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ABSTRACT: Eight major types of umbilical structures can be identified in trochospirally-coiled, latest Cretaceous planktonic
foraminifera. These structures and their variants fall into the broad categories of tegilla and portici. Tegilla have distal secondary
apertures formed by the attachment of tegilla to each other or to the perforate wall around the umbilicus. These distal, secondary
apertures are absent in portici. The various types of tegilla and portici develop from simple lips in early ontogeny and are fundamentally
similar in ultrastructure. Both structures appear to be outgrowths of the chamber wall and are not added secondarilly. Therefore, both
types of umbilical plates are simply elaborate apertural lips. Umbilical covers may function as attachmnent surfaces for rhizopodia over
the umbilicus while simultaneously permitting free communication between chambers in a whorl.
Umbilical structures identify suprageneric groups although convergence is also common. Distinctive portici unite the genera,
Praeglobotruncana, Dicarinella, Marginotruncana and Contusotruncana into a monophyletic group. Globotuncanita and Gansserina
possess derived forms of portici which show that these are valid taxa and may be closely related. Rugoglobigerina, Rugotruncana and
Archaeoglobigerina all carry simple sheet-like tegilla which suggests relationship between these groups. Distinctive elongate tegilla
occur in Archaeoglobigerina cretacea and early globotruncanids, although it cannot be determined whether this reflects convergence
or genetic affinity. Indeed, tegilla with multiple intralaminal apertures have evolved several times as have sheet-like tegilla and those
without intralaminal apertures.
INTRODUCTION
of homeomorphic species such as costellate species of Whitechamber, evidently having been confused by broken structu
inella and Rugoglobigerina (compare Belford 1960 and 1983). (e.g. his plate 260, fig. 6).
Here, I review the umbilical structures found in the majorLongoria and Gamper (1975) presented a classification
groups of Late Cretaceous planktonic foraminifera to assess theumbilical structures, as well as numerous photomicrograp
utility of umbilical structures for phylogenetic classification.
They regarded portici as imperforate, triangular extensions
While umbilical structures have been discussed and figuredthe wall that may fuse into an umbilical cover plate. In th
micropaleontology, vol. 38, no. 2, pp. 165-181, plates 1-7, text-figures 1-2, tables 1-2, 1992
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165
Table 1
Definitions of portici and tegilla and their major variants.
Major variants
Major variants
A) Simple lips--narrow or wide shelf-like structure border- A) Roll-type tegilla--elongate structures fused extensively
ing the primary aperture.
along their distal edges to one another and adjacent
chambers. May have a single intralaminal aperture.
B) Spiral portici--tooth-like structures extending from the B) Strap-shaped tegilla--narrow structures attached at one
primary aperture horizontally over the umbilicus. May or two points distally. May or may not form a single
remain free or may attach distally to other portici.
intralaminal aperture.
C) Imbricate portici--tooth-like structures attached distally C) Multiaperturate tegilla--tegilla forming multiple distal
to other portici whose surfaces dip steeply into the umbi- secondary apertures.
licus.
D) Umbilical sheets--plate or sheet-like structures attachD) Single porticus in adult--a single tooth-like or plate-like ing at many points to the margins of the umbilicus.
porticus.
Both portici and tegilla form two proximal apertures when they
IN UMBILICAL STRUCTURE
truncanella (pl. 2, fig. h). Often, the lip has a serrated edge and
fuse distally with other portici without forming distal secondary
broadens into a flange of roughly uniform width that borders
apertures. A variant includes imperforate to very sparely perfo-
166
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Table 2
Phylogenetic distribution of the main types of umbilical cover plates for taxa examined in this study. Several species may display more than one type
of cover plate. Variability within a genus is also common. References to plates are to illustrations in this paper.
~0
Species
Praeglobotruncana
.D
c'
Figure or Reference
I
I
fit c
stephani (Gandolfi)
Si
t
Li
Hedbergella sp.
1-
gibba Klaus
Dicarinella
Plate 6
concavata (Brotzen)
Plate 6
primitiva (Dalbiez)
II
Marginotruncana
angusticarinata (Gandolfi)
renzi (Gandolfi)
sinuosa (Porthault)
Plate 6
pseudolinneiana Pessagno
Globotruncanita
angulata (Tilev)
calcarata (Cushman)
insignis (Gandolfi)
stuarti (de Lapparent)
stuartiformis (Dalbiez)
I I
X
I:
Plate 1
Plate 1
XIl
I
gansseri (Bolli)
1*1
subspinosa Pessagno
Gansserina
Plate 1
'IX
Contusotruncana
contusa (Cushman)
fornicata (Plummer)
walfischensis (Todd)
IIl
X
I
I
aegyptiaca Nakkady
Plate 2
>(I
arca (Cushman)
bulloides (Volger)
XI
falsostuarti Sigal
linneiana (d'Orbigny)
orientalis El Naggar
'Ix
Globotruncana
Plate 2
I Plate 2
I
I
I
I I IX
Plate 4
X Plate 4
X I
Plate 3
Plate 4
Plate 3
I Plate 4
III I
I
x i
Archaeoglobigerina
blowi Pessagno
cretacea (d'Orbigny)
Plate 3
x I
Plate 3
Plate 3
mateola Huber
Rugoglobigerina
Plate 5
hexacamerata Br6nnimann
Plate 5
rotundata Br6nnimann
Plate 5
rugosa (Plummer)
scotti (Br6nnimann)
Rugotruncana
circumnodifer (Finlay)
X x
Bucherina
Plate 5
citae (Bolli)
havanensis (Voorwijk)
petaloidea (Gandolfi)
Abathomphalus
intermedius (Bolli)
mayaroensis (Bolli)
xxl
Plate 2
Plate 2
Plate 2
Plate 6
Plate 6
167
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A
Primary
Intralaminnl
....
Infralaminal
Comparison of portici (A) and tegilla (B). Numbers refer to successive umbilical cover plates - for instance, structures labeled "2" grew
penultimate chamber in both (A) and (B). Successive portici form only proximal infralaminal apertures between the portici and the perfo
wall, although 'adventitious apertures' may occasionally form in the center of the entire portcullis as in Globotruncanita stuarti (de Lapp
in (A). The tips of portici do not attach to the perforate margins of the umbilicus. Tegilla frequently form multiple distal intralaminal ap
well-formed lips as in Globotruncanafalsostuarti Sigal, shown in (B). The distal margins of tegilla frequently attach to the umbilical wa
fuse distally, but are slightly imbricate and overlap one another
growth of the portici, as the ultimate porticus is often broader
like the steps on a spiral staircase. Each porticus is smooth
thanorthose formed earlier in ontogeny (e.g. pl. 1, fig. d).
weakly pustulose and is not thickened along its margins.
Imbricate spiral portici: Successive portici are strongly imbriSpiral portici: These are essentially the same in structure
and
cate
in some taxa. Instead of being flat-lying in the plane of the
ontogeny as simple portici but with three variations. Spiral
umbilicus, imbricate forms dip very steeply and form a massive
portici are fused at their tips, they may be strongly imbricate,
structure that juts out above the adjacent adumbilical ridges (pl.
and they are occasionally perforate.
1, fig. b). Large individuals may grow a full whorl of large
Plate 1
(DSDP 384-13-6-70-72)
final porticus, (Upper Taylor, near Bristol, Ellis Co.,h, i Gt. atlantica Caron (V34-76-378cm-Amirante PasTexas)
sage, Indian Ocean)
j Gt. calcarata (Cushman) showing typical preservation of imbricate spiral portici (Upper Taylor, near
Bristol, Ellis Co., Texas)
168
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Plate 1
Richard D. Norris
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micropaleontology,
vol.
38,
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no.
169
2,
19
TEXT-FIGURE 2
Comparison of eight types of umbilical structures. A: Simple lip in Hedbergella monmouthensis Olsson-this lip may be modified into
with serrate edge in other taxa; B: Single, tooth-like porticus in Globotruncanella petaloidea (Gandolfi); C: Free spiral portici in Co
contusa (Cushman); D: Imbricate spiral portici in Globotruncanita stuarti (de Lapparent); E: Roll-type tegilla in Globotruncana linneiana
F: Strap-type tegilla in Gt. orientalis El Naggar; G: multiaperturate tegilla in Gt. falsostuarti Sigal; and H: a single, sheet-like tegillum in Rugo
hexacamerata Bronnimann.
portici that do not reach all the way to the center of the
Tegilla
umbilicus. In this case, the portici overlap adjacent portici and Roll-type tegilla: These structures are two to three times longer
leave the center of the umbilicus uncovered (text-fig. la, and than they are wide and assume the shape of a roll or a sausage
pl. 1, fig. g). These central openings are not the same as the (text-fig. 2e). Infralaminal apertures occur at each end of the
intralaminal apertures found in tegilla where the apertures are roll and a short extension near the middle of the tegillum creates
formed by the incomplete fusion of the margins of each tegillum a single, small intralaminal aperture (pl. 3, figs. a, d and f). This
(Compare text-fig. la with Ib). Instead, the 'adventitious apertures' aperture may be filled in by later tegilla or its extension may be
Plate 2
(DSDP 384-13-6-70-72)
(DSDP 384-13-5-66-68).
170
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Plate 2
Richard D. Norris
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1992
171
long but variants that are narrower than they are long are
common (text-fig. 2f). These narrow tegilla often do not form
umbilical area (e.g. pl. 7, fig. d). With further growth the plat
may expand to cover the whole umbilicus and attach at severa
points to form the first tegillum, or remain free as the first
porticus (pl. 7).
usually bordered by a thick collar or rim (e.g. pl. 4, fig. h). The
abundance of apertures means that each tegillum is web-like,
spreading irregularly over the central portion of the umbilicus.
Pores are often sparse, particularly on the earlier formed parts
of the cover plate.
wall and are not added secondarily. This can be seen in juvenile
Umbilical sheets: These include flat or gently undulating single d) and the final porticus of Dicarinella primitava (pl. 6, fig. d)
sheets that cover most or all of the umbilicus (pl. 5, figs. b and In all these cases the umbilical structures merge smoothly with
g). They attach around the margins of the umbilicus leaving a the chamber wall. This suggests that these portici and tegilla ar
series of infralaminal apertures. Intralaminal apertures are few formed simultaneously with the chamber.
(pl. 3, fig. 1). The resulting tegillis nearly completely fills the
umbilicus and is generally sparsely perforate. Umbilical sheets This view is in contrast to that of Banner and Blow (1959), Blow
appear to be partly synonymous with the 'umbilical techo' as (1979) and Banner (1982) who suggest that the porticus is
defined by Longoria and Gamper (1975).
additional to the chamber wall, and is thereby distinquishe
from lips, which are merely extensions of the apertural rim
Some globotruncanids have umbilical structures composed of Blow (1979) shows that portici may overgrow pores on th
multiple overlapping plates here called a compound umbilical chamber testifying to their secondary character. He cites numersheet. Each plate attaches around the margins of the umbilicus ous examples of overgrown pores in Paleogene species and uses
Plate 3
Roll-type and sheet-like tegilla of Globotruncana and Archaeoglobigerina. Scale bars all 100gm.
172
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Richard
D.
Norris
Plate
I m~~~~~~~~~~~~p
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173
even when pores occur they are small and possibly non
tional. Perhaps the scarce pores simply reflect the mech
into the aperture we can see that the umbilical plates are just
sharply reflexed extensions of the aperture lip.
It is likely that tegilla evolved from portici simply by fusing the
distal margin of the first porticus with the opposite side of the
umbilicus. This is illustrated by plate 7, figure e, in which tegilla
living species.
Portici
appear first in Early Cretaceous hedbergellids an
The umbilical structures also permit the development
of wide,
remain relatively
conservative in structure through the Cre
deep umbilical pits without sacrificing sites for attachment
of
ceous.
The
umbilical
structures of Whiteinella, Dicarinella
rhizopodia. In turn, the wide umbilicus allows the test to be
Marginotruncana
and Contusotruncana, are all essentially sim
larger in diameter and flatter without adding substantially
to the
ilar.
genera are united by possessing weakly imbric
overall weight of the skeleton. Direct communication
is These
created
tooth-shaped
between all the chambers, as the apertures all open
into theportici that often fuse distally.
umbilicus beneath the cover plates.
Plate 4
do not form intralaminal apertures (DSDP 313-19-2- the entire umbilicus. The tegillum from the ultimate
12-14)
chamber is broken away (Prairie Bluff Chalk, Braggs,
174
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Richard
D.
Norris
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175
Clearly, umbilical structures do not provide unequivocal evidence of generic relationships. Despite the complexity of these
CONCLUSIONS
tures.
and Archaeoglobigerina cretacea. Whether this similarity reflects convergence or not requires further study. However, the
similarity in umbilical structure together with the variability in
test shape and keel development in globotruncanids, such as Gt.
truncana.
twice.
Plate 5
176
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Richard
D.
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177
REFERENCES
689B and 690C. Proceedings of the Ocean Drilling Program, ScienResults 113:489-513.
BANNER, E T. and BLOW, W. H., 1959. The classificationtific
and
stratigraphic distribution of the Globigerinacea. Palaeontology 2:1-27.
LONGORIA, J. F and GAMPER, M. A., 1975. The classification and
evolution
Cretaceous planktonic foraminifera. Part 1: The superBELFORD, D. J., 1960. Upper Cretaceous foraminifera
fromofthe
Hedbergelloidea.
Revistia Espafola de Micropaleontologia
Toolonga Calcilutite and Gingin Chalk, Western Australia.family
Bureau
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Numero
Especial, enero pp. 61-96.
Mineral Resources, Geology and Geophysics, Bulletin 57,
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and stratigraphy of the Corsicana Formation (Maestrichtian) NorthS., 1977. Test Morphology, organic layers and chamber formation of
Central Texas. Cushman Foundation for Foraminiferal Research
the planktonic foraminifer Globorotalia menardii (d'Orbigny). Jour-
Plate 6
Spiral portici of Dicarinella and Marginotruncana as well as tegilla of Abathomphalus. Scale bars all 100lm.
(DSDP 305-19-6-37-39)
DSDP 305-26-4-39-41
178
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Richard
D.
Norris
Plate
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vol.
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Plate 7
Umbilical structures of juvenile planktonic foraminifera. Scale bars 100l m except for (b), (d) and (f) which are 50glm.
c, d Globotruncana aegyptiaca Nakkady with initial porticus just before growth of the first tegillum (Prairie
Bluff Chalk, Braggs K-T boundary section, Lowndes
Co., Alabama)
f, g Globotruncana sp. (possibly Gt. rugosa (Marie) before keels are well developed) (Prairie Bluff Chalk,
Braggs K-T boundary section, Lowndes Co., Alabama)
180
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Richard
D.
Norris
Plate
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micropaleontology,
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