Some Fossil Echinoids (Echinodermata) from the Cenozoic of Jamaica, Cuba and Guadaloupe

S TEPHEN K. DONOVAN

Department of Geology, University of the West Indies, Mona, Kingston 7, ]amaica

ABSTRACT. – A ‘lost’ cidaroid test from Jamaica, formerly classified as Cidaris melitensis Wright (= Cidaris [Tretocidaris] anguillensis Cutress of Miocene age), is probably not nonspecific and may be Eocene. Echi- nometra Iucunter (Linne) from the parish of St. Elizabeth is either from a well-indurated reef limestone of Sangamonian age or a Holocene beach rock. Clypeaster duchassaingi Michelin is a rare example of a fossil echinoid from Guadaloupe. A mid-Cenozoic Clypeaster sp. from Cuba shows an unusual growth deformity, in which one plate column of most petals ceased growing prematurely and later re-initiated growth.

I NTRODUCTION Tretocidaris anguillensis Cutress, 1980:61. The specimens documented herein are ??Tretocidaris anguillensis Cutress; Donovan, from the Museum of Comparative Zoolo- 1988: Table 1. gy, Harvard, Massachusetts (MCZ). The Cidaris? (Tretocidaris?) anguillensis? Cutress; MCZ has one of the largest and most im- Donovan, 1993:375. portant collections of fossil echinoids from Material, Locality and Horizon. —A single the Caribbean, including those of Louis and test, MCZ 3380, from Jamaica. Locality and Alexander Agassiz, R. T. Jackson and H. L. horizon unknown. Arnold and Clark (1927: Clark, among others (Winsor, 1991). From 11-12) noted that this specimen “. . was this significant accumulation, certain Ce- picked up by a very small boy who was nozoic specimens are presently redocu- too bewilderd [sic] to remember where he mented because they are either of geo- found it, but its condition indicated soft graphic or stratigraphic significance, or earth, perhaps the remains of disintegrated because they show unusual morphological strata now cultivated for bananas.” features. Description .—Test rounded pentagonal in Descriptive terminology used herein outline, but slightly crushed. Moderately follows Melville and Durham (1966). The high, with a rounded ambitus at about 50% classification of echinoids is essentially that of test height. Flattened adapically and of Smith (1981, 1984) and Smith and Wright adorally. (1989). The use of open nomenclature fol- Apical system unknown, but large (about lows the protocol suggested by Bengtson size of peristome) and rounded pentagonal (1988). Where given, synonymy lists refer in outline. to the specimen being discussed. Ambulacra moderately broad (widest ambitally) and moderately sinuous, with S YSTEMATIC P ALEONTOLOGY about 33–35 primary tubercles (=plates) per Class ECHINOIDEA Leske plate column. Poriferous zones in shallow, Subclass CIDAROIDEA Claus depressed grooves. Pore pairs sunken, sub- Order CIDAROIDA Claus conjugate (sensu Cutress, 1980), with in- Family CIDARIDAE Gray terporal partition high adapically, but with Genus Cidaris Leske a shallow groove adorally. Interporal par- Subgenus Cidaris (Tretocidaris) Mortensen tition separates pore pairs by one pore di- Cidaris (Tretocidaris) sp. ameter or slightly less. Both adradial and (Fig. 1A, B) perradial pores circular in outline. Adja- Cidaris melitensis Wright; Arnold and Clark, cent pore pairs within columns separated 1927:11, 12; Cutress, 1980:61. by a low ridge, sloping perradially and

164 CENOZOIC FOSSIL ECHINOIDS 165 orally, and connected to the interporal par- 1980:61 ). This specimen was rediscovered tition adorally. Interporiferous zones about by the present author in March 1993. It is the width of one pore column. Primary important to illustrate (Fig. 1A, B) and de- (=marginal of Fell, 1966:Fig. 235.4) tuber- scribe this test for the first time, as it has cles adjacent to sloping ridges, with one been referred to a species that has not oth- secondary (=internal of Fell, 1966) tuber- erwise been reported from Jamaica. In- cle per ambulacral plate, offset to second- deed, C. (T. ) anguillensis is unknown from ary of adjacent plate, producing a zigzag outside the type area of Anguilla (Cutress, pattern. Ambulacral primary tubercles 1980:63), where it occurs only in the upper smaller than interambulacral scrobicular Lower Miocene. tubercles. MCZ 3380 differs in some important de- Interambulacra about three times wider tails from tests of C. (T. ) anguillensis sensu than ambulacra ambitally and composed of stricto. The ambulacra are gently sinuous, 5-6 plates per plate column. Interambu- possibly close to what Jackson (1922:21) lacral plates higher than wide adapically, called “subundulate,” whereas Cutress high and moderately broad ambitally, rel- (1980:59) considered them to be “nearly atively lower and wider adorally. Primary straight.” Interambulacral plates figured by tubercles conical, noncrenulate and per- both Cutress (1980:P1. 2, Fig. 11) and Cot- forate, with planar platforms and broad, teau (1875:P1. 1, Figs. 4,8) have only a sparse circular (adapically) to elliptical (adorally) covering of granular tubercles outside the areolae. Scrobicular tubercles circular in scrobicular circlet, unlike MCZ 3380. These outline, surrounding primary tubercles as same illustrations also show that the in- a continuous ring, and reaching to or close terporiferous zone between the marginal to the plate edges. Granular tertiary tuber- tubercles is moderately broad, whereas cles in angles of plate and, particularly, these tubercles are close together on the along the interradial margin. Jamaican specimen, even ambitally. These Peristome large, more pentagonal in out- points combine to cast some doubt on the line than periproctal opening and about identity of this specimen, although these half the diameter of the (crushed) test. Ra- differences may be due to its small size dioles not preserved. (specimens examined by Cutress, 1980, had Dimensions. —(Variations in measure- ambital test diameters between 17.5 and ments due to the test having been crushed.) 26.5 mm). Ambital test diameter= 12.4 -14.0 mm; test The Jamaican Miocene is dominated by height = 7.2 mm; peristome diameter = shallow-water limestones (Newport For- 5.0-6.1 mm; periproct diameter = 4.7-6.0 mation), and deeper-water chalks with mm. cherts, bentonites and bioclastic lime- Remarks. —Arnold and Clark (1927) doc- stones presumably derived from a shallow- umented this specimen, the only one from er-water environment by mass flow pro- Jamaica referred to Cidaris melitensis Wright, cesses. Only clypeasteroids have been as being a “typical example” of the species, identified in the Newport Formation (Don- but without formal description or illustra- ovan, 199 1), which is invariably well-in- tion. Although Cutress (1980:59) referred durated. The bioclastic limestones of the to Arnold and Clark (1927:P1. 1, Figs. 12- Montpelier Formation include a more di- 17) as illustrating the specimen, this was a verse, albeit fragmentary, fauna (Donovan, misreference to the Anguillan specimens research in progress), including cidaroids, figured by Jackson (1922). Cutress (1980) but these appear to be closer to Eucidaris differentiated between the Maltese C. mel- Doderlein. Furthermore, the matrix infill- itensis and the Caribbean Tretocidaris an- ing MCZ 3380 is quite unlike either of these guillensis Cutress, which had hitherto been two formations. Rather, its colour is much identified as the same species. However, more reminiscent of the Eocene Yellow Arnold and Clark’s “. . specimen could Limestone Group. This would agree with not be located at the Museum of Compar- Arnold and Clark’s (1927) deduction that ative Zoology or elsewhere . .“ (Cutress, this specimen came from “. . . soft earth, perhaps the remains of disintegrated strata Family ECHINOMETRIDAE Gray “ Of the two cidaroids described from Genus Eckinometra Gray the Yellow Limestone Group (Donovan, Echinometra lucunter (Linne, 1758) 1993), MCZ 3380 is closer to Prionocidaris (Fig. lC) loveni (Cotteau) than FeIlius foveatus (Jack- son), without being nonspecific. Echinometra Iucunter (Linne): Arnold and In conclusion, MCZ 3380 is not suffi- Clark, 1934140; Gordon, 1991:38, Table ciently similar to C, (T, ) anguillensis to be 1, Fig. 2; Donovan, 1993:382, considered nonspecific and it may be Eo- cene, rather than Miocene in age. It is left For a more complete synonymy list of E, in open nomenclature until growth series lucunter, see Weisbord (1969:302-308). of Jamaican Eocene and Miocene cidaroids Material, Locality and Horizon. —A single are available for comparison. test, MCZ 3499, from the parish of St, Eliz- abeth, southwest central Jamaica (precise Subclass EUECHINOIDEA Bronn locality unknown). Probably from the Fal- Order ECHINOIDA Claus mouth Formation sensu Iato, a last inter- CENOZOIC FOSSIL ECHINOIDS 167

FIG. 2. Clypeaster spp. from the Caribbean. A, Clypeaster duckassaingi Michelin, MCZ 1418, apical view, from Guadaloupe. B, C, Clypeaster sp., MCZ 101624, 101625, apical views. B, specimen showing growth deformities in petals (arrowed). C, specimen without growth deformities. Specimens whitened with ammo- nium chloride. Scale bar represents 10 mm.

. 168 S. K. DONOVAN

FIG. 3, Clypeaster sp., MCZ 101624, from Cuba. Growth deformities (arrowed) of ambulacral petals. A, ambulacrum IV, showing an early termination of petal growth in the anterior plate column, with a return to petal growth more adapically. B, ambulacrum V, with a displaced elliptical pore in the posterior plate column (* = fifteenth plate of column, although some small plates may have been lost adjacent to the (missing) apical system). C, ambulacrum I, anterior plate column; compare with A, D and E. D,ambulacrum II, anterior plate column. E,ambulacrum III, petal. On the left (in diagram = right of fossil) one of theinner ambulacral pores is enlarged and obliquely sunken, On the right petal, growth seems to have stopped prematurely and later resumed. In all examples the apical system is towards the top of the page; except for B, the adoral termination of the petal is shown for at least one plate column. Dotted lines are interpreted as cracks in the test, not plate sutures. Dashed lines are inferred (but poorly preserved) plate boundaries.

glacial (=Sangamonian) raised reef (see be- the dominant bioclast. This lithology is low). similar to certain of the well-lithified lime- Description. —See Donovan (1993:381- stone facies of the last interglacial raised 382). reefs of Jamaica that are included within Remarks. —While Echinometra lucunter is the Falmouth Formation sensu late, Upper a common, extant, shallow-water species Coastal Group. However, while fragments in the Caribbean, its fossil record is rela- of Echinometra spp. are locally common in tively poor due to various environmental this unit (Gordon, 1991 ), tests are rare and and taphonomic factors (Gordon, 1991; none are otherwise known from such a Greenstein, 1991; Donovan and Gordon, well-lithified limestone. The alternative 1993). Thus, the precise horizon that yield- possibility is that this specimen is ce- ed this unusually well-preserved specimen mented into beachrock, giving it a Holo- (Fig. lC) is of some interest. The test is cene age. While this is considered less preserved in a small block of porous, grey probable, rare echinoids do occur in beach- limestone (grainstone) in which corals are rock (Donovan et al., 1993). CENOZOIC FOSSIL ECHINOIDS 169

Order CLYPEASTEROIDA A. Agassiz ity. Oligopygus de Loriol is considered to Family CLYPEASTERIDAE L. Agassiz be Eocene (Kier, 1967), while Clypeasfer in Genus Clypeaster Lamarck the Caribbean is Oligocene to Recent (Pod- Clypeaster duchassaingi Michelin, 1861 dubiuk, 1985). (Fig. 2A) Remarks. —While Clypeaster spp. are Material, Locality and Horizon. —A single widespread and common in the fossil rec- test, MCZ 1418. The label reads “Guade- ord of the Caribbean, one of the Cuban loupe. Collected by Duchassaing. L. Ag- specimens is notable in showing obvious assiz Coil. 1859.” The precise locality and growth deformities in all ambulacral petals horizon is unknown, but Mortensen (1948: (Figs. 2B, 3; compare with Fig. 2C). In most 31) regarded this species as Pliocene and instances this has involved a premature it is limited to the “formations madepo- termination of the petal in one ambulacral riques” of Michelin (1861:107; cited by column, followed by a re-initiation of the Jackson, 1922:43). This specimen undoubt- petal more proximally. This is best seen in edly came from Grande-Terre, which is ambulacra IV and I (Figs. 2B, 3A, C). The completely capped by Pliocene-Quater- only petal not effected is V, where the only nary limestones, rather than the igneous break in the normal pattern of growth is a island of Basse-Terre (Andreieff and Cot- slight deflection of one pore pair (Fig. 3B). tez, 1976; Maury et al., 1990). The anterior ambulacrum has one enlarged Remarks. -Jackson (1922:9) noted the dif- and sunken pore (Fig. 2B, 3E). There is no ficulty of obtaining definitive information obvious sign of predation or parasitism, on the fossil echinoids of Guadaloupe and and the reason for these abnormalities is only one test from this island was found not apparent. in the collections of the MCZ. Hence, it Acknowledgments. —I thank Tomasz K. was considered important to figure this Baumiller for his hospitality while I was specimen, despite its somewhat poor pres- visiting the MCZ and Frederick J. Collier ervation. The specimen agrees well with for arranging the loan of these specimens. published diagnoses of C. duchassaingi This paper was written during the period (Jackson, 1922:43; Mortensen, 1948:31), be- of the First Shell Distinguished Research ing domed apically, flattened orally, with Fellowship in Science, which I gratefully petals (admittedly somewhat worn) flush acknowledge. Constructive comments by with the test surface and a well-developed David N. Lewis and an anonymous re- (although somewhat obscured by lime- viewer are gratefully acknowledged. stone) peristome. LITERATURE CITED Clypeaster sp. Andreieff, P., and S. Cottez. 1976. Sur l'age, la struc- (Fig. 2B, C) ture et la formation des iles de Grande-Terre et de Marie Galante (Guadeloupe-FWI). In R. Cauusse, Material, Locality and Horizon. —Collec- (cd.), Trans. 7th Caribb. Geol. Conf., Guadaloupe, tion of 12 tests, labelled “Clypeaster. Oli- 30 June-12 July, 1974, pp. 329-333. BRGM, Or- gocene. Finca Santa Ines. Barrio Majagua, leans. Jatibonico, Camaguey, Cuba. From Mario Arnold, B. W., and H. L. Clark, 1927. Jamaican fossil echini. Mere. Mus. Comp. Zool. Harvard 50:1-75. Sanchez Roig, 1940.” Rocks of the Oligo- —, and —. 1934. Some additional fossil cene outcrop in the region near Jatibonico echini from Jamaica. Mere. Mus. Comp. Zool. Har- and Majagua in western De Avila province vard 54:139-156. (Jimenez et al., 1970), which was part of Bengtson, P. 1988. Open nomenclature. Paleontolo- gy 31:223–227. Camaguey province prior to 1959 (Draper Cotteau, G. H. 1875, Description des Echinides Ter- and Barros, 1994:Fig. 4.1). However, Kier tiaires des iles St. Barthe1emy et Anguilla, K. Sven- (1984:6) noted that many Cuban localities ska Vetensk,-Akad. Handl. 13(6):1-48. that were previously regarded as Oligo- Cutress, B. M. 1980. Cretaceus and Tertiary Cida- cene have now been revised as Miocene. roida (Echinodermata: Echinoidea) of the Carib- bean area. Bull. Am. Paleont. 77(309):1-221, The dating of this sample is further con- Donovan, S. K. 1988. A preliminary biostratigraphy fused by a collection of Oligopygus sp. in of the Jamaican fossil Echinoidea, In R. D. Burke the MCZ, apparently from the same local- et al. (eds.), Echinoderm biology: Proc. 6th Int. 170 S. K. DONOVAN

Echinoderm Conf., Victoria, British Columbia, 23- Maury, R. C., G. K. Westbrook, P. E. Baker, Ph. Bouysse, 28 August, 1987, pp. 125-131. Balkema, Rotterdam. and D. Westercamp. 1990. Geology of the Lesser —. 1991, An Echinolampas (Echinoidea: Cassidu- Antilles. In G. Dengo and J. E. Case (eds.), The loida) marker band in the Newport Formation? J. geology of North America, Vol. H. The Caribbean Geol. Sot. Jamaica 28:43-44. region, pp. 405-432. Geological Society of Amer- —. 1993, Jamaican Cenozoic Echinoidea. In R, ica, Boulder, M. Wright and E. Robinson (eds.), Biostratigraphy Melville, R. V., and J. W. Durham. 1966, Skeletal of Jamaica. Geol. Sot, Am, Mere. 182:371-412. morphology. In R. C. Moore (cd.), Treatise on in- , and C. M. Gordon. 1993. Echinoid taphon- vertebrate paleontology, Part U, Echinodermata omy and the fossil record: supporting evidence 3(l), pp. U220-U251. Geological Society of Amer- from the Plio-Pleistocene of the Caribbean. Palaios ica and University of Kansas Press, New York and 8:304-306. Lawrence. —, R.A. Williams, and J, A. Rocke. 1993. Pres- Michelin, H. 1861, Monographie des Clypeastres ervation of a clypeasteroid echinoid in Holocene fossiles. Mem. Sot. Geo1. France, ser. 2, 7:101-147. beachrock, Jamaica. Caribb. J. Sci. 29:264-267. [Not seen,] Draper, G., and J. A. Barros. 1994. Cuba, In S. K. Mortensen, T. 1948. A monograph of the Echinoi- Donovan and T.A. Jackson (eds.), Caribbean ge- dea. IV. 2. Clypeastroida. Clypeastridae, Arach- ology: an introduction, pp. 65-86. UWI Publishers’ noididae, Fibulariidae, Laganidae and Scutellidae. Association, Kingston. Text. C.A. Reitzel, Copenhagen. 471 pp. Fell, H. B. 1966. Cidaroids. In R. C. Moore (cd,), Poddubiuk, R, H. 1985. Evolution and adaptation in Treatise on invertebrate paleontology, Part U, some Caribbean Oligo-Miocene Clypeasters. In B. Echinodermata 3(l), pp. U312-U339. Geological F. Keegan and B. D. S, O'Connor (eds.), Echinoder- Society of America and University of Kansas Press, mata: Proc. 5th Int. Echinoderm Conf., Galway, New York and Lawrence. 24-29 September, 1984, pp. 19-24. Balkema, Rot- Gordon, C. M. 1991. The poor fossil record of Echi- terdam. nometra (Echinodermata: Echinoidea) in the Carib- Smith, A. B. 1981, Implications of lantern mor- bean region. J. Geol. Sot, Jamaica 28:37-41. phology for the phylogeny of post-Palaeozoic Greenstein, B. J. 1991. An integrated study of echi- echinoids. Paleontology 24:779-801, noid taphonomy: predictions for the fossil record —. 1984. Echinoid palaeobiology. George, Al- of four echinoid families. Palaios 6:519-540. len and Unwin, London, 190 pp. Jackson, R. T. 1922, Fossil echini of the West Indies. —, and C. W. Wright. 1989 (for 1987). British Publ. Carnegie Inst. Wash. 306:1-103. Cretaceus echinoids. Part 1, general introduction Jimenez, A. N., et al. 1970. Atlas national de Cuba. and Cidaroida, Monogr. Palaeontogr. Sot. Lend, Academia de Ciencias de Cuba, La Habana. 143 141(578):1-101. PP. Weisbord, N. E. 1969. Some late Cenozoic Echinoi-

Kier, P. M. 1967. Revision of the oligopygoid echi- dea from Cabo Blancor Venezuela. Bull. Am, Pa- noids, Smithson. Misc. Coil. 152(2):149 pp. leont. 56(252):277-371. —. 1984. Fossil spatangoid echinoids of Cuba, Winsor, M. P. 1991. Reading the shape of nature: Smithson. Contr. Paleobiol. 55:339 pp. comparative zoology at the Agassiz Museum. Uni- Linne, C. 1758. Systems naturae per regna tria na- versity of Chicago Press, Chicago. 324 pp. turae, secundum classes, ordines, genera, species, cum characteribus differentis, synonymis, locis, 10th ed. Holmiae. 824 pp. [Not seen.]