1998 Contr. Tert. Quatern. Geol. 35(1-4) 63-83 37 figs, 1 tab. Leiden, April
Bryozoans from the Pliocene Bowden shell bed of
Jamaica
Paul D. Taylor
AND
Tiffany S. Foster
THE NATURAL HISTORY MUSEUM
LONDON, ENGLAND
Taylor, Paul D. & Tiffany S. Foster. Bryozoans from the Pliocene Bowden shell bed of Jamaica. In: Donovan, S.K. (ed.). The Pliocene
Bowden shell bed, southeast Jamaica.— Contr. Tert. Quatern. Geol.,35(l-4):63-83, 37 figs, 1 tab., Leiden, April 1998.
all of the Recorded for the first Nineteen species of bryozoans are illustrated from the Bowden shell bed, including commoner species. Bassler, 1928, Schizo- time from the Bowden shell bed are ?Plagioecia dispar Canu & Mecynoecia proboscideoides (Smitt, 1872), porella errata (Waters, 1879),Petraliellacf. bisinuata (Smitt, 1873) andSchedocleidochasma porcellanum (Busk, 1860). Most of the
Bowden bryozoans are widespread in the Caribbean and Gulf of Mexico in the Neogene and at the present day. They represent a tropi-
flows. cal shelf fauna, apparently transported without appreciable abrasion into a deeperwater setting in sediment gravity
Key words — Bowden shell bed, systematics, Cyclostomata, Cheilostomata, SEM, ecology.
P.D. Taylor and T.S. Foster, Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, Eng-
land.
Contents tained in Terebripora and the latter being placed in syn-
for onymy with Orbignyopora archiaci (Fischer). Except
Pohowsky (1978), all of the papers describing Bowden Introduction p. 63 bryozoans predate scanning electron microscopy (SEM). Methodsand materials p. 64 The availability of SEM has had a major impact on bryo- Systematic palaeontology p. 64 zoan systematics, especially in permitting easier and Discussion p. 70 more accurate illustrationof morphological details. Acknowledgements p. 71 intended be The present paper is not to a mono- References p. 71 graphic revision of the Bowden bryozoan species: such
of all of an undertaking would require not only restudy
the but also extensive with related types, comparison taxa Introduction of diverse geographical and stratigraphical provenance.
Instead, we aim to provide a short synopsis concentrating
first describe Canu & Bassler (1919) were the to bryozo- on the most common species, the majority of which are
ans from the Bowden Formation (Bowden Marl or Bow- here illustrated for the first time using SEM. Conse-
den shell of Jamaica. described 17 9 and bed) They species, quently, full species descriptions synonymies are not
of these characteristics of which were new. Subsequently, many species given. However, the most important for
redescribed Canu & Bassler the Bowden were by (1923), reproducing distinguishing between commoner species
of the used in their earlier their taxonomic nomenclature and classifi- many photographs they paper, are outlined,
and the of the fauna increased 32 diversity was to spe- cation are updated, and some ecological comments are
cies. Only one paper specifically concerning Bowden given.
bryozoans has been published subsequently, in which The Bowden Formation is part of the Lower Coastal
described further six Canu & Lagaaij (1959) a species. Group and outcrops in a small area of southeastern Ja- of Bassler's (1923) two Bowden species ctenostome maica, around Bowden on the eastern side of Port Mo-
bryozoans represented by borings, Terebripora sinefilum rant Bay. Over 600 species of invertebrates have been
included in the so-called 'Bowden and T. elongata, were Pohowsky's (1978) recorded from the lowermost unit,
monograph of ctenostome borings, the former being re- shell bed', making it the most famous fossiliferous hori- 64
in zon Jamaica. The shell bed comprises a series of from North Carolina as likely to be C. denticulata coarse-grained and conglomeratic layers and lenses (Lamarck). Maturo's figure (1957, fig. 21) shows subcir- within marlstones. Trace fossil and some other evidence cular pseudopores like those of the Bowden shell bed
that the Formation is suggests Bowden a deep-water de- specimen. However, it is impossible to be sure of the exact
with the posit, fossils in the shell bed being allochthonous identity of the Bowden Crisia, which is therefore not as- and from sediment flow deposited a gravity (Pickerill et signed to a species.
al., The Bowden — 1996). age of the shell bed is late Plio- Ecology Species of Crisia have a wide bathymetric cene. distribution; for example, Hayward & Ryland (1985) re-
corded C. denticulatafrom the lower shore, whereas other
of the described Harmelin species genus by (1990) range
down to almost 1,000 m. METHODS AND MATERIALS
Specimens were studied using an ISI ABT-55 scanning Genus Crisulipora Robertson, 1910 electron microscope equipped with an environmental chamber. The micrographs reproduced in this paper are
Remarks — Crisulipora resembles Crisia, but the articu- backscattered electron images of uncoated specimens. lated used in the internodes have between 2 and 5 transverse rows of Specimens present study are in the collec- and the is tions of The Natural History Museum, London (BMNH) autozooids, gonozooid a more complex structure penetrated by autozooidal and the Florida Museum of Natural History, University apertures. of Florida, Gainesville (UF).
Crisulipora sp.
Fig. 7 SYSTEMATIC PALAEONTOLOGY
cf. 1910 Crisulipora occidentalis Robertson, p. 254, pi. 21, Order Cyclostomatida Busk, 1852 figs 22-24. Suborder Articulina Busk, 1859
cf. Robertson — 1937 Crisulipora occidentalis Marcus, p. 21, Crisiidae Johnston, 1847 Family pi. 3, fig. 5. Genus Crisia Lamouroux, 1812 1959 Robertson — Crisulipora occidentalis Lagaaij, p.
484, text-fig. 5.
— Robertson — Soule et Remarks Colonies of Crisia are jointed and erect ('cel- cf. 1995 Crisulipora occidentalis al., p. lariiform'), with internodes comprising biserially-arranged 309, pi. 119, figs A-D. autozooids separated by elastic articulations. Fossil colo-
nies are invariably disarticulated. Gonozooids are bulbous Material— BMNH D41290-41317.
structures situated fixed — usually at a position in the inter- Remarks None of the Bowden specimens possess gono- node. zooids and therefore it is difficult to be certain of their
identity. Recent Crisulipora occidentalis was originally
described from the Pacific coast of North America, but
Crisia sp. subsequently recorded from Brazil and Japan. Soule et al.
(1995) suggested that C. occidentalis was either trans-
cf. 1838 2. into the Atlantic Crisia elongata Milne-Edwards, p. 203, pi. 7, fig. ported as a fouling organism, or that more
1959 Crisia Milne-Edwards — than exists. be excluded in elongata Lagaaij, p. 484, one species Clearly, fouling can
text-fig. 4. the case of the Bowden fossil material. Study of material
cf. 1986 Crisia Milne-Edwards — Winston, 30, elongata p. from various localities is required to resolve the systemat- figs 71-73. ics of Crisulipora and we prefer not to assign the Bowden ,
specimens to a species until this is done. However, the
Material— BMNH D41249. internodes of the material narrow Bowden prompt a tenta-
Remarks — internode Only one of Crisia is known from Crisuliporative comparison with the Japanese species the Bowden shell bed. This has been well which already figured ijimai was described, but not figured, by Okada by Lagaaij (1959, text-fig. 4) as Crisia elongata Milne- (1917).
described from the — Edwards, 1838, a species originally Ecology Crisulipora occidentalis can be found from
Recent of the Red Sea. SEM examination of the Bowden low tide to 86 m (Soule et al., 1995). specimen revealed subcircular pseudopores, whereas a specimen from Panama figured by Winston (1986, figs 71-
73) as C. elongata has slit-like pseudopores. Cook (1985, Suborder Tubuliporina Milne-Edwards, 1838 p. 202) regarded the C. elongata of Maturo (1957, p. 31) Family Diastoporidae Gregory, 1899 65
Genus Plagioecia Canu, 1918 riously variable in vinculariiform cyclostomes (cf. Flor,
1972) and the two species described by Smitt are provi-
— is Remarks Plagioecia a multiserial, sheet-like, typi- sionally regarded as synonyms.
— Recorded from 26-110 in Florida and cally encrusting tubuliporine in which the gonozooid is Ecology m depth
transversely elongate and pierced by autozooidalapertures. Panama.
?Plagioecia dispar Canu & Bassler, 1928 Order CheilostomatidaBusk, 1852
Fig- 1 Suborder Malacostegina Levinsen, 1902
Family Membraniporidae Busk, 1852
Genus Biflustra d'Orbigny, 1852 cf. 1928 Plagioecia dispar Canu & Bassler, p. 159, pi. 31, fig.
10.
— to moder- — Remarks Zooids of have cf. 1982 Plagioecia dispar Canu & Bassler Winston, p. Biflustra slightly
155, fig. 94. ately developed cryptocysts, and the gymnocyst is absent
tubercles. In or present only as well-preserved zooids,
denticles be Ovicells and Material— BMNH D41289. cryptocystal may developed.
avicularia are The ancestrula seen in fos- Remarks— The single Bowden specimen is detached from lacking. (rarely
is in unlike the ara- its substratum and is infertile. Therefore, its identification sils) twinned, as Membranipora, but,
the skeleton of is cal- as Plagioecia can only be tentative. Neither P. dispar nor gonitic Membranipora, Biflustra
similar have been recorded from cific (Taylor & Monks, 1997). Colony form is variable any species previously both within the and within of Bowden; P. dispar was originally described from the Re- genus species Biflustra.
Some colonies are entirely encrusting, whereas others cent of the Straits of Florida, but Winston (1982) noted its
far north Hatteras. develop erect growth commonly in the form of tubular presence as as Cape
branches or or bifoliate Ecology — Winston's (1982) material of P. dispar came ('cavariiform' 'hemescharan') fronds. from 60-90 m depth, and Canu & Bassler's (1928) from
102 m.
Biflustra monilifera (Canu & Bassler, 1919)
2-4 Family Mecynoeciidae Canu, 1918 Figs
Genus Mecynoecia Canu, 1918
? 1873 Biflustra savartii Audouin — Smitt, p. 20, pi. 4,
Remarks —Mecynoecia has erect, narrow branches ('vin- figs 92-95. forma Canu & culariiform') and simple bulbous gonozooids. 1919 Acanthodesia savartii monilifera
Bassler, p. 79, pi. 2, figs 2, 3.
1919 Acanthodesia savartii forma texturata Reuss — Mecynoecia proboscideoides (Smitt, 1872) Canu & 1-5. Bassler, p. 79, pi. 5, figs Figs 8, 9 1923 Acanthodesia savarti forma monilifera Canu &
3. Bassler — Canu & Bassler, p. 32, pi. 2, figs 2,
1872 — Smitt, 11, 5, Entalophora deflexa (Couch) p. pi. 1923 Acanthodesia savarti forma texturata Canu &
28-30. figs Canu & Bassler— Bassler, p. 32, pi. 5, figs 1-5; pi. 1872 Entalophora proboscideoides Smitt, 11, pi. 4, figs p. 46, figs 8, 9. 26, 27.
1928 Smitt— Canu & Mecynoecia deflexa Bassler, p. 160,
— D41113 pi. 31, fig. 1. Material UF 75953-4, BMNH D34301-5,
1928 Entalophora proboscideoides Smitt — Canu & (sample).
34, 11. — revision Bassler, p. 160, pi. fig. Remarks The systematics of Biflustra require
and our assignment of this species to B. monilifera is
Material— UF 75951-52. provisional. The Bowden species, which has tubular
Remarks — Smitt (1872) described two similar species branches, differs from B. savartii, described originally
from the Recent of Florida; Entalophora proboscideoides from the Recent of Egypt, in having a narrower proximal
& arbo- and E. deflexa. The former was described as a new species, cryptocyst (see Taylor Foster, 1994). Biflustra
Canu & 1928 is another similar but the latter is evidently a re-attribution of Tubulipora rescens Bassler, species,
North Africa and the of deflexa Couch, 1842, a European species now assigned to but this form, from Bay Biscay,
bilamellar colonies Our material Entalophoroecia (see Hayward & Ryland, 1985, p. 113), has (Alvarez, 1990).
shows tubercles which are not mentionedor which differs from the Floridan species in having a more gymnocystal
complex gonozooid. Both of Smitt's putative species have figured by Smitt (1873) in his description of supposed B.
from the Recent of Florida. The of Bi- autozooids of about the same size, but his E. deflexa has savartii identity
is it is slightly broader branches. However, branch width is noto- flustra texturata (Reuss, 1848) unclear, although 66
thought unlikely that this species from the Miocene of Remarks — Nellia is characterised by erect articulated
the Paratethys is conspecific with the Bowden specimens. colonies('cellariiform') with quadriserial internodes which
Canu & Bassler had of B. fossilisation. Autozooids (1919) only a single specimen invariably become separated on
from the Miocene 'Bowden horizon' of Santo have shelf-like and monilifera, a proximal cryptocyst a narrow gyn-
the Dominigo (Dominican Republic), but the tubular colony- mocyst with a pair of small avicularia near proximo-
and existence of tubercles lateral of the Ovicells incon- form gymnocystal suggests corners gymnocyst. are very
that this may be the most appropriate name for the abun- spicuous.
dant material ofBiflustra found in the Bowden shell bed
of Jamaica. Nellia tenella (Lamarck, 1816)
Ecology — Tubular colonies of Biflustra grow around Fig. 16
algae or hydroids (see, for example, Cook, 1968, p. 123), but not to these and do 1816 tenella 135. are closely adpressed organisms Cellaria Lamarck, p.
Nellia oculata not make an impression (substrate bioimmuration) of 1852 Busk, p. 18, pi. 64, fig. 6; pi. 65 (bis),
them on the colony underside. Smitt's (1873) tubular fig. 4.
1919 Nellia Busk — Canu & colonies of ‘B. savartii’ from Florida were dredged at a oculata Bassler, p. 82, pi. 2,
figs 5-7. depth of 53 m.
1923 Nellia oculata Busk — & Canu Bassler p. 55, pi. 2,
figs 5-7.
1959 Busk — 1. Nellia oculata Lagaaij, p. 482, text-fig. Suborder Neocheilostomatinad'Hondt, 1985
1966 Nellia tenella (Lamarck) — Cheetham, p. 48, text- Family Calloporidae Norman, 1903 fig. 28. Genus Antropora Norman, 1903
1969 Nellia oculata — 2. Busk Lagaaij, p. 167, fig.
1984 tenella — Nellia (Lamarck) Winston & Cheetham, p.
Remarks — Antropora has encrusting colonies, sometimes 257, figs 1, 2. multilamellar, with autozooids having well-developed
endozooecial cryptocysts, small ovicells, and usually inter- Material— BMNH D41127-D41206. zooecialavicularia (see Osburn, 1950; Cook, 1985).
Remarks — Nellia oculata is regarded as a junior synonym
of N. tenella (Cheetham, 1966). This species has been la-
belled as a living fossil, ranging from the Maastrichtian to Antropora parvicapitatum (Canu & Bassler, 1923) the Recent (Winston & Cheetham, 1984). At the present Figs 5, 6 day N. tenella is widely distributed throughout the tropics
and subtropics (Lagaaij, 1959). 1923 Canu & Membrendoecium parvicapitatum Bassler, p. Ecology — Rucker (1967) found this species to be abun- 36, pi. 12, figs 1, 2. dant on the outer shelf of Venezuela-British Guiana,
mainly in calcareous sand facies. Although the species can
Material — UF BMNH D34282. 75955, at be found in shallow water (less than 4 m), it may occur
Remarks — Although the type specimen of this species depths of up to 1,000 m, and Winston & Cheetham (1984)
comes from the Miocene of Florida, Canu & Bassler's noted that palaeoecological interpretation must take into mentions material from the (1923) original description account the possibility of allochthonous deposition.
Bowden shell bed, and the BMNH collections contain a
Bowden specimen identified and donated by R.S. Bassler.
The has and variable species a pustulose cryptocyst a Family Cupuladriidae Lagaaij, 1952
proximal which include a single median gymnocyst may Genus Cupuladria Canu & Bassler, 1919 tubercle. Several zooids show partial closure plates incor-
porating the operculum. These resemble similar structures Remarks — Cupuladria is a free-living, so-called lunulite
‘Membraniporadescribed by Lagaaij (1963, p. 165) from or 'lunulitiform' bryozoan. The colony has the shape of a
tenuissima’. Small polymorphic zooids are found between Chinese hat, with the zooids opening on the upper convex
the autozooids and to be kenozooids rather than appear surface. Distal to each autozooid is an avicularium (or vi- avicularia. The ovicell is small. Generic is very placement braculum). The underside of the colony is divided into a
uncertain; Membrendoecium has been regarded as a junior series of roughly rectangular sectors with pores.
synonym of Antropora (Bassler, 1953, p. G160), but the
absence of avicularia in the Bowden calls apparent species Cupuladria biporosa Canu & Bassler, 1923 into its question assignment toAntropora. Figs 10-13
— Canu & 1919 Cupuladria canariensis Busk Bassler, p. Quadricellariidae Gordon, 1984 Family 78 (?partim).
Genus Nellia Busk, 1852 1923 Canu & Cupuladria biporosa Bassler, p. 29, pi. 47, 67
Venezuela-British Guiana. Tommasi al. found figs 1,2. of et (1972)
Canu & Bassler — Cook, in the Ilha Grande of Bra- 1965b Cupuladria biporosa p. it at depths of 11-150 m region figs 2A, B, 3A, B, 4A, B, 5, 6A, B; text- 203, pi. 1, zil, where it is particularly common in very fine sand. fig- lg-j-
1994 Cupuladria biporosa Canu & Bassler — Cook &
Chimonides, p. 260, figs 6, 8, 9,11,12,14. Family Candidae d'Orbigny, 1851
Genus Canda Lamouroux, 1816
Material — UF 75956-8, BMNH D34293-6, D41117
(sample). colonies with Remarks — Canda has thinly-calcified, erect
— confused the Remarks Cupuladria biporosa has been in zooids arranged in two series facing obliquely outwards vicari- past with C. canariensis, but the morphology of the median keel and from a having well-developed cryptocysts ous avicularium is different: those of C. biporosa have (Gordon, 1984). Cook & Chi- more extensive proximal gymnocysts (see
described from the monides, 1994, figs 7, 8). Originally Canda caraibica Levinsen, 1909
'Bowden' of the Dominican Republic, the holotype of C.
was Cook & Chimonides biporosa refigured by (1994, fig. Canda caraibica 142. 1909 Levinsen, p. Cook 1, figs 5, 6) figured material of this Levinsen — 9). (1965b, pi. 1959 Canda caraibica Lagaaij, p. 483, text-
from the Bowden shell bed of Jamaica. The name species fig. 3a, b. biporosa is somewhat inappropriate because, as shown by
Cadee (1981, table 2), there are usually more than two Material — BMNH D41207.
basal sector.
pores per — small of this is Remarks Only one fragment species
— This Miocene to Recent species from Ecology ranges known from the Bowden shell bed. This specimen has
13-150 m depth at the day. On the Venezuela- present been adequately illustrated by Lagaaij (1959). British Guiana shelf it is very abundant throughout the Ecology — Lagaaij (1959) referred to this as a shallow- sand facies of the outer shelf calcareous (Rucker, 1967), from 22 while water form, citing two occurrences m depth, while in the Ilha Grande region of Brazil it prefers sandy Osburn (1914) recorded specimens from 17 m in Florida. bottoms deeper than 40 m (Tommasi et al., 1972).
Family Steginoporellidae Hincks, 1884 Genus Discoporella d'Orbigny, 1852 Genus Steginoporella Smitt, 1873
Remarks —Discoporella resembles the related Cupula- Remarks —Steginoporella is characterised by having two
dria, but the zooids have a more extensive frontal shield, and B-zooids. The B-zooids have types of autozooids, A- including a well-developed vestibular arch, and vicarious enlarged mandibleswhich are reflected in the larger size of avicularia are absent (Cook & Chimonides, 1994). the opesium and sometimes in a shelf-like rostrum. The
cryptocyst is well developed and has a median process. Discoporella umbellata(Defrance, 1823) There is no gymnocyst and ovicells are lacking. Fig. 14
Steginoporella parvicella Canu & Bassler, 1919
1823 Lunulites umbellata Defrance, p. 361, pi. 47, figs 1, Fig. 15
la, lb.
1919 Cupularia umbellata (Defrance) — Canu & Bassler, Canu & 1919 Steganoporella parvicella Bassler, p. 89, pi. 17-21. p. 85, pi. 1, figs 5-7; pi. 2, figs 6, figs 6-9. 1923 Cupularia umbellata (Defrance) — Canu & Bassler, 1923 Steganoporella parvicella Canu & Bassler — Canu 15-19. p. 80, pi. 2, figs & 6-9. Bassler, p. 62, pi. 6, figs 1965a Discoporella umbellata (Defrance) — Cook, p. 177, 1979a Steginoporella parvicella Canu & Bassler — Pouyet pi. 1, fig. 7; pi. 3, figs 1, 3, 5, 6; text-fig. 4. & 3. David, p. 789, text-fig. 1985 Discoporella umbellata (Defrance) — Cook, p. 93,
pis 4C, 5A, 5B.
Material— UF 75959, BMNH D34292, D41124 (sample).
— from the Material — BMNH D34286-9, D41118 (sample), BZ Remarks The only species of Steginoporella
3504. Bowden shell bed has unilamellar colonies and relatively
— small zooids & Remarks D. umbellataas presently understood is widely (Pouyet David, 1979a).
— is but the distributed geographically and long-ranging. Cook (1985) Ecology Steginoporella parvicella extinct,
related S. occurs from low regarded it as a species complex. magnilabris (Busk) commonly
— 27 in Florida and be found Ecology Rucker (1967) noted that D. umbellata is water to m (Osburn, 1914), can
abundant in the calcareous sand facies along the outer shelf down to almost 70 m. With one exception, Steginoporella 68
is tropical or subtropical (Pouyet & David, 1979b). erect, bifoliate genus characterised by marginal areolae,
suboral avicularia, and often vicarious and small adventi-
tious avicularia (see Lidgard, 1996). Family Thalamoporellidae Levinsen, 1909
Genus 1887 Thalamoporella Hincks, Adeonellopsis deformis (Canu & Bassler, 1919)
Figs 22, 23
Remarks — Caliper-compass or bow-shaped calcareous
are within the coelomic of zooids in spicules present cavity 1919 Canu & Bracebridgia deformis Bassler, p. 97, pi. 3, Thalamoporella but are not often found in fossils. Auto- , figs 11-16. zooids have extensive cryptocysts pierced by opesiules. 1923 Bracebridgia deformis Canu & Bassler — Canu &
Ovicells and avicularia tend to be and 11-16. are bilobate, large Bassler, p. 160, pi. 8, figs
in & — acuminate or spatulate. pr. Adeonellopsis deformis (Canu Bassler)
Cheetham et al.
Thalamoporella biperforata Canu& Bassler, 1919
Figs 18-20 Material — UF 75963-4, BMNH D34290-1, D41115 (10
specimens).
1919 Canu & Remarks — As with other of Thalamoporella biperforata Bassler, p. 88, many species Adeonellopsis
pi. 6, figs 10-15. there is considerable frontal thickening which obscures the
1923 Thalamoporella biperforata Canu & Bassler -— Canu primary orifice and adventitious avicularia in older
& Bassler, 62, 6, 10-15. p. pi. figs branches. Colonies comprise narrow bifoliate branches
1992 Canu & Bassler — Thalamoporella biperforata ('adeoniform'). 57-59. Soule et al., p. 42, figs
Material — UF BMNH D41126 75960-1, D34297, (sam- Family Lepraliellidae Vigneaux, 1949 ple). Genus Celleporaria Lamouroux, 1821
Remarks — The erect, bifoliate colonies of T. biperforata are sometimes with the two preserved layers separated. Remarks — Celleporaria typically has massive colonies Avicularia and have Soule are long two large opesiules. et formed by frontal budding. Autozooidal orifices are usu- al. able find (1992) were to compass-shaped spicules only. ally non-sinuate (but see below) and ovicells are imperfo- Ovicells were not seen in our material and Soule et al. rate (Gordon, 1984).
(1992) noted only the beginnings of an ovicell in one specimen.
Celleporaria? hemispherica (Canu & Bassler, 1923)
Figs 24-26
Thalamoporella chubbi Lagaaij, 1959
21 Figs 17, 1923 Canu & Holoporella hemispherica Bassler, p. 176,
pi. 3, figs 9,10.
1959 chubbi Thalamoporella Lagaaij, p. 483, text-fig. 2a-c.
chubbi — Soule 1992 Thalamoporella Lagaaij et al., p. 69,
Material— UF BMNH D34283-5. figs 99,100. 75965-6,
Remarks — This species has small hemispherical colonies
formed by piles of frontally budded zooids. Generic attri- Material — UF 75962, BMNH D41208 (holotype), bution of such celleporiform colonies is often difficult in D41209-47 (paratypes).
fossil material because the diagnostic ovicells are rarely Remarks — Thalamoporella chubbi differs from T. biper-
preserved, as in the Bowden species. Although Gordon forata in having larger zooids, tubular, 'cavariiform' colo-
(1984, 115) described Celleporaria as having a non- nies (like those of Biflustra monilifera), and avicularia p. sinuate sinus does orifice, a (or notch) exist in some spe- without opesiules. Unlike T. biperforata, ovicells are very
cies which are traditionally assigned to this genus. Notably common in T. chubbi. The species is known only from the C. brunnea Winston Bowden shell bed of Jamaica. (Hincks), as figured by (1986, fig.
20), has a shallow but distinct sinus and an overall orifice
shape which is very similar to the Bowden species. Conse-
the Bowden Suborder Ascophorina Levinsen, 1909 quently, we assign species questionably to is Infraorder Umbonulomorpha Gordon, 1989 Celleporaria (Holoporella generally regarded as a junior of Family Adeonidae Busk, 1884 synonym Celleporaria ; see Harmer, 1957, p. 664). ? lacks the abundant Genus Adeonellopsis MacGillivray, 1886 Celleporaria hemispherica spatu-
late, interzooecial avicularia found in C. brunnea (see, for
Remarks — Soule Adeonellopsis is an encrusting or more often example, et al., 1995, pi. 101, fig. A). 69
Remarks — This was thoroughly redescribed by Family Cheiloporinidae Bassler, 1953 species Bowden and has Genus Hippaliosina Canu, 1918 Cheetham (1968) based on material from
since been depicted using SEM (Cheetham, 1986). Metra-
autozooids with from the late Miocene (NN Remarks — This encrusting genus has rabdotos lacrymosum ranges
Pliocene granular frontal walls and marginal areolae, an elongate 11) to the late (NN 16) (Cheetham, 1986). orifice and lateralavicularia.
1989 ? Hippaliosina baccata (Canu & Bassler, 1920) Infraorder Lepraliomorpha Gordon, 1883 Fig. 27 Family Schizoporellidae Jullien,
Genus Schizoporella Hincks, 1877
1920 Hippodiplosia baccata Canu & Bassler, p. 397, pi.
Remarks — is and diverse 87, figs 5, 6. Schizoporella a common genus
— & The 1923 Hippodiplosia baccata Canu & Bassler Canu with encrusting, often multilamellar colonies. auto-
3, 1. zooids have frontal orifice with Bassler, p. 131, pi. fig. evenly perforated walls, an
adventitious a distinct proximal sinus, and single or paired
avicularia lateralto the orifice. Material — BMNH D34298-300, D41120.
Remarks — The Bowden material of ? H. baccata consists
errata (Waters, 1879) of unilamellar encrusting colonies with autozooids having Schizoporella
the frontal wall. Ad- 32 a conspicuous granular ornament on Fig.
ventitious avicularia are located singly to one side of the autozooidal orifice. The orifice is not well preserved in the 1879 Lepralia errata, stadiumHemeschara Waters, p. 39,
inferred scanned material, but its probable shape can be by pi. 10, fig. 5.
— & comparison with the related species H. rostrigera (Smitt) 1979 Schizoporella errata (Waters) Hayward Ry-
68. land, p. 170, fig. asHippaliosinadepicted by Winston (1982, fig. 65). ?
baccata differs from this latter species in having more
rounded avicularia. Material — UF 75969.
described from the Lower Remarks — the Hayward & Ryland This species was originally Using key given by the Bowden is apparently S. Miocene of Mississippi and later identified from the Bow- (1979, p. 167), Schizoporella
den shell bed of Jamaica. The Mississippi material figured errata (Waters). This is a widespread warm temperate-
cal- at the Some of the modern by Canu & Bassler (1920) appears to be more thickly subtropical species present day.
cified around the orifice, and the Bowden material may material described by Winston (1984) as S. ?serialis(Hel-
from Belize be the perhaps represent a different species. ler, 1867) may same species.
Family MetrarabdotosidaeVigneaux, 1949 Family Petraliellidae Harmer, 1957
Genus MetrarabdotosCanu, 1914 Genus Petraliella Canu & Bassler, 1927
Remarks — In most species of Metrarabdotos, colonies Remarks — Petraliellatypically has large autozooids with
frontal walls and orifice indented are bifoliate ('adeoniform' or 'eschariform'), although a evenly perforate a large
The basal surface is smooth few are encrusting. The elongate autozooids have large by two or more small sinuses.
rise rootlets in colonies marginal areolae and there are polymorphic gonozooids except for pores that give to living
with costulate ovicells. (Harmer, 1957).
Metrarabdotos lacrymosum Canu & Bassler, 1919 Petraliella cf. bisinuata(Smitt, 1873)
Figs 28, 29 Fig. 33
bisinuata 229. 1919 Metrarabdotos lacrymosum Canu & Bassler, p. 96, cf. 1873 Escharella Smitt, p. 59, pi. 12, fig.
& — pi. 3, figs 1-10. ? 1967 Hippopetraliella marginata (Canu Bassler)
Canu & Bassler — Canu 14h. 1923 Metrarabdotoslacrymosum Rucker, p. 826, fig.
& 164, 8, 1-10. Bassler, p. pi. figs
1968 Metrarabdotos (Biavicularium) lacrymosum Canu &
Material — UF 75970.
— 1. Bassler Cheetham, p. 87, pi. 9, fig. 5; pi. 12, fig.
Remarks — The Bowden material differs from Recent P. 1986 Metrarabdotos lacrymosum Canu & Bassler — orifice bisinuata in having a proportionally larger Cheetham,figs 6C, F, 7C. (Smitt) (cf. Canu & Bassler, 1928, pi. 16, fig.l), and more closely
resembles Rucker's (1967) figure of Hippopetraliella mar- Material — UF 75967-8, BMNH D34309-12, D41123 (3 ginata (Canu & Bassler). However, Canu & Bassler (1928) pieces). 70
described the orifice of this latter species as having a the calcareous sand facies of the outer Venezuela-British
straight proximal border (the serrations in Canu & Guiana shelf.
Bassler's due figures are to retouching; A.H. Cheetham,
pers. comm., December 1996), whereas the Bowden mate-
when well shows rial, preserved, two small sinuses. Family Phidoloporidae Gabb & Horn, 1862
— The from the Gulf of Ecology petraliellids Mexico re- Genus Schedocleidochasma Soule et al., 1991
corded Canu & Bassler by (1928) have a depth range of
14-55 m. to Harmer Remarks — Colonies of According (1957, p. 693) petraliellid Schedocleidochasma are encrust-
colonies '...often which their with loosely encrust Sponges, to ing autozooids having mamillate frontal walls and a rootlets attached. Tubular are Hemescharan colonies, as few marginal pores. The orifice is keyhole shaped. Avicu-
found in laria short and Hippopetraliella magna, may commence as an are acute, and ovicells globose and imperfo- encrustation and be hollow prolonged as tubes beyond the rate (Soule et al., 1991). attached portion'.
Schedocleidochasma porcellanum (Busk, 1860)
Fig. 36
Family Mamilloporidae Canu & Bassler, 1927
Genus Smitt, 1873 Mamillopora 1860 3. Lepralia porcellana Busk, p. 283, pi. 31, fig.
1873 Lepralia cleidostoma Smitt, p. 62, pi. 11, figs 217-
Remarks — Like Cupuladria, the 'lunulitiform' coloniesof 219.
Mamillopora resemble a Chinese hat in shape, with the 1928 Hippoporinacleidostoma (Smitt) — Canu & Bassler,
autozooids 104, pi. 9, pi. 32, fig. 18. opening on the upper convex surface. The large p. fig. 7; 5; text-fig.
1964 Cleidochasma — Cook, 11, orifice has paired condyles, and is surrounded by a strongly porcellanum (Busk) p.
4A-E. frontal wall which ad- pi. 1, fig. 4; pi. 2, figs 1, 2; text-figs pustulose cryptocystal may support 1982 Cleidochasma porcellanum — Winston, ventitious avicularia. The mamillate underside of the col- (Busk) p. 147, 80. contains fig. ony large chambers which in living colonies rep-
1991 Schedocleidochasma porcellanum (Busk) — Soule et resent the points of origin of rootlets for anchoring the col- 481. al., p. ony into the sediment.
Material— UF 75973.
Remarks — The Bowden fossil retains the porcellanous Mamillopora tuberosa (Canu & Bassler, 1919) appearance from which this extant species takes its name. Figs 30, 31, 34, 35, 37
— This Ecology species has a circumtropical distribution
in the Atlantic at the present day. Depth ranges from 9- ? 1873 Mamillopora cupula Smitt, p. 33, pi. 7, figs 146, 220m(Shier, 1964; Cook, 1964). 147a-c.
1919 tuberosa Canu & Stichoporina Bassler, p. 98, pi. 1,
figs 20-23; pi. 6, figs 16-19; pi. 7, figs 1-8.
1923 Mamillopora tuberosa (Canu & Bassler) — Canu & DISCUSSION
Bassler, p. 192, pi. 6, figs 16-19; pi. 7, figs 1-8.
1928 Mamillopora tuberosa & — Canu & The (Canu Bassler) Bowden bryozoan fauna now stands at 43 species
Bassler, p. 152. (Table 1), of which 19 are considered herein and 17 are
figured. In the UF samples, the following species were
Material — UF BMNH D41122 found 75971-2, D34306-7, to be particularly common: Biflustra monilifera,
(sample). Cupuladria biporosa, Thalamoporella biperforata, T.
Remarks — Among the specimens used by Canu & chubbi, Adeonellopsis deformis and Metrarabdotos lacry-
Bassler in their of this (1919) original description species mosum. Three of these have erect bifoliate colonies (T.
from the Bowden shell bed of Jamaica were examples (pi. biperforata, A. deformis, M. lacrymosum), two erect tubu- 7, It is notable that these Bowden figs 1-8). specimens are lar colonies (B. monilifera, T. chubbi), and one a free- the retained in M. tuberosa only figured specimens by living lunulitiform colony (C. biporosa). Encrusting colo- & Bassler the Costa Rican Canu (1928); material they nies, which often reach high diversities, may be under-
transferred to M. cavernulosa Canu & Bassler, and in the 1928, represented samples available to us because they
the Dominican material to M. cupula. Cook & Chimonides included few shell substrates. Cheilostomes very are
found it decide whether M. tuberosa (1994) impossible to strongly dominant, comprising 37 (86%) of the species; and M. cavernulosa of M. and are junior synonyms cupula, only four cyclostome species have been recognised. Some
we have retainedM. tuberosa here. provisionally species (e.g., Biflustra monilifera, Thalamoporella chubbi,
Ecology — The of M. cupula was Petraliella cf. depth range given by bisinnata) probably grew loosely around
Canu & Bassler 48-110 and Rucker (1928) as m (1967) soft-bodied animals or plants.
noted that the is abundant and distributed in The species widely Bowden bryozoans clearly represent a shelf as- 71
the 1873-76. Part 1. The Cheilosto- semblage. Indeed, Lagaaij (in Racz, 1971) noted that living Challenger during years the Scientific Results of the of mata. — on Voyage representatives of the Bowden bryozoan species ranged in Report H.M.S. 'Challenger', Zoology, 10(30): 1-216. and inferred that of depth from 0 to 130 m, a depth deposi- I. canariensis Cadee, G.C., 1981. Notes on Bryozoa, Cupuladria tion of between 44 and 64 metres was most probable for
from the British Pliocene Coralline Crag. — Mededelingen the Bowden shell bed. The surface preservation of the Tertiaire en Kwartaire 18: van de Werkgroep voor Geologie, Bowden bryozoans is generally excellent, with little sign of 3-9.
the erect and of the colo- wear, although many free-living Canu, F., 1914. Les Bryozoaires fossiles des terrains du Sud- of fresh, nies are fragmented. It is evident that the bodies Ouest de la France. VIII. Rupelien de Gaas. — Bulletin de la
animals can be for dis- undecayed transported appreciable Societe geologiquede France, (3)14: 465-474. without disarticulation in tances appreciable abrasion, or Canu, F., 1918. Les ovicelles des bryozoaires cylostomes. Etudes
of animals with multielement skeletons anciennes. — Bulletin de la the case (Allison, sur quelquesfamilies nouvelles et
fossils is de 324-335. 1986). Therefore, lack of surface wear in not a Societe geologique France, (4)16: Bassler, 1919. Fossil from the West good indicator of in situ deposition. Paradoxically, the un- Canu, F. & R.S. Bryozoa
Indies. — Publication of the Carnegie Institute of Washing- abraded condition of the Bowden bryozoans may imply
ton, 29: 73-102. quite the opposite because autochthonous bryozoan depos- Canu, F. & R.S. Bassler, 1920. North American early Tertiary its tend to contain an admixture of colonies in various
Bryozoa. — Bulletin of the United States National Museum, how much time states of abrasion depending on they spent 106: 1-879. on the sea-bed before becoming finally buried. A suitable Canu, F. & R.S. Bassler, 1923. North American later Tertiary and model to explain the origin of the Bowden bryozoan fauna Quaternary Bryozoa. — Bulletin of the United States Na-
be one of turbidity currents sweeping the shelf, pick- may tional Museum, 125: 1-244.
colonies and the distal branches of erect ing up free-living Canu, F. & R.S. Bassler, 1927. Classification of the cheilosto-
of the United States Na- colonies, and transporting and burying them in a deeper matous Bryozoa. — Proceedings water environment. tional Museum, 69(14): 1-42.
Canu, F. & R.S. Bassler, 1928. Fossil and Recent Bryozoa of the
Gulf of Mexico Region. — Proceedings of the United States
National Museum, 72: 1-199. ACKNOWLEDGEMENTS Cheetham,A.H., 1966. Cheilostomatous Polyzoa from the Upper
Bracklesham Beds (Eocene) of Sussex. — Bulletin of the The authors are greatly indebted to Abigail Marrero who British Museum (Natural History), Geology, 13: 1-115. assisted with this project as part of the Nuffield Science Chectham, A.H., 1968. Morphology and systematics of the bryo- Scheme, to Doug Jones and Roger Portell (Florida Bursary zoan genus Metrarabtodos. — Smithsonian Miscellaneous Museum of Natural for material, to History) providing Collections, 153(1): 1-121.
Kevin Tilbrook for and Alan Cheetham for discussion, to Cheetham, A.H., 1986. Tempo of evolution in a Neogene bryo-
rates within and across species helpful reviewer's comments. zoan: of morphologic change
boundaries. — Paleobiology, 12:190-202.
Cheetham, A.H., J.B.C. Jackson, J. Sanner & Y. Ventocilla, in
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Bowden reference This First Originalname This paper
& 1919 Canu & Bassler, 1919 AcanthodesiaAcanthodesia savartii forma texturata Reuss,Reuss, 1848 Biflustra monilifera (Canu & Bassler, 1919)
Canu & Bassler, 1919 Adeona heckeli Reuss, 1848 I
Canu & Bassler,Bassler, 1923 Aimulosia brevis Canu & Bassler, 1923 |
Canu & Bassler,Bassler, 1919 Bracebridgia deformis Canu & Bassler, 1919 Adeonellopsis deformis (Canu & Bassler, 1919)
Canu & Bassler, 1923 Callopora dumerilii Audouin, 1826 I
Canu & Bassler,Bassler, 1919 Callopora tenella Hincks, 1880 I
Lagaaij, 1959 Canda caraibica Levinsen, 1909 Canda caraibica Levinsen, 1909
Canu & Bassler, 1923 Conopeum lacroixii Busk, 1872 I
Canu & Bassler, 1919 Conopeum ovale Canu & Bassler, 1919 I
1959 Crisia 18381838 Crisia Lagaaij, Crista elongalaelongata Milne-Edwards, sp.
1959 occidentalis 1910 Lagaaij, 1959 Crisulipora Robertson, Crisulipora sp.
Canu & Bassler, 1919 Cupuladria canariensis Busk, 18591859 Cupuladria biporosabiporosa Canu & Bassler, 1923
Canu & Bassler, 1919 Cupularia umbellata (Defrance, 1823) DiscoporellaDiscoporella umbellata (Defrance, 1823)1823)
Canu & Bassler, 1923 CycloperiellaCycloperiella rubrarubra Canu & Bassler, 1923 I
Canu & Bassler, 1919 Gemellipora punctata Canu & Bassler, 1919 I
CanuCanu&& Bassler, 1919 HemiseptellaHemiseplella grandicella Canu & Bassler, 1919 |
Canu & Bassler, 1923 HippodiplosiaHippodiplosia baccata Canu & Bassler, 1920 ?HippaliosinaIHippaliosina baccata (Canu & Bassler, 1920)
Canu & Bassler, 1919 Holoporella albirostris Smitt, 1873 I
Canu & Bassler, 1923 Holoporella hemisphericahemispherica Canu & Bassler,Bassler, 1923 Celleporaria?Celleporaria ? hemispherica (Canu & Bassler,
1923)
Canu & Bassler, 1923 Mastigophoragranulosa Canu & Bassler, 1923
This paper Mecynoecia proboscideoides (Smitt, 1872)
Canu & Bassler, 1923 Membranipora osburni Canu & Bassler,Bassler, 1923 | -
Canu & Bassler, 1923 MembrendoeciumMembrendoecium parvicapitatumparvicapilatum Canu & Bassler, Antropora parvicapitatumparvicapilatum (Canu & Bassler, 1923)
1923
Canu & Bassler, 1919 MetrarabdotosMetrarabdolos lacrymosum Canu & Bassler, 1919 MetrarabdolosMetrarabdotos lacrymosum Canu & Bassler, 1919
Lagaaij, 1959 Nellia oculata Busk, 1852 Nellia tenella (Lamarck, 1816) 75
This Canu & 1928 paper ?PlagioeciaIPlagioecia dispar Bassler,
This Petraliella cf. bisinuata 1873) paper (Smitt, 1873)
1 1
Canu & Bassler,Bassler, 1923 Rhamphostomella granulosagranulosa Canu & Bassler, 1923 I
Canu & Bassler, 19191919 RhamphostomellaRhamphoslomella laticella Canu & Bassler, 1919 |
Canu & Bassler, 19231923 RhyncozoonRhyncozoon verruculatum (Smitt, 1873)
This Schedocleidochasma paper porcellanumporcellanum (Busk, 1860)
Canu & Bassler, 1919 Schizopodrella unicornis Johnston, 1847 possibly Schizoporella errata (Waters,(Waters, 1879)
This Schizoporella errata (Waters, 1879) paper errata (Waters,
Canu & Bassler,Bassler, 1923 Smittina ophidiana Waters, 1878 I
Canu & Bassler, 1919 Steganoporella [sic] parvicellaparvicella Canu & Bassler, 1919 Steginoporella parvicella Canu & Bassler, 1919
CanuCanu & Bassler,Bassler, 1923 Stephanosella biaperta Michelin, 1841 I
Canu & Bassler, 1919 Stichoporina luberosatuberosa CanuCanu & Bassler, 1919 Mamillopora luberosatuberosa (Canu & Bassler, 1919)
Canu & Bassler, 1923 Stylopoma minuta Canu & Bassler, 1923 I
Canu & Bassler, 1919 Stylopoma spongites Pallas, 1766 I
Canu & Bassler, 1923 TerebriporaTerebripora elongata Canu & Bassler,Bassler, 1923 |
Canu & Bassler, 1923 Terebripora sinefilum Canu & Bassler, 1923 |
Canu & Bassler, 1919 Thalamoporellabiperforatabiperforata CanuCanu & Bassler, 1919 Thalamoporella biperforata Canu & Bassler, 1919
Lagaaij, 1959 Thalamoporella chubbi Lagaaij, 1959 Thalamoporella chubbi Lagaaij, 1959
Table 1. Bryozoan species recorded from the Bowden shell bed. 76 77
Figs 1-6. Bryozoans from the Pliocene Bowden shell bed, southeast Jamaica; 1 - ?Plagioecia dispar Canu & Bassler, BMNH D41289,
infertile 53; 2-4 - of UF 75954, colony, x Biflustra monilifera (Canu & Bassler), UF 75953, part a tubular branch, x 45 (2),
of abraded 80 UF zooid with 160 6 - Antro- group slightly zooids, x (3), 75953, well-preserved cryptocystal denticles, x (4); 5,
& UF with closure pora parvicapitatum (Canu Bassler), 75955, variably-shaped infertile autozooids, some partial plates incor-
porating the operculum, and small kenozooids, x 90 (5), BMNH D34282, autozooid with small ovicell (top left) and infertile
autozooid, x 150 (6).
7-9. from the Bowden shell southeast 7 - BMNH drawn Figs Bryozoans Pliocene bed, Jamaica; Crisulipora sp., D41290, internode by
9 - UF 80 UF Lagaaij (1959, text-fig. 5), x 38; 8, Mecynoecia proboscideoides (Smitt), 75951, autozooids, x (8), 75952, gono-
zooid, x 75 (9). 78 79
Figs 10-15. Bryozoans from the Pliocene Bowden shell bed, southeast Jamaica; 10-13 - Cupuladria biporosa Canu & Bassler, UF 75958,
27 of autozooids with distal 70 UF upper surface of lunulitiformcolony, x (10), UF 75956, group interzooidalavicularia, x (11),
75958, large vicarious avicularium with distal interzooidal avicularium, x 170 (12), UF 75957, underside of colony showing
sectors with 30 14 - umbellata BMNH BZ 3504, of autozooids with cribrate pores, x (13); Discoporella (Defrance), group
- A- frontal shields and distal avicularia, x 75; 15 Steginoporella parvicella Canu & Bassler, UF 75959, group of and B-zooids,
the latterwith longer opesia, x 42.
Figs 16, 17. Bryozoans from the Pliocene Bowden shell bed, southeast Jamaica; 16 - Nellia tenella (Lamarck), BMNH D41133, internode, x
40; 17 - Thalamoporella chubbi Lagaaij, BMNH D41208, holotype specimen drawn by Lagaaij (1959, text-fig. 2A) showing
autozooids and two spatulate avicularia, x 38. 80
Figs 18-23. Bryozoans from the Pliocene Bowden shell bed, southeast Jamaica; 18-20 - Thalamoporella biperforata Canu & Bassler, UF
35 UF 95 UF 85 75960, group of autozooids and two avicularia, x (18), 75961,autozooids, x (19), 75961, avicularium, x (20);
21 - Thalamoporella chubbi Lagaaij, UF 75962, autozooids, x 58; 22, 23 - Adeonellopsis deformis (Canu & Bassler), UF
75963, surface ofbifoliate branch, 75964, autozooids with x 50 (22), UF depressed suboral avicularia, x 140 (23). 81
Figs 24-29. Bryozoans from the Pliocene Bowden shell bed, southeast Jamaica; 24-26 - Celleporaria? hemispherica (Canu & Bassler), UF avicularia and 75965, top of a hemispherical colony, x 5 (24), UF 75965, autozooids, x 75 (25), UF 75966, suboral primary ori-
fice with 27 - ? baccata & BMNH D sinus (lower right), x 160 (26); Hippaliosina (Canu Bassler), 34298, group of autozooids
- with small avicularia, x 80; 28, 29 Metrarabdotos lacrymosum Canu & Bassler, UF 75968, autozooids, x 81 (28), UF 75967,
gonozooids with abraded ovicells leaving large holes, x 35 (29). 82
31. the tuberosa & Figs 30, Bryozoans from Pliocene Bowden shell bed, southeast Jamaica; Mamillopora (Canu Bassler), UF 75972, upper
surface of a colony, x 19 (30), UF 75971, colony underside with four prominent rootlet chambers, x 12 (31).
Figs 32-37. Bryozoans from the Pliocene Bowden shell bed, southeast Jamaica; 32 - Schizoporella errata (Waters), UF 75969, autozooids
avicularia lateral the - cf. and with adventitious to orifice, x 75; 33 Petraliella bisinnata, UF 75970, autozooids sparse adventi-
tious avicularia (top) and ovicell vestiges on the three proximal zooids, x 35; 34, 35, 37 - Mamillopora tuberosa (Canu &
later autozooids with autozooid avicularium, Bassler), UF 75972, early autozooids, x 105 (34), avicularia, x 75 (35), orifice and
x 230 (37); 36 - Schedocleidochasma porcellanum (Busk), UF 75973, autozooids, avicularia and two abraded ovicells, x 115. 83