A Late Cretaceous terebratulid brachiopod from Jamaica, and its significance for Mesozoic brachiopod palaeobiogeography

and evolution

Michael R. Sandy*, David A. T. Harper t, Stephen K. Donovan# and David J. Miller#

SANDY, M. R. , HARPER, D. A. T. , DO OVAN, S. K. & MILLER, D. J. 1997. A Late Cretaceous terebratulid brachiopod from Jamaica and its significance for Mesozoic brachiopod palaeobiogeography and evolution. Proceedings of the Geologists' Association, 108, 201-207. Recent fieldwork has accumulated collections of terebratulid brachiopods from Late Cretaceous strata on Jamaica referrable to the genus Dyscritothyris Cooper. This is confirmed by the preparation of serial sections of the internal structures of one well-preserved specimen. The only other record of th is genus is from the Late Cretaceous rocks of Cuba. The specimens from Jamaica and Cuba are considered to belong to the same species, confirming oceanographic links between these islands during the Late Cretaceous, already apparent from similarities between their rudist bivalve and echinoid faunas . The small-sized Dyscritothyris is considered to be a Late Cretaceous derivative of the Cretaceous terebratulid Capillithyris which has been described primarily from Europe. Such a relationship would support models for the dispersal and evolution of brachiopods during the continued opening of the Central Atlantic Ocean during the Cretaceous.

* Departmelll of Geology, University of Dayton, Dayton, Ohio 45469-2364, USA. t Departme/11 of Geology, University College Galway, Galway, Ireland. # Departmei!T of Geography and Geology, University of the West Indies, Mona, Kingston 7, Jamaica.

1. INTRODUCTION

Fossi l brachiopods are rare in the Mesozoic rocks of the Antillean region. This paper represents the first detailed investigation of Cretaceous terebratulid brachiopods from Jamaica. Records of Cretaceous brachiopods from Jamaica are uncommon (Trechmann, 1927; Harper & Donovan, 1990; Harper, 1993) and they are certainly not abundant, Dyscritothyris being recorded from only one locality to date (Harper & Donovan, 1990) in the Central Inlier (Fig. 1). This , however, represents an important record of a Cretaceous terebratulid from the Caribbean region .

Cooper's original description of Dyscritothyris from Cuba ( 1979) appears to be the only other record of the genus. There are, however, a reasonable number of con­firmed records of other terebratulid brachiopods from the Cretaceous of North America (for example, Roemer, 1852; Stanton, 1895; Warren, 1937; Stephenson, 1941 ; Cooper, 1955; Ager, Underwood & DeFord, 1963; Owen, 1976, 1981 ; and other references in Sandy, Murphy & Rodda, 1995). Records from the South American Cretaceous are, by comparison, rare (e.g. Sandy, 1991 ). Most of these records are Early Cretaceous in age, not Late Cretaceous, making the material from Jamaica all the more noteworthy. Cooper ( 1979), however, recorded a few additional Late Cretaceous terebratulid genera from Cuba; Cruralina, Orthothyris, Rugia and Terebratulina.

In an effort to clarify the affinities of the Jamaican material , tr:J.nsverse serial sections have been prepared of

Proceedings of the Geologists' Association, 108, 20 1-207.

Fig. 1. Outline map of the area around Frank:field, parish of Clarendon, central Jamaica, showing the position of the locality(+) discussed in the tex t (redrawn after Donovan et al., 1995, fig. 14). Key: thick lines = principal roads; thin line = Rio Minho (note that there are numerous streams that feed thi s river, but which are not shown for clarity). The inset map shows the position of the Central Inlier (stippled) and the Frankfield region · (F) with in Jamaica.

0016-7878/97 $10·00 © 1997 Geologists ' Association

202 M . R. SANDY ET AL.

one well -preserved specimen. The techniques and equ ip­ment used are the same as those described by Sandy ( 1989) . The seri al-sectioning programme confi rms ass ignment to Dyscritothyris and moreover, sugges ts the small -s ized genus is a Late Cretaceous derivative of the Cretaceous terebratulid Capillithyris which has been described mainly from Europe. Specimens documented in the present paper are deposited in the James Mitchell Museum , Unversity College Galway (JMM) and the Natural Hi story Museum, London (NHM).

2. GEOLOGICAL SETTING

The Central Inlier, in the centre of the Clarendon Block (Fig. I), has the second larges t outcrop area of Cretaceous rocks in Jamaica, after the Blue Mountain Inlier to the east. The stratigraphic succession of the Central Inlier has most recently been reviewed by Robinson ( 1988, 1994). The Guinea Corn Formation is a sequence of grey, rubbly lime­stones and carbonate mudrocks interbedded with reworked volcaniclastics and some thin lignites. The siliciclastics are generally unfossili fero us, but the limestones have yielded a diverse marine fauna. This fauna is best known for its di verse assemblage of rudi st bivalves (Chubb, 197 1), but it also inc ludes other benthic molluscs, corals, echinoids, foraminiferans and ostracodes, with rare brachiopods, bar­nac les and brachyuran crabs. Ca lcareous nannofossil s indicate the age of this formation to be late Campanian to earl y Maastrichtian (Jiang & Robinson 1987a, b).

The D. cube1uis hori zon is in the bank of the Rio Minho, adjacent to the B3 road west of Frankfield, parish of Clarendon. Thi s is Stop 3 of Donovan, Jackson, Dixon & Doyle ( 1995, p. 38-39), about I km west of Frank field ( I :50 000 topographic new series sheet II ' Mandeville ' , NGR 099 670; Fig. I). The Guinea Corn Formation is exposed as steeply-dipping beds on the opposite side of the river to the road. Monospecific, autochthonous horizons of the small , gregarious rudist Biradiolites are a di stinctive feature of this sequence and one has been taken as the top of the measured section ; the base is a fau lt (Fig. 2) . The secti on is essentiall y a series of alternating nodu lar to mass ive limestone beds interbedded with mudstones . The limestones may be at least part ly secondary in origin, as indicated by lateral gradations from nodul ar to massive limestones and the interstitial sed iment between nodules, which may be similar to that of the mudrocks. However, unlike the mud­rocks, the limestones are rich in fossi ls, particu larly benthic molluscs (mainly rudists and mouldic gastropods) and echinoid spines, and are commonly microsparry to sparry with bands of bioclastic debris. Mudrocks are either grey, blocky, sticky muds or orange-brown, thinly (20-50 mm) bedded, mudrocks that are generally poorly fossi li ferou s, but which include some thin bioclas tic horizons. This succession agrees well with the migrating carbonate bank sequences (Skelton , Donovan & Dixon, 1994: 27) noted from locali ties 4 and 5 of Donovan et al. ( 1995), although Titanosarcolites giganteus (Whitfield) is not commonly seen in the admittedly more limi ted exposure at the D.

KEY ~ Nodular limestone

~ Massive limestone

~ Radiolitid limestone

[.-:<:_-~;·: I Mudstone

- Muddy sandstone

·.· .... ... :. ·.: . . :··:":.·.-:-.=: -FAULT

1111111 MSfmcPC ~

SST

Fig. 2. Strip log of measured secti on through pan of the Upper Cretaceous Guinea Corn Formation at GR 099 670, I krn west of Frankfield, parish of Clarendon, Jamaica. The Dyscri10rhyris cubensis horizon is the base of the muddy lens at the top of unit 8. The section is measured from a prominent, but thin , unit with in situ Biradiolires (horizon I), while the base is fau lted (with a prominent shell of Tiranosarco!ires giganreus (Whitfie ld) in the overhang above the fault). Key: M=mudstone; S=siltstone; f, m, c SST=fine-, medium- , coarse-g rained sandstone; P=pebble conglomerate; C=cobble conglomerate.

A TEREBRATULID BRA HIOPOD F R OM J A M A l A 203

cubensis locality. The presence of a 2 em thick li gnite bed in horizon 5 is indicative of ' .. . occasional hyposalinity and nutrient flux ' (Skelton et al., 1994). D. cubensis has onl y been found at the top of the muddy lens towards the ba e of horizon 2. Specimens are very rare and those documented herein have been collected since 1986, when the fi rst specimen was found by C. R. C. Paul.

3. BRACHIOPOD PALAEOECOLOGY

Dyscritothyris is a small , biconvex peduncul ate species suggested by the presence of an open pedicle foramen. To date no other brachiopods have been reported from the Rio Minho locality despite intensive searches over a number of years. The brachiopods occur near the base of horizon 2, within a muddy lens, associated with nodul ar limestones. The majority of brachi opod pedicles found in extant brachiopods are either short stout plenipeduncul ate stalks or longer thread-like rhizopeduncul ate structures. Although there is no direct ev idence, Dyscritothyris may have been attached by a short stout pl enipeduncul ate structure, providing a strong bond with the substrate in the re lati vely hi gh energy conditions associated with migrating she ll banks of rudi sts and other molluscs. Dyscrirorhyris is less than I em in length, comparable to the taxa Surlyk ( 1972) associated with small substrates such as shell fragments in carbonate banks. Habitats within the interstices between li ving and skeletal molluscan material may have supported a cryptic li fe mode for Dyscritorhyris. Elsewhere, small shells of the brachiopod Pseudorhaetina have been reported from interstices between the individual coral lites of coloni al corals from Triass ic strata in Sonora, Mex ico (S tanley, Gonzalez- Leon, Sandy, Senowbari-Daryan, Doy le, Tamura & Erwin , 1994). To date, however, Podichnus, the trace fossil associated with pedicle etchings on she ll substrates (Bromley & Surlyk, 1973) has not been reported from the rudi sts at Rio Minho.

4. PALAEOBIOGEOGRAPHIC AND EVOLUTIONARY SIGNIFICANCE

Dyscritorhyris was orig inally recorded from two Late Cretaceous localities in Cuba by Cooper ( 1979). The material described here in from Jamaica i considered to be referrable to the Cuban spec ies, namely Dyscritorhyris cubensis. This certainl y agrees with the close proximity of these islands during the Late Cretaceous (Draper, Jackson & Donovan, 1994). It is also possible, however, for terebra­tulid brachiopods to di sperse over considerable di stances; therefore, they should not be taken as reliable indicators of geographic proxi mity on the ir own. The sum of geological ev idence, such as strat igraphy and sedimentology, as well as incorporation of palaeomagnetic data , a ll he lp in palaeo­geographic reconstructions in which palaeontology may play an important ro le. Other palaeonto logie ties between the Late Cretaceous of Jamaica and Cuba inc lude the sim­ilarities between the spectacular rudist bivalve faunas of the two islands (Chubb, 197 1) and certain of the

Campanian- Maastrichtian echinoids (Donovan , 1993), amongst others.

The small-sized Dyscritothyris is considered to be a Late Cretaceous derivative of the terebratulid genus Capillithyris Katz, 1974 which ranges from the Early to Late Cretaceous. Capillithyris was originally described from the Cretaceous of England by Middlemiss ( 1959), under the generic name Platythyris (see di scuss ion by Middlemi ss, 199 1: 232-234). The similarity in the form of the brachidium (lophophore support) in both Dyscritothyris and Capillithyris suggests that they may be closely re lated. In fact , Dyscritothyris may be derived from Capillithyris; possibly by paedomorphosis (suggested by the small size of Dyscritothyris). Alter­natively, but less likely, Dyscrirothyris, may simply be a small -s ized species with an origin elsewhere.

Based on current distributional information, it would appear likely that the west Atlantic species, Dyscritorhyris cubensis, was derived from European form that migrated across the opening Atlantic Ocean. Thi s dispersal route has been suggested for other brachiopods during the Cretaceous (Sandy, 199 1; Sandy, Murphy & Rodda, 1995). The dispersal and diversification of terebratulid brachiopods wou ld therefore appear to be linked to a major tectonic event, that is, the continued opening of the central Atlantic Ocean during the Cretaceous.

5. SYSTEMATIC DESCRIPTION

Order: TEREBRATULIDA Waagen, 1883 Suborder: TEREBRAT LIDINA Waagen, 1883

Famil y: Uncertain Subfamily: CAPILLJTHYRIDI AE? Cooper, 1983

Genus: DYSCRITOTHYRIS Cooper, 1979

Type species

Dyscritothyris cubensis Cooper, 1979, p. 16, fig. I , from the Upper Cretaceous of Cuba. The only species known.

Dyscritorhyris cubensis Cooper, 1979 Figs 3-4

1979 Dyscritothyris cubensis Cooper: 16, fig. I ; Discritothyris cubensis Cooper (sic), Pl. 7, figs 3-8.

1983 Dyscritothyris cubensis Cooper; Cooper: 188, Pl. 61 , figs I 0-13.

1990 Dyscritorhyris sp. cf. D. cubensis Cooper; Harper & Donovan: 27.

1993 Dyscritothyris sp. cf. D. cubensis Cooper; Harper, p. II 0, figs 3.20-23.

Revised diagnosis

Small , ubcircular to rounded Dyscritothyris species of subpentagonal outline; sides well rounded. Val ves nearly equal in convex ity. Anterior commissure uniplicate in adult specimens. She ll surface generally smooth, may bear fine

204 M. R. SANDY ETAL.

Fig. 3. Dyscritothyris cubensis Cooper, 1979, from the Central lnlier, Jamaica, Guinea Corn Formation, Upper Cretaceous: (a) ventral (ped icle); (b) dorsal (brachial); (d) posterior and (e) anterior views of conjoined pair, NHM BC I 000, all x 8 (also figured Harper, 1993, fig . 3, 20-23); (c) ventral view of conjoined pair, JMM Br2042, x 18; (f) dor al view of conjoined pair, JMM Br2043, x 10; (g-j ) various views of conjoined sectioned specimen (see also Fig. 4); (g) ventral (x 3.25); (h) dorsal (x 3.25); (i) lateral view (x 3.25); U) anterior (x 2.75) views; (k) ventral view of conjo ined pair, JMM Br2044, x 12; (I) ventral view of conjoined pair, JMM Br2045, x II ; figs c, f, k and I are scanning electron micrographs.

capillation and fine concentric growth lines. Ventral valve interior lacks dental plates.

Material and occurrence

Cooper ( 1979) recorded the species from the Upper Cretaceous, Habana Province, Cuba, from conglomeratic

sand beds under Eocene-Oligocene chalk, 7.3 (?) km E-SE of Madruga, 2.4 km S of Grua Esperanza and at a second locality 150m N of Grua Esperanza, 6 km E of Madruga.

The Jamaican material , which is considered conspecific, is from the Central Inlier, Guinea Corn Formation, Upper Campanian-Lower Maas trichtian , Upper Cretaceous:

A TEREBRATULI D B R AC HJOPOD FROM JAMA ICA 205

o~~~ ~ao o~ ~ q

~ 0.7 - 08 • { ) . 1.0

~oo 00 ''

c ,,

Fig. 4. Transverse seri a l sections through a spec imen of Dyscritothyris cubensis Cooper, 1979. Ini tial section at 0 .0 mm represents ti p of the pedicle valve umbo. Cumulati ve distance from initial section given in mm. Features observed include: initial section through brachial va lve umbo (0.7 mm); broad concave cardinal process (0.8); incipient hinge plates ( 1.4); brachidium traced to 2.5 mm . Magni fication of sections: 0.0-0.5, x 3.3; 0.7-2.3, x I 0; 2.3, 2.5, x 20. Spec imen sectioned perpendicular to max imum length . JMM Br2044, Central Inlier, Jamaica, Guinea Corn Formation, Upper Cretaceous. Dimensions of specimen: length 8.0; width 7.1; thickness 3.9 mm.

rubbly limestone horizon at GR 4 19 455 , in the Rio Minho west of Guinea Corn, on the B4 Frankfie ld to Spaldings Road (the ex posure is on the opposite side of the ri ver from the road), pari sh of Clarendon, Jamaica. The brachiopods are rare. The fauna is dominated by rudi sts (Chubb, 1971 ; Kauffman & Sohl , 1974) and other molluscs, corals, echinoids, foraminifers and ostracods. About 30 conjoined pairs of valves are known from the Rio Minho locality.

Description

(Adapted from Cooper, 1979, p. 16 and Harper, 1993, p. II 0 .) Small , subcircul ar to rounded subpentagonal in outline with maximum width at about midvalve; hinge width about three-fifths maximum width ; s ides we ll rounded; anterior margin broadly to somewhat narrowly rounded depending on age, subnasute in old age. Valves nearly equal in depth. Anterior commissure rectimarginate in young growth stages, uniplicate in adult specimens; beak small and low, pedicle fo ramen moderate ly large, submesothyrid. Deltidi al pl ates disjunct (Cooper recorded as conj unct). Shell surface generall y smooth but fine capillation (Figs 3a,e) and fine concentric growth lines may be present.

Ventral (pedicle) valve about nine- tenths as wide as long, gently and evenly convex in lateral profile with max imum convex ity near rnidvalve; anterior profile broadly domed;

median region somewhat narrowed, sides slopi ng moder­ate ly. Umbonal region broadl y convex , ri sing to midvalve that is swollen; anterior somewhat fl attened to fo rm shallow sulcus and short rounded tongue. Valve interior lacks dental plates, but has moderately strong teeth .

Dorsal (brachial) valve about nine-tenths as wide as long, moderate ly convex in lateral profile, convex ity slightly less than that of opposite valve; anteri or profi le broadly and moderate ly rounded, not medially narrowed. Umbonal and posteromedian regions fairly strongly swollen; anterior half somewhat fl attened and narrowed anteriorly to form low, poorly defined fo ld. Valve interior with strong socl<et ridges, flattened outer hinge pl ates and loop about one-third the length of the dorsal valve, median ridge low and obscure, transverse band of loop not seen.

Discussion

The first detailed description of the species by G. A. Cooper ( 1979) was based on material collected by R. H. Palmer from the late Cretaceous rocks of Cuba. Harper 's ( 1993) description of the Jamaican materi al was based on a few conjoined valves and lacked detailed data on the valve interiors. On the bas is of the available in fo rmation the Jamaican material was compared with Cooper 's species although a limited Principal Component Analys is revealed no external shape differences between the two samples; the Cuban fo rm, however, was significantly larger (Harper, 1993).

Cooper ( 1979) attempted to excavate infi lling matrix from the Cuban specimens of this species but was unsuc­cessful. He then seri ally sectioned two specimens (Cooper, 1979: 15, fi g. 1). Like the specimen sectioned herein , he did

Table 1. Measurements and stati stics for a sample of Dyscritothyris cubensis Cooper, 1979 from the Guinea Corn Formation, Centra l Ln I ier, Jamaica

slpv slbv mwi hwi pmwi dpv dbv

Measurements NHM BC IOOO 5.2 4.5 4.8 3.5 3.0 1.4 1.0 NHM BC IOI9 6.7 5.9 5.8 3.0 4.0 1.8 1.5

Statistics (n= I 0) Vector means 6.6 4.9 5. 1 3.3 3.2

Variance 2.8 2.5 2.6 1.3 1.6 Covariance 2.3 2.4 1.3 1.5 Matri x 2.7 1.4 1.6

0.9 0.7 1.1

Abbreviations: slpv : sagittal length of pedicle valve; slbv : sagittal length of

brachial valve; mwi :maximum width; hwi :hinge width; pmwi : position of maximum width measured anteriorly from posterior margins; dpv : depth of

pedic le valve; dbv : depth of brachial valve; N : sample size.

206 M . R. SANDY ET AL.

not encounter the anterior transverse band of the brachi­dium. However, he indicated that the structure of the socket ridges and the hori zontal tapering style of the hinge pl ates defined a unique brachiopod. Another terebratulid with horizonta l tapering hinge pl ates, Platythyris Middlemi ss, 1959 (now referred to Capillithyris), was considered by Cooper to differ markedly in both external and internal characters. As Cooper pointed out ( 1983 : 188), Dyscritothyris lacks the capillate external ornament of Capil/ithyris. Despite Cooper 's comments, the re lationship between Capillithyris, which spans much of the Cretaceous (Hauterivian to Campanian, see Midd lemiss, 1984), and Dyscritothyris, requi res closer examination. internally both genera possess a fl at cardinal process with a di stinct 'U'­shaped dorsa l valve fl oor/socket ridge outline j ust anterior of thi s (compare Fi g. 4 with Cooper, 1979, fig. I, and the sections of Capillithyris cristobali in Middlemi ss, 1978, fi g. 7) . Attached crural bases are not obvious features in Dyscritothyris, but are present in some spec ies of Capillithyris (see Midd lemiss, 1978). In fac t, the develop­ment of cuneate hinge pl ates with attached crural bases was interpreted as a gerontic feature in Capillithyris diversa (Middlemiss, 1978, fi g. 4 ). The general form of the development of the crural processes, hinge pl ates and free crura are not markedly different between Capillithyris and Dyscritothyris. However, such simjlarity might be readily evident between forms that do not develop hinge plates immediately and have reduced crural bases. As stated previously, the transverse band has not been observed in Dyscritothyris. From both internal and external morpho­logica l considerations, it is possible that Dyscritothyris could be a micromorphic terebratu lid deri ved, paedo­morphically, from Capillithyris.

Capill ate o rnament is not an obvious fea ture of Dyscritothyris, but fine capillation can be di scerned in some specimens (Fig. 3a,e). It is a vari able feature among species of Capillithyris in which the shell may be 'capillate, or smooth with more or less obvious longitudinal stri ae' (Oieni & Middlemiss, in Dieni , Middlemi ss & Owen, 1975: 195).

The other terebratulid that Cooper di scussed in relation to Dyscritothyris was Gryphus. ' In spite of the incompleteness of the spec imens of Dyscritothyris the ensemb le of structures suggest those seen in Gryphus' (Cooper, 1979: 15). However, crural processes with attached crural bases have not been observed in the seri a l sections of Dyscritothyris (see Cooper, 1979, Pl. I, fi g. 35, and serial sections of the genus), obviating the poss ibili ty of a close relationship.

Despite its overall larger size the medium-sized tere-

bratulid Pseudogibbithyris arabica Owen, 1995 has much in common with Dyscritothyris. Owen described Pseudo­gibbithyris fro m the Maastrichtian of the United Arab Emirates-Oman border region. The fo ld of the anterior commissure is more pronounced in Pseudogibbithyris than in Dyscritothyris but outline and profile are s imil ar, although Pseudogibbithyris is more biconvex in profil e. These differences could be related to the larger size of Pseudogibbithyris. The transverse seri a l secti ons of Pseudogibbithyris (Owen, 1995, fi g. 2) have many close simil arities to those of Dyscritothyris (Cooper, 1979, fig. I; Fig. 4 herein). The hinge plates seen in Owen's sections taper medi ally as they do in Cooper 's specimen B. A cardinal process and complete transverse band are present in Pseudog ibbithyris but are not seen in any of the series of sections taken of Dyscrilothyris. Their absence in sections of Dyscrilothyris does not prove their absence in the genus. The series of secti ons of Pseudogibbithyris and Dyscrito­thyris are suffic iently close to suggest a very close re lati on­ship between the two genera; e ither a c losely related ancestry, or that Pseudogibbithyris is a junior subjecti ve synonym of Dyscritothyris. If the latter is the case, a broadly distr ibuted low-latitude Tethyan distribution is suggested fo r this latest Cretaceous brachiopod.

ACKNOWLEDGEMENTS

We thank the numerous geologists who have helped collect the Jamaica n spec imens of thi s spec ies since 1986, inc luding Chri s Paul , Eamon Doy le, Hal Di xon, Niall Hoey, and students of S K.D 's GL IOO and GLIOB fi e ld c lasses. We thank Cornelis J. Veltkamp (Uni versity of Li verpool) for taking the scanning e lectron micrographs and Frank Middlemiss and Elli s Owen for comments that improved the manuscript. MRS thanks Dr Frank A. Middlemiss for introducing him to Cretaceous terebratulids and gratefully acknowledges the donors of The Petroleum Research Fund, administered by the American Chemical Society, and the Uni versity of Dayton Student Fellow's Program for support. Lindsey Gri ffith made the seri a l secti ons. OATH acknow ledges support from the Curry Fund of the Geologists' Association to visit Jamaica in 1994. Part of this research was undertaken by SK.D when he was a Vi siting Professor at the Uni versity of Portsmouth during March 1996. This study was completed during tenure of Grant 5722-96 from the National Geographic Society. We thank Simon Mitchell (UWI , Mona) fo r allowing us to benefi t from his knowledge of the sedimentology of the Guinea Corn Formation.

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Manuscript received 26 May 1996; revised typescript accepted 7 August 1996