Page 1 of 40 Geological Journal

Page 1 of 32 1 2 3 Neogene echinoids from the Cayman Islands, West Indies: regional 4 5 6 implications 7 8 9 10 1 2 3 11 STEPHEN K. DONOVAN *, BRIAN JONES and DAVID A. T. HARPER 12 13 14 15 1Department of Geology, Naturalis Biodiversity Center, Leiden, the Netherlands 16 17 2Department of Earth and Atmospheric Sciences, University of Alberta, Edmonton, Canada, T6G 2E3 18 For Peer Review 19 3 20 Department of Earth Sciences, Durham University, Durham, UK 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 *Correspondence to: S. K. Donovan, Department of Geology, Naturalis Biodiversity Center, 49 50 Darwinweg 2, 2333 CR Leiden, the Netherlands. 51 52 E-mail: [email protected] 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 2 of 40

Page 2 of 32 1 2 3 The first fossil echinoids are recorded from the Cayman Islands. A regular echinoid, Arbacia? sp., the 4 5 spatangoids Brissus sp. cf. B. oblongus Wright and Schizaster sp. cf. S. americanus (Clark), and the 6 7 clypeasteroid Clypeaster sp. are from the Middle Miocene Cayman Formation. Test fragments of the 8 9 mellitid clypeasteroid, Leodia sexiesperforata (Leske), are from the Late Pleistocene Ironshore 10 11 Formation. Miocene echinoids are preserved as (mainly internal) moulds; hence, all species are left 12 13 14 in open nomenclature because of uncertainties regarding test architecture. All Miocene taxa are 15 16 recorded from single specimens apart from the 27 assigned to Brissus. Schizaster sp. cf. S. 17 18 americanus (Clark) is compared to a species from the Oligocene of the south-east USA. Brissus sp. cf. 19 For Peer Review 20 B. oblongus is close in gross morphology to a taxon from the Miocene of Malta. Leodia 21 22 sexiesperforata is identified from fragments with confidence, being the only extant Antillean sand 23 24 dollar with elongate ambulacral petals that is limited to carbonate substrates. The Miocene 25 26 27 echinoids of Grand Cayman, although of limited diversity, are mainly comprised of genera common 28 29 in comparable mid-Cenozoic carbonate environments. 30 31 32 33 Received 15 December 2014 34 35 36 37 KEY WORDS systematics; biodiversity; Miocene; Pleistocene; Brissus; Schizaster; Clypeaster 38 39 40 41 42 43 44 1. INTRODUCTION 45 46 47 48 In a region renowned for its fossil faunas, it is surprising to note that echinoids have hitherto not 49 50 been recorded from the rock record of the Cayman Islands in the Greater Antilles (Fig. 1). The first 51 52 53 echinoid fossils were only discovered by B.J. and his graduate students in 1995 and further 54 55 specimens have been slow to accumulate. 56 57 Apart from any biogeographic significance, these echinoids are also noteworthy for being 58 59 60 http://mc.manuscriptcentral.com/gj Page 3 of 40 Geological Journal

Page 3 of 32 1 2 3 mainly from the Miocene. This interval was called the ‘age of echinoids’ by Ager (1993, p. 27), but 4 5 this comment was presumably based upon that author’s observations in the Mediterranean region. 6 7 In the Caribbean, perhaps only in the Paleocene are fossil echinoids less well documented than the 8 9 Miocene-Pliocene, with the notable exception of the Miocene of Anguilla (Poddubiuk and Rose, 10 11 1985; Poddubiuk, 1987). Otherwise, echinoids are not a particularly well-known component of the 12 13 14 Miocene fauna of the Antilles (see comments in Donovan et al., 2005). There are historical 15 16 difficulties of correlating Miocene (or Oligocene?) echinoids from Cuba with confidence: “Many of 17 18 the [purported Oligocene] species may be Miocene. In the last 20 years some workers … have placed 19 For Peer Review 20 most of what was thought to be the Cuban Oligocene into the Miocene. Unfortunately, Sánchez Roig 21 22 usually did not state from what formation his echinoids were collected” (Kier, 1984, p. 6). The Lower 23 24 Miocene echinoids of Jamaica are best preserved in allochthonous blocks of coralliferous reef 25 26 27 limestone, and are mainly fragmentary and/or overgrown by calcite (Donovan, 1993, 2004; Donovan 28 29 and Portell, 2000; Donovan et al., 2005). The Dutch ABC islands, Aruba, Bonaire and Curaçao, had a 30 31 small fauna of echinoids described from the Miocene-Pliocene Seroe Domi Formation (de Busonje, 32 33 1974) which has recently been doubled in an unpublished M.Sc. thesis (Schelfhorst, 2013). The 34 35 echinoids of the Middle Miocene Grand Bay Formation of Carriacou, the Grenadines (Donovan, 36 37 2012) and the Miocene-Pliocene August Town Formation of Jamaica (Donovan, research in progress) 38 39 40 similarly await adequate publication. Scattered occurrences of Miocene echinoids are known from 41 42 other islands, for example, Barbados (Donovan and Veale, 1996, table 1). 43 44 45 46 47 48 2. GEOLOGICAL SETTING 49 50

51 52 53 The Cayman Islands are the only sub-aerially exposed parts of the Cayman Ridge, which lies parallel 54 55 to the Oriente Transform Fault that defines the north boundary of the Cayman Trough (see Jones, 56 57 1994, fig. 5.1, for map). This tectonic system was most probably part of the Late Cretaceous to 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 4 of 40

Page 4 of 32 1 2 3 Paleogene magmatic arc (Case et al., 1984, pp. 16-17, sheet 1). Bedrock exposed on each of the 4 5 Cayman Islands is comprised of the Bluff Group, which includes the unconformity bounded Brac 6 7 Formation (Lower Oligocene), Cayman Formation (Middle Miocene) and the Pedro Castle Formation 8 9 (Pliocene), that is unconformably overlain by Pleistocene strata that belong to the Ironshore 10 11 Formation (Fig. 1; Jones et al., 1994a, b). The Ironshore Formation is divided, from oldest to 12 13 14 youngest, into Units A to F (Vézina et al., 1999; Coyne et al., 2007). Fossil echinoids are herein 15 16 recognized for the first time from the Cayman Formation (Locality 1) and Ironshore Formation 17 18 (Locality 2) on Grand Cayman (Fig. 1A). 19 For Peer Review 20 21 22 2.1. Locality 1: Miocene 23 24 The Cayman Formation is composed largely of finely crystalline dolostones that are commonly 25 26 27 fossiliferous with corals, gastropods, red algae, foraminifers and rhodolites being locally numerous 28 29 (Jones and Hunter, 1989; Jones, 1994, p. 89). Any fossils that originally had aragonitic skeletons are 30 31 now represented by moulds. The echinoids described herein (Arbacia? sp., Schizaster sp. cf. S. 32 33 americanus (Clark) and Brissus sp. cf. B. oblongus Wright) were collected from the upper part of the 34 35 Cayman Formation that is exposed in High Rock Quarry, which is located in the east-central part of 36 37 Grand Cayman (Fig. 1). All of these specimens came from the eastern side of the quarry, which is 38 39 40 now weathered and largely covered with grass on the quarry floor, and bushes and trees along the 41 42 quarry walls. Although the age of the Cayman Formation is open to some debate, it is generally 43 44 regarded as being Middle Miocene. 45 46 Poorly preserved echinoids have been noted by B.J. in the Cayman Formation at two other 47 48 localities on Grand Cayman. In Pedro Castle Quarry (Fig. 1A), where the type sections of the Cayman 49 50 Formation and Pedro Castle Formation are located (Jones and Hunter, 1989, fig. 2), one poorly 51 52 53 preserved echinoid, probably Brissus (not collected) was seen in the in large boulder from the 54 55 Cayman Formation that had been blasted from the quarry wall. In a disused quarry just north of 56 57 Spotts (Fig. 1A), an external mould of the apical surface of Clypeaster sp. in a large boulder of 58 59 60 http://mc.manuscriptcentral.com/gj Page 5 of 40 Geological Journal

Page 5 of 32 1 2 3 dolostone from the Cayman Formation was noted in the early 1980s (see below). This quarry is now 4 5 overgrown and inaccessible. 6 7 8 9 2.2. Localities 2A and B: Pleistocene 10 11 Exposures of the Ironshore Formation along the western shoreline of North Sound (Fig. 1A) are 12 13 14 formed of Unit D that is unconformably overlain by Unit F (Coyne et al., 2007). Coral-rich patch reefs, 15 16 well-preserved bivalves and gastropods (Coyne et al., 2007), and oolitic grainstones that are 17 18 characterized by numerous burrows (Pemberton and Jones, 1988) characterize Unit D in this area. 19 For Peer Review 20 This unit, which is 123-147 ka old, formed during the Marine Isotope Stage (MIS) 5e highstand 21 22 (Coyne et al., 2007). Unit F is formed largely of cross-bedded oolitic grainstones that contain few 23 24 fossils. Although dating of this unit is problematic because of the scarcity of well-preserved fossils, 25 26 27 one Th/U date from Unit F suggests it formed 87 ± 3 ka in association with the MIS 5a highstand 28 29 (Coyne et al., 2007). 30 31 Despite extensive searching, no echinoids have ever been found in Unit D, which is widely 32 33 exposed on Grand Cayman and Cayman Brac. At two localities, located to the south of Morgan’s 34 35 Harbour on the west coast of North Sound (2 in Fig. 1), fragments of a mellitid sand dollar, Leodia 36 37 sexiesperforata (Leske), were found in the lower part of the cross-bedded ooid grainstones of unit F. 38 39 40 The only other fossils found in this unit are rare bivalves, conch shells and very rare corals. 41 42 43 44 45 46 3. MATERIALS AND METHODS 47 48 49 50 The Miocene echinoids of the Cayman Islands in the present collection are preserved as natural 51 52 53 moulds, mainly internal moulds. These were painted with black food colouring to give a uniform 54 55 body tone and coated with ammonium chloride for photography. Pleistocene specimens are rarer 56 57 and more fragmentary, but preserve tests. These were similarly coloured and coated with 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 6 of 40

Page 6 of 32 1 2 3 ammonium chloride for photography. Measurements were taken using electronic callipers. 4 5 Descriptive terminology used herein follows Melville and Durham (1966), Durham and 6 7 Wagner (1966) and Smith (1984). The classification follows Kroh and Smith (2010) and Smith and 8 9 Kroh (2011). The use of open nomenclature follows the protocol recommended by Bengtson (1988). 10 11 Fossils specimens are deposited in the Department of Earth and Atmospheric Sciences, University of 12 13 14 Alberta, Edmonton, Canada (UA) and, in the example of one Eocene specimen from Jamaica, the 15 16 Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (MCZ). Recent 17 18 echinoids used in the present study have been presented to the Department of Invertebrate 19 For Peer Review 20 Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). 21 22 The Miocene material of Brissus from the Cayman Islands is moderately well preserved and 23 24 nearly 30 specimens were available for measurement; however, only some 20 presented useful data. 25 26 27 Only two of the 20 specimens yielded a complete measurement set, yet a further 10 of the 28 29 specimens were included in the multivariate analysis, their missing values for some variates being 30 31 interpolated by iterative imputation. The dataset thus included 12 specimens of Brissus from the 32 33 Cayman Islands comprising a near complete data matrix, plus 46 specimens of Brissus from the 34 35 Recent of Jamaica (National Museum of Natural History, Smithsonian Institution) and one specimen 36 37 from the Eocene, the latter probably representing a new species, albeit only known from one 38 39 40 specimen (MCZ 3469; see Donovan and Veale, 1996). The multivariate (Principal Components) shape 41 42 analysis interrogated a correlation matrix of eleven variates (see Donovan and Harper, 2000) 43 44 measured on all 59 specimens: test length (TL); test width (TW); test height (TH); height at apical 45 46 system, measured from apical system to labrum or equivalent height (Tha); distance from apical 47 48 system to anterior (AA); length of petal V (Lv); length of petal IV (Liv); number of pore pairs in outer 49 50 half ambulacrum of petal V (PPv); number of pore pairs in anterior half ambulacrum of petal IV 51 52 53 (PPiv); periproct height (PH); and periproct width (PW). The multivariate analyses were implemented 54 55 on the software program PAST (Hammer et al., 2001; Hammer and Harper, 2006). For ‘Results of 56 57 PCA’, see Brissus sp. cf. B. oblongus, below. 58 59 60 http://mc.manuscriptcentral.com/gj Page 7 of 40 Geological Journal

Page 7 of 32 1 2 3 4 5 6 7 4. SYSTEMATIC PALAEONTOLOGY 8 9 10 11 Class ECHINOIDEA Leske, 1778 12 13 14 Subclass EUECHINOIDEA Bronn, 1860 15 16 Order ARBACIOIDA Gregory, 1900 17 18 Family ARBACIIDAE Gray, 1855 19 For Peer Review 20 Genus Arbacia Gray, 1835 21 22 23 24 Type species. Cidaris pustulosa Leske, 1778, p. 150 [=Echinus lixula Linné, 1758, p. 664], by 25 26 27 subsequent designation of Agassiz and Clark (1908, p. 67; Smith and Kroh, 2011). 28 29 Diagnosis. See Smith and Kroh (2011). 30 31 Remarks. The sole internal mould of a regular echinoid (Fig. 2) from this site provides less 32 33 diagnostic data than for either of the spatangoids. 34 35 Range. Miocene to Recent, Europe, North Africa, North and South America; Recent, Atlantic, 36 37 Caribbean, West Pacific (Smith and Kroh, 2011). 38 39 40 41 42 43 44 Arbacia? sp. 45 46 Figure 2 47 48 49 50 Material. A single test, UA P1669, preserved as an internal mould. 51 52 53 Locality and horizon. Middle Miocene Cayman Formation, Grand Cayman (Locality 1). 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 8 of 40

Page 8 of 32 1 2 3 Description. Internal mould of regular echinoid. Circular outline, low domed profile, ambitus 4 5 below mid-height of test. Apical system rounded pentagonal in outline. Three lozenge-like genital 6 7 plates apparent, but circlet imperfectly preserved. 8 9 Ambulacra narrower than interambulacra. Details of ambulacral plating not apparent, 10 11 ambulacral plates moderately high. Ambulacra preserved as paired, diverging ridges on the apical 12 13 14 surface, but it impossible to determine confidently if it these are pore pairs in columns or columns of 15 16 tubercles immediately adjacent to ambulacra as in Arbacia. Pore pairs not distinguishable. 17 18 Interambulacral plates larger (both higher and wider) than ambulacral plates, with a small central 19 For Peer Review 20 tubercle. Peristome obscured by rock. 21 22 Remarks. The gross morphology of this specimen, which preserves too few fine details, is 23 24 nonetheless essential for determining its probable systematic position. It is certainly neither a 25 26 27 cidaroid nor a diadematoid, both of which have bun-like tests with the ambitus at about mid-height. 28 29 It is not close to the more elliptical test of Echinodermetra Gray. Rather, it is more like to be an 30 31 arbacioid or a toxopneustid. Assuming the specimen to be an adult, the test diameter (<20 mm) is 32 33 more in the range of modern Antillean Arbacia punctulata (Lamarck) or Lytechinus williamsi Chesher 34 35 (albeit with a more pentagonal test) than other toxopneustids (Hendler et al., 1995, pp. 214-222, fig. 36 37 134A-G). Assuming it to be fully grown, it is tentatively classified as an Arbacia? sp. because this is an 38 39 40 Antillean Neogene genus that is at least close in gross morphology to the test, although it is more 41 42 inflated than, for example, Pliocene Arbacia improcera (Conrad) (Lewis and Donovan, 1991). 43 44 45 46 Cohort IRREGULARIA Latreille, 1825 47 48 Order SPATANGOIDA L. Agassiz, 1840 49 50 Suborder PALEOPNEUSTINA Markov and Solovjev, 2001 51 52 53 Family SCHIZASTERIDAE Lambert, 1905 54 55 Genus SCHIZASTER L. Agassiz, 1835 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 9 of 40 Geological Journal

Page 9 of 32 1 2 3 Type species. Schizaster studeri L. Agassiz, 1835, p. 185, by the subsequent designation of 4 5 the International Commission on Zoological Nomenclature (1954, p. 388) (Fischer, 1966, p. U569). 6 7 Diagnosis. (After Smith and Kroh, 2011.) Test ovate with deep anterior sulcus; slightly 8 9 pointed to rear. Apical disc ethmolytic, with four gonopores. Anterior ambulacrum deeply sunken; 10 11 pore-pairs in adapical portion large and specialized. Other ambulacra also deeply sunken, anterior 12 13 14 petals longer and more flexed than posterior petals. Periproct small and marginal, on near-vertical 15 16 truncate face. Bound by interambulacral plates 5a/b on oral side. Peristome opening facing anterior; 17 18 kidney-shaped. Labral plate short and wide; not extending beyond half-way along the first 19 For Peer Review 20 ambulacral plate; in broad contact with sternal plates. Sternal plates large and symmetric; sternal- 21 22 episternal suture at rear of ambulacral plate 5. Aboral tuberculation fine, uniform and dense. Oral 23 24 tubercles also dense and uniform. Well-developed peripetalous and lateral fascioles. Peripetalous 25 26 27 fasciole indented by three plates behind anterior petals; crosses ambulacrum III at plate 7 or 8. 28 29 Latero-anal fasciole branches off about one-third up from the ends of the anterior petals on plates 30 31 1.b.5 and 4.a.5 32 33 Remarks. Schizaster is a common echinoid taxon in the Antillean region from the Eocene to 34 35 Recent (Kier and Grant 1965; Kier 1984; Donovan, 1993), particularly the former. 36 37 Range. Eocene to Recent (Fischer, 1966, p. U569). 38 39 40 41 42 Schizaster sp. cf. S. americanus (Clark in Clark and Twitchell, 1915) 43 44 Figure 3 45 46 47 48 Material. A single test, UA P1670, preserved as an internal mould. 49 50 Locality and horizon. Middle Miocene Cayman Formation, Grand Cayman (Locality 1). 51 52 53 Description. Preserved as an internal mould in dolostone. Test globular, about 27.2 mm long, 54 55 27.5 mm wide and 22.5 mm high; highest posterior of centre, widest about centre. Four genital 56 57 pores, but apical system poorly preserved otherwise. Peristome anterior, but concealed by 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 10 of 40

Page 10 of 32 1 2 3 dolostone; periproct rounded (elliptical?) on concave posterior surface and elevated, in upper half of 4 5 test. Oral surface with central keel. Anterior ambulacrum (III) not petaloid, depressed, moderately 6 7 broad and forming a shallow, albeit distinct anterior sulcus. Short, sunken petals developed apically. 8 9 Anterior ambulacra (II, IV) with about 10-12 pore pairs per petal, petal length c. 8.1 mm. Posterior 10 11 petals (I, V) shorter with about 7 pore pairs per ambulacral column, length c. 3.8 mm. Ambitus low 12 13 14 with largest interambulacral plates in 1 and 4. Plastron moderately broad and shield-like. External 15 16 features of test unknown. 17 18 Remarks. The Cayman Islands Miocene Schizaster is considered closest to the Oligocene 19 For Peer Review 20 Schizaster americanus (Clark) from the south-east USA, described and illustrated by Clark and 21 22 Twitchell (1915, p. 176, pl. 82, figs 3A-D), Cooke (1959, p. 72, pl. 30, figs 5-8) and Oyen and Portell 23 24 (2002, pl. 2, figs G, H). The Cayman specimen, apart from being younger, is relatively more inflated 25 26 27 and highest at the apical system, whereas the holotype of S. americanus is highest posterior to the 28 29 apical system. The apical system of the holotype is within a keel in the posterior interambulacrum 30 31 (5), although this may be an artefact produced by the test being slightly crushed laterally (see, in 32 33 particular, Cooke, 1959, pl. 30, fig. 6). Certainly, the geometry of the ambulacral petals, positions of 34 35 periproct and peristome, and posterior concavity of the test is similar between S. americanus and UA 36 37 P1670. 38 39 40 Miocene Schizaster species from Cuba are less similar to UA P1670. Schizaster egozcue 41 42 Lambert has a wedge-shaped gross morphology, and petals II and IV are distinctly closer to 43 44 ambulacrum III; Schizaster fernandezi Sánchez Roig, like S. egozcue, has a more posterior apical 45 46 system, and also longer petals II and IV in a relatively narrower test (Kier, 1984, pl. 18). Similar 47 48 aspects of gross morphology also differentiate Mio-Oligocene S. rojasi Sánchez Roig and S. 49 50 sanctamariae Sánchez Roig from UA P1670 (Kier, 1984, pl. 25). Schizaster sp. from the Upper 51 52 53 Pliocene of Jamaica is too poorly preserved for worthwhile comparison (Donovan and Portell, 1998). 54 55 Poddubiuk and Rose (1985, table 3) recognized two species of Schizaster from the Lower 56 57 Miocene Anguilla Formation of Anguilla, namely S. clevei Cotteau, 1875, and S. loveni Cotteau, 1875. 58 59 60 http://mc.manuscriptcentral.com/gj Page 11 of 40 Geological Journal

Page 11 of 32 1 2 3 Schizaster clevei is strongly heart-shaped, unlike the rounded UA P1670; compare Fig. 3 herein with 4 5 Cotteau (1875, pl. 5, figs 7, 8). Schizaster loveni is a little more rounded than S. clevei (Cotteau, 1875, 6 7 pl. 5, figs 9-13), with a posterior keel behind the apical system and, in larger specimens, moderately 8 9 long anterior petals curving towards ambulacrum III. 10 11

12 13 14 Suborder BRISSIDINA Stockley et al., 2005 15 16 Family BRISSIDAE Gray, 1855 17 18 Genus BRISSUS Gray, 1825 19 For Peer Review 20 21 22 Type species. Spatangus brissus unicolor Leske, 1778, p. 248, by the subsequent designation 23 24 of International Commission on Zoological Nomenclature (1954, p. 387) (Fischer, 1966, p. U582). 25 26 27 Diagnosis. (After Donovan and Veale, 1996, p. 635; based on Mortensen, 1951, p. 506-508; 28 29 Fischer, 1966, p. U582-U583; Kier, 1984, p. 81.) Test elongate ovoid in outline, moderate to large as 30 31 adult. Test more or less highly arched aborally; posterior interambulacrum may be raised as a keel. 32 33 Oral surface flattened to greatly convex. Anterior sinus not developed, posterior truncate. Apical 34 35 system anterior ethmolytic (Smith, 1984, fig. 3.22), with four genital pores (genital 5 absent); 36 37 posterior pores larger than anterior. Anterior ambulacrum narrow, not petaloid. Paired ambulacral 38 39 40 petals sunken; anterior petals approximately transverse, posterior petals not widely divergent. First 41 42 plate of interambulacrum 1 followed by single plate. Plastron large, ultramphisternous. Peripetalous 43 44 and subanal fascioles developed; subanal fasciole broad with lateral lobes. Periproct lenticular in 45 46 outline, on upper part of truncate posterior margin. Peristome near anterior, semilunate, with 47 48 broad, but not prominent, labrum; conspicuous phyllodes developed. Tuberculation dense. 49 50 Pedicellariae usually of five types; globiferous, tridentate, rostrate, ophicephalous and triphyllous. 51 52 53 Remarks. Since the `Modern' echinoid fauna of the Caribbean region first emerged in the 54 55 Eocene-Oligocene, certain genera have shown a propensity for extreme plasticity in overall form 56 57 (most notably Clypeaster spp.), while others have shown near-stasis in gross morphological features 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 12 of 40

Page 12 of 32 1 2 3 (such as the holectypoid Echinoneus cyclostomus group and the spatangoid Brissus unicolor group). 4 5 The influence of environment, dispersion and ecophenotypic variation on these evolutionary 6 7 patterns is unknown. 8 9 Range. Eocene to Recent (Fischer, 1966, p. U583). 10 11

12 13 14 Brissus sp. cf. B. oblongus Wright, 1855 15 16 Figure 4 17 18 19 For Peer Review 20 Material. Twenty seven specimens, UA P1671-P1695 (complete or incomplete internal 21 22 moulds), UA P1696, P1697 (partial external moulds). 23 24 Locality and horizon. Middle Miocene Cayman Formation, Grand Cayman (Locality 1). 25 26 27 Description. Inflated, elliptical Brissus, rounded anteriorly, but more broadly in larger tests 28 29 (Fig. 4A, D, G, K); widest about centre to posterior of centre; flattened posteriorly; oral surface 30 31 convex; no anterior sulcus. Anterior of test wedge-shaped, steeply sloping (Fig. 4E, F, H, I, O); test 32 33 highest about two-thirds to three-quarters of test length from anterior. Posterior interambulacrum 34 35 (5) raised as a keel with a rounded ridge between the posterior petals (Fig. 4B, C, L, M). Ambitus low, 36 37 rounded and sloping forwards (Fig. 4E, H, O). 38 39 40 Apical system poorly preserved, but anterior of centre. Four genital pores apparent. 41 42 Peristome on anterior oral surface, close to anterior margin, kidney-shaped (Fig. 4D, P), 43 44 moderately broad, sunken and with a raised posterior margin; that is, the peristome is directed 45 46 anteriorly. Periproct on flattened posterior margin at about mid-height, lensoid and higher than 47 48 wide (best seen in Fig. 4C). 49 50 Four ambulacral petals on apical surface; anterior ambulacrum (III) not petaloid (Fig. 4A, G, 51 52 53 K). Petals sunken, short, not extending to ambitus. Anterior paired petals (II and IV) short, extending 54 55 in a near-straight line laterally on either side of the apical system, petals straight to slightly curved. 56 57 Posterior paired petals (I and V) longer, narrowly diverging, straight or curving away from 58 59 60 http://mc.manuscriptcentral.com/gj Page 13 of 40 Geological Journal

Page 13 of 32 1 2 3 interambulacrum 5. Ambulacral pores of petals rounded, arranged in two closely-spaced columns of 4 5 pore pairs. Anterior ambulacrum (III) narrow, flush with test surface, narrower than petals. 6 7 Interambulacra broader than ambulacra, but details not preserved. 8 9 Ambulacra converge to peristome on oral surface; single pored adjacent to peristome. 10 11 Plastron (interambulacrum 5) with closely packed, asymmetrical tubercles (Fig. 4N), flanked by 12 13 14 curved ambulacra I and V. 15 16 Given the mouldic preservation, fine details of the test surface, particularly tuberculation 17 18 and distribution of fascioles, are not apparent except on the two partial external moulds and 19 For Peer Review 20 specimen UA P1675 which preserves the plastron. These show densely packed primary tubercles, 21 22 but no fascioles are discernible. They were considered too friable for casting. 23 24 Results of PCA. The specimens were plotted against the first and second principal 25 26 27 components or eigenvectors (Fig. 6). Whereas the first discriminated the specimens with respect to 28 29 size, the specimen from the Eocene of Jamaica being by far the largest, the second represents 30 31 variation in shape. Four variates have marked contributions to the second eigenvectors, PPV, PIV, PH 32 33 and PV (see Materials and Methods, above). Specimens with positive scores of the second 34 35 eigenvector have relatively larger values for the first two variates and smaller for values for the 36 37 second two. Thus, the cluster of specimens from the Cayman islands (BS prefix), including the 38 39 40 Eocene specimen (EOC), that have negative scores on the second eigenvector, are characterized by 41 42 relative large values for PH and PV (the size of the periproct) and smaller values for the numbers of 43 44 PPV and PIV, the pore pairs in the ambulacral region of petals V and VI. 45 46 Remarks. Species of Brissus show considerable similarity in time and space. For example, 47 48 Chesher (1972) compared Brissus spp. from either side of the Isthmus of Panama. Biometric analysis 49 50 showed that specimens from the Bay of Panamá (Brissus obesus Verrill) and the tropical Western 51 52 53 Atlantic (Brissus unicolor (Leske)) differed in the number of plates between the mouth and posterior 54 55 petals, and the position of the lateral portion of the peripetalous fasciole. Specimens from the Gulf 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 14 of 40

Page 14 of 32 1 2 3 of California, however, were unexpectedly within the range of morphology of B. unicolor and not B. 4 5 obesus. 6 7 The features used by Chesher (1972) cannot be determined for many fossil specimens, 8 9 which may preserve details of the test surface only poorly or, as in the case of the Cayman 10 11 specimens, almost not at all. Donovan and Veale (1996) used a range of biometric criteria to 12 13 14 compare Recent (Fig. 5A, B) and fossil Brissus from the Antillean region, demonstrating that gross 15 16 test morphology has remained similar at least since the Middle Eocene. The statistical analysis used 17 18 herein (Fig. 6) is built on that of Donovan and Harper (2000), who described a Lower Miocene(?) 19 For Peer Review 20 Brissus from Jamaica. 21 22 Comparison of the Cayman taxon with the extant species from the shallow waters of the 23 24 Caribbean, Brissus unicolor (Leske) (Fig. 5A, B), shows a number of dissimilarities. These include the 25 26 27 relatively longer petals of B. unicolor, its more divergent posterior petals, more sinuous anterior 28 29 petals, less rounded sides and more inflated test. A closer comparison with the Miocene species 30 31 from Malta, Brissus oblongus Forbes MSS in Wright, 1855, suggests that it may be closely related to 32 33 the Cayman species. The holotype is certainly similar in gross test outline to the smaller Cayman 34 35 specimens (compare Figs. 5C, D, with 4G, K). The Maltese taxon has longer anterior petals and 36 37 relatively more divergent posterior petals, but, overall, is closer to the Cayman specimens than B. 38 39 40 unicolor and other Antillean Brissus spp. Other nominal Brissus spp. from the Miocene of Malta 41 42 (Rose, 1975) are either synonymous with or markedly different in morphology from B. oblongus. 43 44 The closest Brissus from the Miocene of the Antilles to the Cayman species is Brissus exiguus 45 46 Cotteau, 1875 (pp. 35-36, pl. 6, figs 16-18), but the illustrated specimens are more teardrop-shaped 47 48 in plan view than any specimen in Fig. 4. Kier (1984) discussed five species of Brissus from the 49 50 Cenozoic of Cuba, most of which are broad, inflated and blunt anteriorly. Closest to the Cayman 51 52 53 species is Brissus caobaense Sánchez Roig, 1953 (Kier, 1984, pp. 83-84, fig. 31, pl. 44, figs 1-4), which 54 55 has a broadly similar profile, although not as markedly high in the posterior interambulacrum on the 56 57 apical surface, is blunter anteriorly and posteriorly, has a particularly large periproct and a more 58 59 60 http://mc.manuscriptcentral.com/gj Page 15 of 40 Geological Journal

Page 15 of 32 1 2 3 hexagonal outline in apical view. The blunt anterior is a feature also seen in the lateral views of three 4 5 extant species of Brissus, including B. unicolor, illustrated by Schultz (2005, fig. 722); all have a 6 7 relatively higher and more steeply sloping anterior than the Cayman species. It may be, of course, 8 9 that the Cayman tests represent a new species. Given the overall similarity of all Brissus spp., it is 10 11 considered best not to erect a new nominal taxon based on internal moulds. 12 13 14 15 16 17 18 Superorder MICROSTOMATA Smith, 1984 19 For Peer Review 20 Order CLYPEASTEROIDA L. Agassiz, 1835 21 22 Suborder SCUTELLINA Haeckel, 1896 23 24 Family MELLITIDAE Stefanini, 1912 25 26 27 Genus LEODIA Gray, 1851 28 29 30 31 Type species. Leodia richardsoni Gray, 1851, p. 36 (= Echinodiscus sexiesperforatus Leske, 32 33 1778, p. 199) by original designation (Durham, 1966, p. U485; Mooi, 1989, p. 41; Smith and Kroh, 34 35 2011). 36 37 Diagnosis. See Smith and Kroh (2011). 38 39 40 Remarks. This genus was monotypic until the description of Leodia divinata Mooi and 41 42 Peterson, 2000, from the Lower Pliocene of Venezuela. 43 44 Range. Pliocene to Recent, Cape Hatteras, North Carolina, to Florida Keys, Central and South 45 46 America, and the Antilles (Mooi, 1989, p. 41; Hendler et al., 1995, pp. 234-235). 47 48 49 50 Leodia sexiesperforata (Leske, 1778) 51 52 53 Figure 7 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 16 of 40

Page 16 of 32 1 2 3 Material. UA P1698-P1700, Locality 2A; UA P1701-P1704, Locality 2B. All specimens 4 5 fragmentary. 6 7 Locality and horizon. Unit F, Pleistocene Ironshore Formation, Grand Cayman (Localities 2A 8 9 and B). 10 11 Description. See Hendler et al. (1995, pp. 234-235 and references therein). 12 13 14 Remarks. Comparison with other Late Pleistocene sites in the Antilles strongly indicates that 15 16 these fragments, undoubtedly a mellitid, are derived from an extant taxon. At the present day, the 17 18 mellitids of the Gulf of Mexico and Antilles include only four species, all shallow water (Serafy, 1979; 19 For Peer Review 20 Hendler et al., 1995). There are two species of Encope L. Agassiz, 1840, have ambulacral notches, not 21 22 lunules, and these can be easily disregarded. Mellita isometra Harold and Telford, 1990, has four 23 24 long, slender ambulacral lunules, but prefers siliceous sediment substrates (Mooi, 1989, p. 41; 25 26 27 Hendler et al., 1995, p. 236, fig. 126). Only Leodia sexiesperforata has five long, slender ambulacral 28 29 lunules and a similar anal lunule, and is restricted to carbonate substrates (Mooi, 1989, p. 41). Extant 30 31 L. sexiesperforata occurs in 0-60 m water depth and only on carbonate substrates (Serafy, 1979, 32 33 table 2; Mooi, 1989, p. 41). 34 35 36 37 Suborder CLYPEASTERINA L. Agassiz, 1835 38 39 40 Family CLYPEASTERIDAE L. Agassiz, 1835 41 42 Genus CLYPEASTER Lamarck, 1801 43 44 45 46 Type species. Echinus rosaceus Linné, 1758, p. 665, by the subsequent designation of 47 48 Desmoulins (1835, p. 183) (Durham, 1966, p. U462). 49 50 Diagnosis. See Smith and Kroh (2011). 51 52 53 Remarks. Clypeaster is the commonest and most easily recognizable echinoid genus in the 54 55 Oligocene and Neogene of the Antilles, in part because of the test which is particularly robust. 56 57 Fragments are easily identified to genus and, in some instances, species (see, for example, Dixon and 58 59 60 http://mc.manuscriptcentral.com/gj Page 17 of 40 Geological Journal

Page 17 of 32 1 2 3 Donovan, 1998). 4 5 Range. Late Eocene to Recent, widespread in tropical to temperate regions (Smith and Kroh, 6 7 2011); first appeared in the Antillean region in the Late(?) Oligocene (Poddubiuk, 1985). 8 9 10 11

12 13 14 Clypeaster sp. 15 16 Figure 8 17 18 19 For Peer Review 20 Material. External mould of the apical surface of one test, not collected (Fig. 8). 21 22 Locality and horizon. Spotts Bay Quarry, Grand Cayman. Middle Miocene Cayman Formation, 23 24 Grand Cayman, now overgrown. 25 26 27 Remarks. Poddubiuk (1985) rationalized the 60+ nominal species of Clypeaster described 28 29 from the Upper Oligocene-Lower Miocene strata of the Antilles to just seven names. The specimen 30 31 from Grand Cayman is close to Clypeaster concavus Cotteau, 1875 (Cotteau, 1875, p. 16, pl. 2, figs 4- 32 33 8; Jackson, 1922, pp. 34-36, pl. 2, figs 10-12) and descriptions of Clypeaster cubensis Cotteau, 1875, 34 35 although good illustrations of the latter species are unavailable (but note the comment of Jackson, 36 37 1922, p. 37, that it “… has very large petals”). However, an identification to genus based on an image 38 39 40 of a partial external mould is considered adequate in this instance. 41 42 43 44 45 46 5. DISCUSSION 47 48 49 50 Until now, fossil echinoids have not been described from the Cayman Islands, an anomaly if 51 52 53 compared to the wealth of nominal taxa known from, for example, nearby Cuba and Jamaica. 54 55 Although specimens described herein are either mouldic (Miocene) or fragmentary (Pleistocene), 56 57 three of the five species have been confidently assigned to genus and a fourth identified to species. 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 18 of 40

Page 18 of 32 1 2 3 Most taxa are Miocene. Apart from the Lower Miocene Anguilla Formation of Anguilla, echinoids 4 5 from the other Antillean islands have either been recognized as Oligocene or Miocene, requiring 6 7 stratigraphic revision (such as Cuba; Kier, 1984), or are Miocene-Pliocene, such as the Seroe Domi 8 9 Formation of Aruba, Bonaire and Curaçao (Jackson and Robinson, 1994), and August Town 10 11 Formation of Jamaica (Robinson, 1994) (see comments in Donovan et al., 2005, pp. 91-92). 12 13 14 Comparison with other mid-Cenozoic echinoid-bearing sites and horizons from Malta and 15 16 the Antilles is revealing (Table 1). Most of these records are based on multiple sites collected by 17 18 echinoderm specialists over many years, with the exception of the Lower Miocene of Jamaica (Table 19 For Peer Review 20 1, site 5). Even the latter essentially samples more than one location; well-preserved echinoids 21 22 mainly come from allochthonous blocks (shallow water), with different taxa occurring as fragments 23 24 in the enclosing chalks (deep water). But four species from two localities (Locality 1, Spotts Bay 25 26 27 Quarry) is certainly comparable with six from the Lower Miocene of Jamaica. Indeed, S.K.D.’s 28 29 experience of collecting echinoids from the Upper Oligocene of Jamaica (Table 1, Site 1) and Antigua 30 31 (Site 2) suggests that four to six taxa from any one site is good. 32 33 Multiple taphonomic and collector biases lie behind the data compiled in Table 1. The 34 35 Miocene of Grand Cayman has only yielded moulds and fragmentary remains are unknown. The mid- 36 37 Cenozoic echinoids of Antigua, Anguilla and Malta are, in contrast, well-preserved, locally common 38 39 40 and moderately easy to collect, and have been the subject of numerous research programmes since 41 th 42 at least the mid-19 Century (for example, Wright, 1855, was working on Maltese specimens 160 43 44 years ago). The Jamaican Oligocene echinoids occur in well-indurated limestones and were largely 45 46 ignored until the work of the late Hal Dixon of the University of the West Indies, Mona, in the 1990s 47 48 (Dixon and Donovan, 1998). Similarly, the Lower Miocene of the same island was collected and 49 50 prepared by Roger Portell and colleagues of the Florida Museum of Natural History, University of 51 52 53 Florida, Gainesville, including fragments and tests in crystal apple preservation (that is, each plate is 54 55 overgrown by a single calcite crystal; see Donovan and Portell, 2000, for explanation; Donovan et al., 56 57 2005). That is, the better preserved localities have been collected and described for at least 160 58 59 60 http://mc.manuscriptcentral.com/gj Page 19 of 40 Geological Journal

Page 19 of 32 1 2 3 years, whereas the less well-preserved faunas, such as that of the Cayman Formation, have only 4 5 been examined for about the 25 years we have looked for new localities and new ways to study 6 7 fossil echinoids. 8 9 None of the other Sites (1-5) in Table 1 includes an Arbacia, so the tentative assignment of 10 11 UA P1669 to this genus remains uncertain, although the (dissimilar) arbacioid Coelopleurus is known 12 13 14 from Malta and Pliocene Arbacia is well-known from the Antilles (Lewis and Donovan, 1991; 15 16 Donovan and Paul, 1998). In contrast, the irregular echinoids identified from Grand Cayman are 17 18 widely distributed, particularly Clypeaster. Again, a taphonomic bias is easily determined, as 19 For Peer Review 20 Clypeaster undoubtedly has the most robust test of any echinoid genus; there is no wonder that it is 21 22 known from all six sites (Table 1). More surprisingly, relatively more delicate taxa, Schizaster and 23 24 Brissus, have been found at most sites and, where they are unknown, they may unknowingly be 25 26 27 represented by indeterminate spatangoid remains (such as Donovan et al., 2005, p. 108, pl. 8). 28 29 The Pleistocene record of the Antilles is enriched by the identification of L. sexiesperforata 30 31 from the Ironshore Formation. Sand dollars are poorly known from the Pleistocene of the Antilles 32 33 (Jackson, 1922, p. 52; Donovan and Embden, 1996, p. 490; Donovan, 2001, p. 184), yet are common 34 35 in coeval deposits in Florida (Donovan, 2003, table 1). Leodia sexiesperforata was previously 36 37 unknown from the Pleistocene of the Caribbean islands and other genera, namely Encope and 38 39 40 Mellita, included only taxa left in open nomenclature. As Donovan (2003, p. 5) noted, clypeasteroids 41 42 are a common component of many Eocene to Pliocene successions of the Antilles and include taxa 43 44 with tests that are especially robust (Smith, 1984, p. 23; Nebelsick, 1994, 1995, 1999; Nebelsick and 45 46 Kroh, 2002). 47 48 It may be that Pleistocene clypeasteroids were living further offshore than the carbonate 49 50 successions that are currently available for study. For example, the only horizons that are rich in 51 52 53 sand dollars from the Jamaican Pliocene-Pleistocene are demonstrably allochthonous (storm) 54 55 deposits (Donovan et al., 1994a, b). The presence of L. sexiesperforata only in the cross-bedded unit 56 57 F of the Ironshore Formation (see above) may similarly demonstrate at least some local transport. It 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 20 of 40

Page 20 of 32 1 2 3 is unlikely to have been derived from a siliciclastic environment (and thus be M. isometra) in the 4 5 carbonate platform setting of the Cayman Islands. 6 7 8 9 10 11 ACKNOWLEDGEMENTS 12 13 14 15 16 Financial assistance enabling fieldwork by B.J. and his students was provided by a Natural Sciences 17 18 and Engineering Research Council of Canada operating grant which is gratefully acknowledged. We 19 For Peer Review 20 thank the Photographic Unit of the Natural History Museum, London, for the images in Figs. 2-4 and 21 22 7. S.K.D. thanks three institutions for their support during the lengthy gestation of this paper, namely 23 24 the University of the West Indies, Mona, the Natural History Museum, London, and the Naturalis 25 26 27 Biodiversity Center, Leiden. Comprehensive reviews by David N. Lewis (the Natural History Museum, 28 29 London) and Jeffery R. Thompson (University of Southern California, Los Angeles) are gratefully 30 31 acknowledged. 32 33 34 35 36 37 REFERENCES 38 39 40 41 42 Agassiz, A., Clark, H.L. 1908. Hawaiian and other Pacific Echinii. The Salenidae, Arbaciadae, 43 44 Aspidodiadematidae, and Diadematidae. Memoirs of the Museum of Comparative Zoology, 45 46 Harvard College 34, 43-132. 47 48 Agassiz, L. 1835. Prodrome d’une Monographie des Radiares ou Echinodermes. Mémoires de la 49 50 Société des Sciences Naturelles de Neufchâtel 1, 168-199. 51 52 53 Agassiz, L. 1840. Catalogus systematicus Ectyporum Echinodermatum fossilium Musei Neocomiensis, 54 55 secundum ordinem zoologicum dispositus; adjectis synonymis recentioribus, nec non stratis et 56 57 locis in quibus reperiuntur. Sequuntur characters diagnostici generum novorum vel minus 58 59 60 http://mc.manuscriptcentral.com/gj Page 21 of 40 Geological Journal

Page 21 of 32 1 2 3 cognitorum. Petitpierre: Neuchâtel; 20 pp. 4 5 Ager, D.V. 1993. The Nature of the Stratigraphical Record. Third edition. Wiley: Chichester; xiv+151 6 7 pp. 8 9 Bengtson, P. 1988. Open nomenclature. Palaeontology 31, 223-227. 10 11 Bronn, H.G. 1860. Die Klassen und Ordnungen des Thier-Reichs, wissenschaftlich dargestellt in Wort 12 13 14 und Bilt. Zweiter Band. Actinozoen. C.F. Winter’sche Verlagshandlung: Leipzig and Heidelberg; 15 16 434 pp. 17 18 Buisonjé, P.H. de. 1974. Neogene and Quaternary geology of Aruba, Curaçao and Bonaire 19 For Peer Review 20 (Netherlands Antilles). Uitgaven Natuurwetenschappelijke Studiekring voor Suriname en de 21 22 Nederlandse Antillen 78, 1-291. 23 24 Case, J.E., Holcombe, T.L., Martin, R.G. 1984. Map of geologic provinces in the Caribbean region. In: 25 26 27 The Caribbean-South American Plate Boundary and Regional Tectonics, Bonini, W.E., Hargraves, 28 29 R.B., Shagam, R. (eds). Geological Society of America Memoir 162, 1-30. 30 31 Chesher, R.H. 1972. The status of knowledge of Panamanian echinoids, 1971, with comments on 32 33 other echinoderms. In: The Panamic Biota: Some Observations Prior to a Sea-Level Canal, Jones, 34 35 M.L. (ed.). Bulletin of the Biological Society of Washington 2, 139-158. 36 37 Clark, W.B., Twitchell, M.W. 1915. The Mesozoic and Cenozoic Echinodermata of the United States. 38 39 40 Monographs of the U.S. Geological Survey 4, 1-341. 41 42 Cooke, C.W. 1959. Cenozoic echinoids of eastern United States. U.S. Geological Survey Professional 43 44 Paper 321, 1-106. 45 46 Cotteau, G.H. 1875. Description des echinides Tertiaires des îles St. Barthélemy et Anguilla. Kongliga 47 48 Svenska Vetenskaps-Akademiens Handlingar 13 (6), 1-47. 49 50 Coyne, M.K., Jones, B., Ford, D. 2007. Highstands during Marine Isotope Stage 5: evidence from the 51 52 53 Ironshore Formation of Grand Cayman, British West Indies. Quaternary Science Reviews 26, 536- 54 55 559. 56 57 Desmoulins, C. 1835. Premier mémoire sur les Échinides. Prodrome dúne nouvelle classification des 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 22 of 40

Page 22 of 32 1 2 3 ces animaux. Actes de la Société Linnéenne de Bordeux 7, 167-245. 4 5 Dixon, H.L., Donovan, S.K. 1998. Oligocene echinoids of Jamaica. Tertiary Research 18, 95-124. 6 7 Donovan, S.K. 1993. Jamaican Cenozoic Echinoidea. In: Biostratigraphy of Jamaica, Wright, R.M., 8 9 Robinson, E. (eds). Geological Society of America Memoir 182, 371-412. 10 11 Donovan, S.K. 2001. Evolution of Caribbean echinoderms during the Cenozoic: moving towards a 12 13 14 complete picture using all of the fossils. Palaeogeography, Palaeoclimatology, Palaeoecology 15 16 166, 177-192. 17 18 Donovan, S.K. 2003. Completeness of a fossil record: the Pleistocene echinoids of the Antilles. 19 For Peer Review 20 Lethaia 36, 1-7. 21 22 Donovan, S.K. 2004. Echinoderms of the mid-Cainozoic White Limestone Group of Jamaica. In: The 23 24 Mid-Cainozoic White Limestone Group of Jamaica, Donovan, S.K. (ed.). Cainozoic Research 3 (for 25 26 27 2003), 143-156. 28 29 Donovan, S.K. 2012. Taphonomic criteria for recognizing downslope transport of echinoderms: 30 31 Neogene of the Antilles. Geological Society of America Abstracts with Programs 44 (7), 529. 32 33 Donovan, S.K., Embden, B.J. 1996. Early Pleistocene echinoids of the Manchioneal Formation, 34 35 Jamaica. Journal of Paleontology 70, 485-493. 36 37 Donovan, S.K., Harper, D.A.T. 2000. The irregular echinoid Brissus Gray from the Tertiary of Jamaica. 38 39 40 Caribbean Journal of Science 36, 332-335. 41 42 Donovan, S.K., Paul, C.R.C. 1998. Echinoderms of the Pliocene Bowden shell bed, southeast Jamaica. 43 44 In: The Pliocene Bowden shell bed, southeast Jamaica, Donovan, S.K. (ed.). Contributions to 45 46 Tertiary and Quaternary Geology, 35: 129-146. 47 48 Donovan, S.K., Portell, R.W. 1998. A spatangoid echinoid from the Upper Pliocene of southeast 49 50 Jamaica. Caribbean Journal of Science 34, 320-322. 51 52 53 Donovan, S.K., Portell, R.W. 2000. Incipient ‘crystal apples’ from the Miocene of Jamaica. Caribbean 54 55 Journal of Science 36, 168-170. 56 57 Donovan, S.K., Veale, C. 1996. The irregular echinoids Echinoneus Leske and Brissus Gray in the 58 59 60 http://mc.manuscriptcentral.com/gj Page 23 of 40 Geological Journal

Page 23 of 32 1 2 3 Cenozoic of the Antillean region. Journal of Paleontology 70, 632-640. 4 5 Donovan, S.K., Dixon, H.L., Littlewood, D.T.J., Milsom, C.V., Norman, Y.J.C. 1994a. The 6 7 clypeasteroid echinoid Encope homala Arnold and Clark, 1934, in the Cenozoic of Jamaica. 8 9 Caribbean Journal of Science 30, 171-180. 10 11 Donovan, S.K., Dixon, H.L., Pickerill, R.K., Doyle, E.N. 1994b. Pleistocene echinoid (Echinodermata) 12 13 14 fauna from southeast Jamaica. Journal of Paleontology 68, 351-358. 15 16 Donovan, S.K., Portell, R.W., Veltkamp, C.J. 2005. Lower Miocene echinoderms of Jamaica, West 17 18 Indies. Scripta Geologica 129, 91-135. 19 For Peer Review 20 Durham, J.W. 1966. Clypeasteroids. In: Treatise on Invertebrate Paleontology, Part U, 21 22 Echinodermata 3 (2), Moore, R.C. (ed.). Geological Society of America and University of Kansas 23 24 Press: New York and Lawrence; U450-U491. 25 26 27 Durham, J.W., Wagner, C.D. 1966. Glossary of morphological terms applied to echinoids. In: Treatise 28 29 on Invertebrate Paleontology, Part U, Echinodermata 3 (1), Moore, R.C. (ed.). Geological Society 30 31 of America and University of Kansas Press: New York and Lawrence; U251, U253-U256. 32 33 Fischer, A.G. 1966. Spatangoids. In: Treatise on Invertebrate Paleontology, Part U, Echinodermata 34 35 3(2), Moore, R.C. (ed.). Geological Society of America and University of Kansas Press: New York 36 37 and Lawrence; U543-U628. 38 39 40 Gray, J.E. 1825. An attempt to divide the Echinida or sea eggs, into natural families. Annals of 41 42 Philosophy (series 2) 10, 423-431. 43 44 Gray, J.E. 1835. On the genera distinguishable in Echinus. Proceedings of the Zoological Society of 45 46 London 3, 57-60. 47 48 Gray, J.E. 1851. Description of two new genera and some new species of Scutellidae and 49 50 Echinolampidae in the collection of the British Museum. Proceedings of the Zoological Society of 51 52 53 London 19, 34-38. 54 55 Gray, J.E. 1855. An arrangement of the families of Echinida, with descriptions of some new genera 56 57 and species. Proceedings of the Zoological Society, London 23, 35-39. 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 24 of 40

Page 24 of 32 1 2 3 Gregory, J.W. 1900. The Echinoidea. In: A Treatise on Zoology. Part III. The Echinodermata, 4 5 Lankester, E.R. (ed.). A. and C. Black: London; 282-332. 6 7 Haeckel, E. 1896. Systematische Phylogenie der wirbellosen Thiere (Invertebrata). Vol. 2. George 8 9 Reimer Verlag: Berlin; xviii + 720 pp. 10 11 Hammer, Ø., Harper, D.A.T. 2006. Palaeontological Data Analysis. Blackwell, Oxford, 351 pp. 12 13 14 Hammer, Ø., Harper, D.A.T., Ryan, P.D. 2001. PAST – palaeontological statistics software: package 15 16 for education and data analysis. Palaeontologia Electronica 4, 9 pp. 17 18 Harold, A.S., Telford, M. 1990. Systematics, phylogeny and biogeography of the genus Mellita 19 For Peer Review 20 (Echinoidea: Clypeasteroida). Journal of Natural History 24, 987-1026. 21 22 Hendler, G., Miller, J.E., Pawson, D.L., Kier, P.M. 1995. Sea Stars, Sea Urchins, and Allies: 23 24 Echinoderms of Florida and the Caribbean. Smithsonian Institution Press: Washington, D.C.; 25 26 27 xi+390 pp. 28 29 International Commission on Zoological Nomenclature. 1954. Opinion 209. Validation of, and 30 31 designation of type species for, “Brissus” Gray, 1825, “Echinocardium” Gray, 1825, and 32 33 “Spatangus” Gray, 1825 (Class Echinoidea) under the plenary powers, and designation, under 34 35 those powers, of a type species for “Schizaster” Agassiz (L.), 1836, and in so far as necessary, for 36 37 “Moira” Agassiz (A.), 1872. Opinions and Declarations Rendered by the International Commission 38 39 40 on Zoological Nomenclature 3(28), 367-392. 41 42 Jackson, R.T. 1922. Fossil Echini of the West Indies. Carnegie Institution of Washington, Publication 43 44 306, iv+122. 45 46 Jackson, T.A., Robinson, E. 1994. The Netherlands and Venezuelan Antilles. In: Caribbean Geology: 47 48 An Introduction, Donovan, S.K., Jackson, T.A. (eds). University of the West Indies Publishers’ 49 50 Association: Kingston; 249-263. 51 52 53 Jones, B. 1994. The Cayman Islands. In: Caribbean Geology: An Introduction, Donovan, S.K., Jackson, 54 55 T.A. (eds). University of the West Indies Publishers’ Association: Kingston; 87-109. 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 25 of 40 Geological Journal

Page 25 of 32 1 2 3 Jones, B., Hunter, I.G. 1989. The Oligocene-Miocene Bluff Formation on Grand Cayman. Caribbean 4 5 Journal of Science 25, 71-85. 6 7 Jones, B., Hunter, I.G., Kyser, K. 1994a. Stratigraphy of the Bluff Formation (Miocene-Pliocene) and 8 9 the newly defined Brac Formation (Oligocene), Cayman Brac, British West Indies. Caribbean 10 11 Journal of Science 30, 30-51. 12 13 14 Jones, B., Hunter, I.G., Kyser, K. 1994b. Revised stratigraphic nomenclature for Tertiary strata of the 15 16 Cayman Islands, British West Indies. Caribbean Journal of Science 30, 53-68. 17 18 Kier, P.M. 1984. Fossil spatangoid echinoids of Cuba. Smithsonian Contributions to Paleobiology 55, 19 For Peer Review 20 336 pp. 21 22 Kier, P.M., Grant, R.E. 1965. Echinoid distribution and habits, Key Largo Coral Reef Preserve, Florida. 23 24 Smithsonian Miscellaneous Collections 149 (6), 1-68. 25 26 27 Kroh, A., Smith, A.B. 2010. The phylogeny and classification of post-Palaeozoic echinoids. Journal of 28 29 Systematic Palaeontology 8, 147–212. 30 31 Lamarck, J.B.P.M. de. 1801. Système des animaux sans vertèbres, ou Tableau général des classes, 32 33 des ordres et des genres des ces animaux. Deterville: Paris; 432 pp. 34 35 Lambert, J.M. 1905. Échinides du sud de la Tunise environs de Tatahouine. Bulletin de la Société 36 37 Géologique de France (série 4) 5, 569-577. 38 39 40 Latreille, P.A. 1825. Famille naturelles de Régne Animal, exposées succinctement et dans un ordre 41 42 analytique avec l’indication des leurs genres. Paris; 570 pp. 43 44 Leske, N.G. 1778. Iacobi Theodori Klein Naturalis disposito Echinodermatum, Edita et aucta a N.G. 45 46 Leske. Lipsiae; 278 pp. 47 48 Lewis, D.N., Donovan, S.K. 1991. The Pliocene Echinoidea of Tobago, West Indies. Tertiary Research 49 50 12, 139-146. 51 52 53 Linné, C. 1758. Systema Naturae per Regna tria Naturae, secundum Classes, Ordines, Genera, 54 55 Species, cum Characteribus Differentis, Synonymis, Locis. Holmiae. 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 26 of 40

Page 26 of 32 1 2 3 Markov, A.V., Solovjev, A.N. 2001. Echinoids of the family Paleopneustidae (Echinoidea, 4 5 Spatangoida): morphology, taxonomy, phylogeny. Geos 2001, 1-109. 6 7 Melville, R.V., Durham, J.W. 1966. Skeletal morphology. In: Treatise on Invertebrate Paleontology, 8 9 Part U, Echinodermata 3 (1), Moore, R.C. (ed.). Geological Society of America and University of 10 11 Kansas Press: New York and Lawrence; U220-U251. 12 13 14 Mooi, R. 1989. Living and fossil genera of the Clypeasteroida (Echinoidea: Echinodermata): an 15 16 illustrated key and annotated checklist. Smithsonian Contributions to Zoology 488, iii+51 pp. 17 18 Mooi, R., Peterson, D. 2000. A new species of Leodia (Clypeasteroida: Echinoidea) from the Neogene 19 For Peer Review 20 of Venezuela and its importance in the phylogeny of mellitid sand dollars. Journal of 21 22 Paleontology 74, 1083-1092. 23 24 Mortensen, T. 1951. A Monograph of the Echinoidea. V. 2. Spatangoida. II. Amphisternata. II. 25 26 27 Spatangidae, Loveniidae, Pericosmidae, Schizasteridae, Brissidae. Text and atlas. C.A. Reitzel: 28 29 Copenhagen; 593 pp. 30 31 Nebelsick, J.H. 1994. Taphonomy of Clypeaster humulis and Echinodiscus auritus (Echinoidea, 32 33 Clypeasteroida) from the Red Sea. In: Echinoderms through Time: Proceedings of the 8th 34 35 International Echinoderm Conference, Dijon, 6-10 September, 1993, David, B., Guille, A., Feral, J.- 36 37 P., Roux, M. (eds). Rotterdam: A.A. Balkema; 803-808. 38 39 40 Nebelsick, J.H. 1995. Comparative taphonomy of clypeasteroids. Eclogae Geologicae Helvetiae 88, 41 42 685-693. 43 44 Nebelsick, J.H. 1999. Taphonomy of Clypeaster fragments: preservation and taphofacies. Lethaia 32, 45 46 241-252. 47 48 Nebelsick, J.H., Kroh, A. 2002. The stormy path from life to death assemblages: the formation and 49 50 preservation of mass accumulations of fossil sand dollars. Palaios 17, 378-393. 51 52 53 Oyen, C.W., Portell, R.W. 2002. Oligocene and Miocene echinoids. Florida Fossil Invertebrates 2, 1- 54 55 22. 56 57 Pemberton, S.G., Jones, B. 1988. Ichnology of the Pleistocene Ironshore Formation, Grand Cayman, 58 59 60 http://mc.manuscriptcentral.com/gj Page 27 of 40 Geological Journal

Page 27 of 32 1 2 3 British West Indies. Journal of Paleontology, 62, 495-505. 4 5 Poddubiuk, R.H. 1985. Evolution and adaptation in some Caribbean Oligo-Miocene Clypeasters. In: 6 7 Echinodermata: Proceedings of the Fifth International Echinoderm Conference, Galway, 24-29 8 9 September, 1984, Keegan, B.F., O'Connor, B.D.S. (eds). Rotterdam: A.A. Balkema; 19-24. 10 11 Poddubiuk, R.H. 1987. Sedimentology, echinoid palaeoecology and palaeobiogeography of Oligo- 12 13 14 Miocene eastern Caribbean limestone (in 2 volumes). Unpublished Ph.D. thesis, Royal Holloway 15 16 and Bedford College, University of London, 419 + 148 pp. 17 18 Poddubiuk, R.H., Rose, E.P.F. 1985. Relationships between mid-Tertiary echinoid faunas from the 19 For Peer Review 20 central Mediterranean and eastern Caribbean and their palaeobiogeographic significance. 21 22 Annales Géologiques des Pays Hélleniques 32 (for 1984), 115-127. 23 24 Robinson, E. 1994. Jamaica. In: Caribbean Geology: An Introduction, Donovan, S.K., Jackson, T.A. 25 26 27 (eds). University of the West Indies Publishers’ Association: Kingston; 111-127. 28 th 29 Rose, E.P.F. 1975. Oligo-Miocene echinoids of the Maltese islands. VI Congress Regional Committee 30 31 on Mediterranean Neogene Stratigraphy, Bratislava, 75-79. 32 33 Sánchez Roig, M. 1953. Neuvos equinodos fosiles de Cuba. Anales de la Academis de Ciencias 34 35 36 Medicas, Fisicas y Naturales de la Habana 91, 135-176. 37 38 Schelfhorst, R. 2013. The Echinodermata of the Pliocene of Curaçao. Unpublished M.Sc. 39 40 thesis, University of Leiden. 41 42 43 Schultz, H. 2005. Sea Urchins: A Guide to Worldwide Shallow Water Species. Heinke and 44 45 Peter Schultz Partner: Hemdingen, Germany; xii+484 pp. 46 47 Serafy, D.K. 1979. Echinoids (Echinodermata: Echinoidea). Memoirs of the Hourglass Cruises 48 49 50 5 (3), 1-120. 51 52 Smith, A.B. 1984. Echinoid Palaeobiology. George Allen and Unwin: London; 191 pp. 53 54 Smith, A.B., Kroh, A. (eds) 2011. The Echinoid Directory. World Wide Web electronic publication. 55 56 http://www.nhm.ac.uk/research-curation/projects/echinoid-directory [accessed 21 August 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 28 of 40

Page 28 of 32 1 2 3 2014]. 4 5 Stefanini, G. 1912. Osservazioni sulla distribuzione geografica, sulla origini e sulla filogenesi degli 6 7 Scutellidae. Bolletino della Societe Geologica Italiana 30 (for 1911): 739-754. 8 9 Stockley, B., Smith, A.B., Littlewood, D.T.J., Lessios, H.A., MacKenzie-Dodds, J.A. 2005. 10 11 Phylogenetic relationships of spatangoid sea urchins (Echinoidea), taxon sampling density and 12 13 14 congruence between morphological and molecular estimates. Zoologica Scripta 34, 447-468. 15 16 Vézina, J.L., Jones, B., Ford, D.C. 1999. Sea-level highstands over the last 500,000 years: evidence 17 18 from the Ironshore Formation on Grand Cayman, British West Indies. Journal of Sedimentary 19 For Peer Review 20 Research 69, 317-327. 21 22 Wright, T. 1855. On fossil echinoderms from the island of Malta; with notes on the stratigraphical 23 24 distribution of the fossil organisms of the Maltese beds. Annals and Magazine of Natural History 25 26 27 (series 2) 15, 101-127, 175-196, 262-277. 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 29 of 40 Geological Journal

Page 29 of 32 1 2 3 Table 1. A comparison of some mid-Cenozoic echinoid occurrences in carbonate sequences from the 4 5 Antilles and Malta. Key: 1 = Upper Oligocene of central north Jamaica (Dixon and Donovan, 1998; 6 7 Donovan, 2004); 2 = Upper Oligocene of Antigua (Poddubiuk and Rose, 1985, table 1); 3 = Oligocene- 8 9 Miocene of Malta (Rose, 1975, table 12); 4 = Lower Miocene of Anguilla (Poddubiuk and Rose, 1985, 10 11 table 3); 5 = Lower Miocene of central north Jamaica (Donovan et al., 2005); 6 = Middle Miocene of 12 13 14 Grand Cayman (herein); + = present; ? = presence uncertain. 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 30 of 40

Page 30 of 32 1 2 3 Table 1. 4 5 1: U. 2: U. 3: Oligo- 4: L. 5: L. 6: 6 Oligo. Oligo. Miocene Miocene Miocene Miocene 7 Jamaica Antigua Malta Anguilla Jamaica Cayman 8 CIDAROIDA 9 Histocidaris + 10 Phyllacanthus + + 11 Prionocidaris + + + + + 12 Stylocidaris? + 13 Tretocidaris + 14 ARBACIOIDA 15 Arbacia? + 16 Coelopleurus + 17 CAMARODONTA 18 Arbacina + 19 ECHINOIDA For Peer Review 20 Echinometra + + + 21 Psammechinus + + 22 Schizechinus + 23 Tripneustes + 24 regular echinoids indet. + 25 ECHINONEIOIDA 26 Echinoneus + + + + 27 CLYPEASTEROIDA 28 Clypeaster + + + + ? + 29 Echinocyamus + 30 Scutella + 31 Sismondia + + 32 CASSIDULOIDA 33 Apatopygus + 34 Echinolampas + + + + 35 Pliolampas + 36 Studeria + 37 SPATANGOIDA 38 Agassizia + + 39 Antillaster + 40 Brissopsis + + 41 Brissus + + + 42 Eupatagus + + + + 43 Hemiaster + 44 Heterobrissus + 45 Lovenia + + + 46 Meoma + + + 47 Pericosmus + + 48 Schizaster + + + + 49 Spatangus + 50 Trachypatagus + 51 spatangoid indet. + + 52 TOTALS 8 11 26 17 6 4 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 31 of 40 Geological Journal

Page 31 of 32 1 2 3 FIGURE CAPTIONS 4 5 6 7 Figure 1. (A) Geological map of Grand Cayman (modified after Jones et al., 1994b, fig. 1). Key: * = 8 9 numbered fossil locality. Note that Localities 2A and 2B are close together; greater detail can be 10 11 12 gleaned from Coyne et al. (2007, fig. 3). (B) Schematic stratigraphic succession of the Cayman 13 14 Islands, showing stratigraphic units, dominant lithologies, fossil biota and nature of fossil 15 16 preservation (after Jones et al., 1994b, fig. 2). Key: VC = very common; C = common; LC = locally 17 18 common; R = rare. 19 For Peer Review 20 21 22 Figure 2. Arbacia? sp., UA P1669, Middle Miocene, Cayman Formation, Grand Cayman. Internal 23 24 25 mould of regular echinoid. (A) Lateral view. (B) Apical view. (C) Oral view. Scale bar represents 10 26 27 mm. Specimen whitened with ammonium chloride. 28 29 30 31 Figure 3. Schizaster sp. cf. S. americanus (Clark in Clark and Twitchell, 1915), UA P1670, Middle 32 33 Miocene, Cayman Formation, Grand Cayman. Internal mould of irregular echinoid. (A) Anterior view. 34 35 (B) Apical view. (C) Right lateral view, anterior to right. (D) Oral view. (E) Posterior view. Scale bar 36 37 38 represents 10 mm. Specimen whitened with ammonium chloride. 39 40 41 42 Figure 4. Brissus sp. cf. B. oblongus Wright, 1855, Middle Miocene, Cayman Formation, Grand 43 44 Cayman. Internal moulds of irregular echinoids. (A-E) UA P1671, apical, anterior, posterior, oral and 45 46 right lateral views. (F, K) UA P1672, right lateral and apical views. (G-I, L, M, P) UA P1678, apical, left 47 48 lateral, right lateral, posterior, anterior and oral views. (J, O) UA P1673, oral and left lateral views. 49 50 51 (N) UA P1675, oral view, preserving part of test; note tuberculation of plastron. Scale bar represents 52 53 10 mm. Specimen whitened with ammonium chloride. 54 55 56 57 Figure 5. (A, B) Brissus unicolor (Leske, 1778), USNM E44054 (two specimens), Recent, the 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 32 of 40

Page 32 of 32 1 2 3 Palisadoes, parish of St Andrew, Jamaica (after Donovan and Veale, 1996, figs 4.1, 4.2). Scale bar 4 5 represents 10 mm. Specimens painted with dark food colouring and whitened with ammonium 6 7 chloride. (A) Apical surface of test #1. (B) Oral surface of test #2. (C, D) Brissus oblongus Wright, 8 9 1855, holotype (after Wright, 1855, pl. 5, figs 2a, b, respectively). (C) Apical surface. (D) Oral surface. 10 11 Presumed x 1. 12 13 14 15 16 Figure 6. The multivariate (Principal Components) shape analysis, interrogated a correlation matrix 17 18 of eleven variates (see Donovan and Harper, 2000) measured on 59 specimens. The specimens were 19 For Peer Review 20 plotted against the first and second principal components (eigenvectors). The following fields are 21 22 discriminated: Recent Brissus unicolor (Leske, 1778) from Jamaica, Brissus sp. cf. B. oblongus Wright, 23 24 1855, and a single test, Brissus sp. nov.(?), from the Eocene of Jamaica (Donovan and Veale, 1996, 25 26 27 pp. 635-638, figs 2.5, 2.6; MCZ 3469). 28 29 30 31 Figure 7. Leodia sexiesperforata (Leske, 1778), UA P1704, Pleistocene Ironshore Formation, oral 32 33 surface of incomplete test, orientation uncertain. Both ambulacral lunules are of similar size and 34 35 shape, but that at 10 o’clock is partly occluded by limestone. Scale bar represents 10 mm. Specimen 36 37 whitened with ammonium chloride. 38 39 40 41 42 Figure 8. Clypeaster sp., photograph taken in the field of an external mould of the apical surface. 43 44 Spotts Bay Quarry, Grand Cayman. Middle Miocene Cayman Formation, Grand Cayman. Lens cap for 45 46 scale. 47 48 49 50

51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 33 of 40 Geological Journal

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 Figure 1. (A) Geological map of Grand Cayman (modified after Jones et al., 1994b, fig. 1). Key: * = 47 numbered fossil locality. Note that Localities 2A and 2B are close together; greater detail can be gleaned 48 from Coyne et al. (2007, fig. 3). (B) Schematic stratigraphic succession of the Cayman Islands, showing 49 stratigraphic units, dominant lithologies, fossil biota and nature of fossil preservation (after Jones et al., 50 1994b, fig. 2). Key: VC = very common; C = common; LC = locally common; R = rare. 51 150x213mm (300 x 300 DPI) 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 34 of 40

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 Figure 2. Arbacia? sp., UA P1669, Middle Miocene, Cayman Formation, Grand Cayman. Internal mould of 31 regular echinoid. (A) Lateral view. (B) Apical view. (C) Oral view. Scale bar represents 10 mm. Specimen 32 whitened with ammonium chloride. 33 110x75mm (300 x 300 DPI) 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 35 of 40 Geological Journal

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 32 33 Figure 3. Schizaster sp. cf. S. americanus (Clark in Clark and Twitchell, 1915), UA P1670, Middle Miocene, 34 Cayman Formation, Grand Cayman. Internal mould of irregular echinoid. (A) Anterior view. (B) Apical view. 35 (C) Right lateral view, anterior to right. (D) Oral view. (E) Posterior view. Scale bar represents 10 mm. 36 Specimen whitened with ammonium chloride. 37 142x111mm (300 x 300 DPI) 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 36 of 40

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 Figure 4. Brissus sp. cf. B. oblongus Wright, 1855, Middle Miocene, Cayman Formation, Grand Cayman. 47 Internal moulds of irregular echinoids. (A-E) UA P1671, apical, anterior, posterior, oral and right lateral 48 views. (F, K) UA P1672, right lateral and apical views. (G-I, L, M, P) UA P1678, apical, left lateral, right 49 lateral, posterior, anterior and oral views. (J, O) UA P1673, oral and left lateral views. (N) UA P1675, oral 50 view, preserving part of test; note tuberculation of plastron. Scale bar represents 10 mm. Specimen 51 whitened with ammonium chloride. 52 140x204mm (300 x 300 DPI) 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 37 of 40 Geological Journal

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 Figure 5. (A, B) Brissus unicolor (Leske, 1778), USNM E44054 (two specimens), Recent, the Palisadoes, 41 parish of St Andrew, Jamaica (after Donovan and Veale, 1996, figs 4.1, 4.2). Scale bar represents 10 mm. 42 Specimens painted with dark food colouring and whitened with ammonium chloride. (A) Apical surface of 43 test #1. (B) Oral surface of test #2. (C, D) Brissus oblongus Wright, 1855, holotype (after Wright, 1855, pl. 44 5, figs 2a, b, respectively). (C) Apical surface. (D) Oral surface. Presumed x 1. 150x150mm (300 x 300 DPI) 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 38 of 40

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 Figure 6. The multivariate (Principal Components) shape analysis, interrogated a correlation matrix of eleven 32 variates (see Donovan and Harper, 2000) measured on 59 specimens. The specimens were plotted against the first and second principal components (eigenvectors). The following fields are discriminated: Recent 33 Brissus unicolor (Leske, 1778) from Jamaica, Brissus sp. cf. B. oblongus Wright, 1855, and a single test, 34 Brissus sp. nov.(?), from the Eocene of Jamaica (Donovan and Veale, 1996, pp. 635-638, figs 2.5, 2.6; MCZ 35 3469). 36 198x140mm (300 x 300 DPI) 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Page 39 of 40 Geological Journal

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 Figure 7. Leodia sexiesperforata (Leske, 1778), UA P1704, Pleistocene Ironshore Formation, oral surface of 37 incomplete test, orientation uncertain. Both ambulacral lunules are of similar size and shape, but that at 10 38 o‘clock is partly occluded by limestone. Scale bar represents 10 mm. Specimen whitened with ammonium 39 chloride. 40 92x82mm (300 x 300 DPI) 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj Geological Journal Page 40 of 40

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 For Peer Review 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 Figure 8. Clypeaster sp., photograph taken in the field of an external mould of the apical surface. Spotts Bay 47 Quarry, Grand Cayman. Middle Miocene Cayman Formation, Grand Cayman. Lens cap for scale. 48 113x166mm (240 x 240 DPI) 49 50 51 52 53 54 55 56 57 58 59 60 http://mc.manuscriptcentral.com/gj