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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.

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