earth-encrusted surface of some
soil-i nhah i t i n ~ harwstmt.n. Some mites and inscLt larv<te also receive ca rriage. In
tl ll'L"i!Sc ,,r solllL'IlliiL'S. it is not clear if the mit es arc simply riding on th e harvestmen
or actually benl'liting the Jwrwstman by groom ing il.
Hunt 11 Y/9) foun d at least two species of mites on the exoskeleton of the tri-
il eiHJilyrh id IHI!'\'eslnwn l:"tjllilius tlorim from southeastern Australia. While one of
thl s pcril's is pantsitic. l-luntll9 79) stated that the other species "may have a symbi-
otic rel<~tionship with the Jwr\'l'slman ."" Conccntnltions frequ ently occur on the
scutum ncar the orxni ngs of the odoriferous gla nds. and they may feed on non-
\'tllatile rtlmponen ts of their st.-cretions. Presu mably the mit es have some "immunity
to the <ll"ti\'C componen t.
r\ gonyleptid from Argent ina. Pod!!Jioidcllus lmthri. had numerous astigmatid
mlle dcutonymphs locat ed on its chelicerae a nd pcdipa lps lJ. C. Cokendolpher. pers.
obs.). Two species of mites \\"L're observed on the ncmastomatid OrtlwltiSIIIll kvipcs
from the western USt\ ~ J. C. Cokendolpher. pers. obs.). Some of th e mites were the hy-
popal stage of an t\ not:l idae. whereas the sc..<:ond Spt..'"<.'ies remains unidentilled. The
tlnnctids were observed to move about freely on the chelicerae and the mouth area of
the harvestmen. It seems odd that these mites would frequent an area where there
might be food scraps if they did not feed during transport (see also Mi tov. 1 997a:
Fi gure 9.5 1\ ). The sel~o nd species of mite was found on the dorsum of the abdomen.
These appeared to be Hgrazin g"' on the layer of dirt and fungi that is found on the sur-
fan: of many soil-dwelling nemastomatids. It is possible that both of these species
were feeding while in tran sporl and therefore should more properly be con sidered as
a form of cpizoism
Sc\'enl speci mens of Cyplwpl! /lwlmus a rf. leyrovski collected in a Yugoslavian
n tvc bad what appear to be arthropod eggs attached to their appendages (Figure
9.5B). Additional phot ographs show that the end of the structure was open. sug-
gesti ng that the eggs lmd hatched. Further research is needed to determine if this
was an accidental event or the species laying the eggs normally chooses harvestmen
to lay it s eggs u pon for transport Cyplwplulwfmus aff. teyrovski is a small harvestman
!lull is not hi ghly mobile. suggesting that this might have been more of a chance en-
counter.
Opiliones presumably carry many species of bacteria. fungal cleistothecia/spores.
and planl pollen and seeds. which they pick up while crawling through their environ-
Figure 9.5. Epizoics of Opiliones. (A) Phoretic mite Anoetus sp. (Anoetidae) on the anal operculum of the won1d
harvestman Siro sp. {photo: P.G. Mitov). (B) Unidentified arthropod egg on the sironid harvestman Cypl!ophiiJrllmus
all. reyrovski (photo: I.M. Karaman). (C) Fern sporangia (Polysticlwm sp.) epizoically attached to the leg tarsus of the
phalangiid harvestman Leptobunus parvulus (photo: J. C. Cokendolpher). (D) Phol'etk Mallophaga attach('d to thE."
leg tarsus of the sderosomatid harvestman Leiobunum rumelicum (photo: P.G. Mitov)
370 Nauuat Enern1es

llll'lll l i\l;idi<ldo L'l a l. . 1000: l\li tuv. l lJlJl). Much of their grooming removes these
ilt'llls. hlll t111!y <tt"tl-r tlll'Y t r<lllSptlrl tlw1n Stlme distance. Bet:ausc such transport gcn-
L'rtdly l'ill l only lx obst:rwd with ;1 wmpou nd microscope, only a few fern
SJ"ltll'<ln~i<t / spores <IIlli fu n~ill ddstolhct'i;J hilve been rq>orted in the lilcruture to be
carried by hilr\'l'Slrnen tl'okemlolpher. !lJH)b: Figure 9.5C). Phoretic Ma llophaga
haw <Jiso been l'l'l"lmled tFigurL' lJ. 5D).

Endozoism
l lnlikl nwny <lfilchnids. Opilioncs h1ck <I pu mpin g sto mach, and therefore they
dll'w thei r fuud ISL'e L'lmpll'rs 1 <111<1 XI ;:md often consume oocysts. spores. and other
l'Olll<t min tlnt s on their foods. The frequent groomin g of the legs by the harvestmen
may <J iso lead to the ing..stion of otlll'r microorga nisms
,\!though it is unrLportcd in thl literature. there is no doubt thnt ha rvestmen
l"ilfr)' many spt,:des of barteria. yeast. mul fun ga l sporcs as t hey pass through t he di-
gest in system. Exa mination of thl gu t con ten t of many harvestmen reveals the
preseJKl' of a variety of yeasts and fungal spores and hyphae a nd bacteria (J. C. Cok-
cndulpher. pcrs. nbs. I. It is u nknown if thl'SC enter via food (contaminates on foods)
ur <~s li1m!. or if they arl' in gested ucddcntally during groomi ng. Digestion docs not
Sl'em to kill some of theSl' organisms. and thl'Y remai n active nft er being passed in
thc fcccs t ). l' . l'uken dolph cr. pers. obs)

Epizoism
Earlier under t he heHdi ng "" Phorcsy."" collcmbolans and severa l mites were men-
tioned tlwtm ight be more corrcctly listed here. Furthcr study wou ld have to be done
to see if thcsl' uninHl ls were actua lly feed ing. In the cases of photosynthetic
cyanobartcria. <dgae. 11m\ li verworts reported from Opi liones (Gruber. 199 3:
Machado & Vit<Jl. 20t JI ), there is no doubt that these organisms nrc using the sur-
fare of the harvestmen as a substrate for living (sec also Figure 9.SC). Saprophytic
fun gi are a lso observed on harvestmen that are from very humid conditions. The
mil l'S may be bcnclkialto the harvestmen by grooming them. whereas the photo-
synthctir organisms possibly provide camouOage

GENERAL PATTERNS

Bcnmse of the great vmicty in habits ;:md morphologies of Opiliones nnd the diverse
habitats they occupy. it is no wonder that they arc allacked by and associated with a
wide variety of organisms. Most harvestmen share some characteristics that make
them vulnerable to attack by certain enemies. Unlike most arachnids. all harvestmen
masticate their food. ingestin g particles rather thnn using a pumping stomach to cat
liquefied diets. Because of this. they are exposed to parasites and pathogens that
might ot herwise be filt ered out by the feeding mechanism of most other arachnids.
 Natural Enemies 37 1

This is especially true of the gregarines. which are abundan t in harvestmen and un -
common in other arachnids. The omnivorous feeding habits of many harvestmen
place them in proximity to contaminated materials that could result in contact with
pathogens or infective stages of some parasites (sec Chapter 8). Although harvestmen
occur in arid regions. most seem to prefer moister environments (sec Chapter 7).
Living in moist, dark situations provides suitable habitat for exposure to some
pathogens such as fungi. as well as infective stages of some nemat od es. Because of the
activity of some harvestmen. they arc exposed to detection by predators that hunt by
sight or sound . Highly mobile harvestmen also seem to travel more and appear to be
more vulnerable to attack by parasitic mites. Higher mobility is linked to lon ger legs.
so this also results in greater exposu re to aboveground predators and parasites.
Harvestmen a lso share some characteristics that seem to prevent atl<lck by some
enem ies. As already mentioned. a preference for moist habittlls can be detrimenta l.
but this detriment also occurs for other arthropods that might be preda tors or panl-
sitoids of harvest men. This preference can a lso be positive in that such environments
generally offer complex habitats that provide a suitable substmtc for hiding from
some predators. Diurnal huntin g predat ors arc not as successful tlt hunting hcl r-
vestmen as other arthropods because most harvestmen are nocturnc1 l or cryptic
living under coveri ng objects or within the tiller (sec Chapter 101. Whereas some
morphologica l traits such as hardened cu ticle and spines deter some predators. tht
use of exocri ne secretions may have a dual result. Some exocrine Sl't'rel ion s arl' t~n
timicrobial in nature and could help rid the har vestmen of externa l p<Jthogcns <llld
parasites. as well as servi ng as a defensive agen t <J ga inst larger Cit tackers (Sl'C Chapter
10). Fina lly. a ll harvestmen arc free-li vin g Cllld thercli 1re lwvc not hosttd or tr:m s-
mitted the various rickettsial and viral discasts that have dcvclnpl'd in p<~rasi t il"
arachnids {mites and ticks).
Preda tors of harvestmen arc in general the samt as tiHlSe th;1 t at1;1ck spiders. As
wit h other arachn ids. predators of har vest men orten show <t prl'fcrcncc for sizt. o1s
well as time and site of activity. However. many major groups of pa rasites and p<lra-
sitoids of spiders and h arvestmen differ. Crcgarines arc conlllHlllly tncnunt t'fl'd in
harvestm en. but not in spiders (Levine. I IJH5L Ectopar:lsilic mil t'S <l l"l' ronmwn on
ha rvestm en. but much less so on spiders (Sou thl"tllt. I 'JlJ I: l'<,ke ll d{llphcr. llJlJ 3.
Foeli x. 199()). Par<Jsitoids of eggs t~nd t~d u lt spiders. sud 1 <IS IJipll'I"<L ll ymenoptl't"<l.
and Man tispidac. arc unknown li-tm1 Opilioncs. Like scorpion:> !i\kl'ormil"k l'\; l'oli s.
1990). solifu gcs IPunzo. 199H). and spidtrs (1-'oelix. llJlJh). h<~rws t men <l fl' predii -
tors u pon you n ger members of their own spedes. :1s well as ot her splcies of tlll'ir
order (sec Clmpter 81.
The general lack of harvestman parasi toi(ls is inlncslilll! bn"H USl't hty a re rl'la-
tively common in other groups of c1rm:h nids (especia lly spiders!. llarwstnwn tc1dul ts
and eggs) a rc not exploited by wasps. whercc1s spiders <l rl' illlill"kttl by ;1 ~re<ll
num ber of parasitoids of eggs und mlult s. includ in g represl'n1atin-s of the fc11nilil's
lchncumonidac. Sphccidac. l1ompil ith1c. ;1ml Scclionid<IL'. as ll'l'il ns of 1hl' stlpl'l"-
family Chalcidoidca (S haw. JlJlJ4: (;ibson ct ;1l .. 1997: (J" Nl'ill. 2! Ill 1 l. S(llllL' (If tlll'se
372 N;lturat Enelll ies

para~ituids also L'x ploit SU illl' insects. butnotlwrvcstmen. The sa me si tu a lion appar-
l'lltly lk.'l'Urs with till' diptl'ran t'gg predators {Chloropidae. Phoridae. and Ephy-
llridal'\. 'l'lll' rl'asuntsl why harwst mL'Il a rL' not oli cn att;u,;kcd by parasitoids is un -
cert:li n. but ll'l' might SJK't'Ulil tc here. Most lw rvcstmen prefer moist and shady
lwhitats. Possibly thL' pan1silnids prefe r more open conditions, which arc possibly
h~.:tter for hunting and present fewe r pa thogenic fun gi tlwt m ight attack the wasps or
Ilks. ~lust ha rwst men ha\'<' a small body size. This is a n adva nt age that allows its oc-
l'lll'l't'lll'l' in spatia lly rl'stricted soi l a nd litter m i1.: rohabi tats where il is not readily dis-
nrnihk hy many predators ;md p11rasitoids. Possibly the smaller biomass of these
sm;d l species is not en ough fo r full dcvclopmL'Ill of parasitoid larvae. The long legs of
11111st surfacL-act iVL'll<lrwstml'n 1\'tlll ld probably also be an obstacle for wasps to suc-
cessfu lly <I tt ark the hndy or to m;m ipu lat c the host. l'v lost harvestmen arc noct urnal ,
and soml' arc altil'l' durin g otlll'r low-light ev~.: nt s (cloud s. mornings. evenings). Al-
th ough harvcstmL'Il <U'l' capublc of n1pid movclnl!nl. me my wil l move over the land-
SC <IPL' slow ly. possibly not ac ti va tin g the Clllm:k bdwvior of the W<ISps. Possibly the
ansi\'L'r is also in the exocrine glc111d l'lll!ln k al s cmd integumentary morphology
!thick cuticle and hard urm<~mcnt ) of harvcst ll ll!ll. Mor~.:over. many hc1rvcstmcn arc
cryptie. liv ing in Sllil. under stones. and on tree trunks (sec Chapt er 7). Some of them
arc covered with soil frugmcnts tmd so arc well protectl!d (sec Chapter 10). The
eggshells tlf har vcst mc11 arl! th ick. unlikl' th e thin shdl of spiders, a nd better defend
th e embryos. Har\'l'slmcn eggs arc also g~.: n e rally pl<Kt.'<l in a jelly mass or are de-
posed in soils. umkr stoncs. in plant st~.:m s. or in voids such as empty gastro!XJ<I
shells ISL'l' Chapt er l l).

CONCLUDING REMARK S

Studies of enemies and phorct ic and cndozoic/epizoie associations. not mere listing of
new pamsitc or predator rcmrds. constitute prolitable Jiclds for future research. Besides
basic understanding of these interactions { lif~.: cycles and recognit ion of new species of
parasites). studies of the impact of enemies and associates on harvestman individuals
tllld populat ions arc highly desirable. The relative case of laboratory culture of har-
vestmen (sec Chapter I 5) suggests the potential for detailed laboratory studies on para-
sites and pat hog~.:ns. but these would be less in forma tive about predators.

ACKNOWLEDGMENTS

We thank Dr. S. Keller and F. Gschmeidler of Ferdinand Berger und Sohne GMBH
Verlag, Horn, for allowing the reproduction of Figure 9.2A: B. A. Buzatto, J. Warfel.
and Drs. I~ Gnaspini. G. Machado. and J. Spatafora for providing other photographs:
Dr. G. Poinar fo r his photographs. literature. comments, and identi6cations of var-
ious pat hogens and parasites: Dr. I. M. Karama n ror providing Figure 9.58, as well as
new records and literature: J. Owen and Dr. G. Perry for allowing access to Cuban
 Natural Enemies 373

tree frog data and gut contents: Dr. A. A. Giarelta for providing data on the diet of a
leptodactylid frog: Dr. M. Prysby for sending living. parasitized Opiliones that appear
in Figures 9.1A--C: Drs. C. Delchev, A. Gyonova, and T. Lyubomirov for identifying
some spiders. ants. and wasps reported herein: A. Perez Gonzfi lez for identification of
the host in Figure 9.2B: D. Palmer for observations and photographs of mites on Eu-
rybwws harvestmen: Drs. T. Blick. J. Gruber. R. Holmberg. A. KriStin. G. Machado. R.
Madas-Ord6ii.ez. T. Michev. I. Mih<'i.l. P. H. Pache. R. J. Raven, W. Reeves. and \ Z
RUZiCka for information. records. and literature (some very difficult to obtain) on
harvestmen enemies: and Dr. B. P. Nikolov for his great effort in checking over J .000
publications on the foods of birds-his help made our coverage on th is topic very
complete.