Formation of the Isthmus of Panama

The formation of the Isthmus of Panama stands as one of the greatest natural events of the Cenozoic, driving profound biotic transformations on land and in the oceans. Some recent studies suggest that the Isthmus formed manymillions of years earlier than the widely recognized age of approximately 3 million years ago (Ma), a result that if true would revolutionize our understanding of environmental, ecological, and evolutionary change across the Americas. To bring clarity to the question of when the Isthmus of Panama formed, we provide an exhaustive review and reanalysis of geological, paleontological, and molecular records. These independent lines of evidence converge upon a cohesive narrative of gradually emerging land and constricting seaways,withformationof theIsthmus of Panama sensustricto around 2.8 Ma. The evidence used to support an older isthmus is inconclusive, and we caution against the uncritical acceptance of an isthmus before the Pliocene.Facultad de Ciencias Naturales y Muse

Mellita L. Agassiz 1841

<i>Mellita</i> L. Agassiz, 1841 <p> <i>Des Scutelles. Monographies des Echinoderms Vivans et Fossiles. Monograph 2</i>, 1–151.</p> <p> <b>Type species.</b> <i>Echinodiscus quinquiesperforatus</i> Leske, 1778 (= <i>Mellita testudinata</i> Klein, 1734), by subsequent designation of Pomel, 1883.</p> <p> <b>Assigned species.</b> (<i>sensu</i> Coppard <i>et al</i>. 2013) <i>M. quinquiesperforata</i> (Leske, 1778), <i>M. tenuis</i> Clark, 1940 (= <i>M. isometra</i> Harold & Telford, 1990), <i>M. notabilis</i> Clark, 1947 (= <i>M. kanakoffi</i> Durham, 1961, = <i>M. eduardobarrosoi</i> Caso, 1981).</p> <p> <b>Distribution.</b> Early Pliocene to Recent. Caribbean Sea, tropical and subtropical Atlantic and Pacific coasts of the Americas.</p>Published as part of <i>Coppard, Simon E., 2016, A new genus of mellitid sand dollar (Echinoidea: Mellitidae) from the eastern Pacific coast of the Americas, pp. 158-166 in Zootaxa 4111 (2)</i> on page 160, DOI: 10.11646/zootaxa.4111.2.4, <a href="http://zenodo.org/record/256474">http://zenodo.org/record/256474</a&gt

Metalia Gray 1855

Metalia Gray, 1855 Catalogue ot the recent Echinida, or sea eggs, in the collection ot the British Museum. Pt 1. Echinida Irregularia. Lon- don, by order of the Trustees, p. 51. Type species: Spatangus sternalis Lamarck, 1816, by original designation. Assigned species: Mortensen (1951 a) lists seven Recent species (M. dicrana Clark, 1917; M. latissma Clark, 1925; M. nobilis Verrill, 1867; M. robillardi (de Loriol, 1876); M. spatagus (Linnaeus, 1758); M. sternalis (Lamarck, 1816); M. townsendi Bell, 1904) in this genus, and raises doubts as to whether any fossil form can without question be attributed to this genus. Distribution: Miocene of Egypt to Recent, Indo-Pacific. Brissalius gen. nov. (gender: masculine) Type species: Brissalius vannoordenburgi gen. nov., sp. nov. here designated (gender: masculine) Figs 1, 2 and 9. Diagnosis: Test outline broadly-ovate with indented anterior lateral ambulacra and sunken anterior ambulacrum forming a shallow frontal notch. Broadest point, halfway along the anterior petals, vertex mid-way along the posterior petals. Apical system ethmolytic, central, with four gonopores. Petals small, compact, the anterior pair slowly flex outwards, the posterior pair confluent for two thirds of their length proximally, the last third flexing outwards, with only the six distal pore-pairs of the inner series developed. The plastron is amphisternous, the labrum having a narrow posterior prolongation reaching the end of the first adjoining ambulacral plate. Peripetalous, anal and subanal fascioles are well developed. The subanal fasciole is shieldshaped resulting in a single sanitary funnel. Clavulae of the peripetalous and subanal fascioles have a ‘crown’ distally. Lobed funnel building tube feet are present in the anterior ambulacrum within the peripetalous fasciole and also within the subanal fasciole. Penicillate tube feet present only around the peristome. Plastron primary spines distinctly spatulate with sloping collars. Primary spines slender, curved, and ridged, widening and flattening dorso-ventrally distally. Miliary spines on both the oral and aboral surface only slightly curved, while those inside the peripetalous fasciole have a relatively broad shaft and a globular head with serrated ridges. Typical spatangoid rostrate, ophicephalous and triphyllous pedicellariae are present. Fanged openbladed globiferous pedicellariae occur in large numbers on the posterior of the oral ambulacra, fanged fistulate globiferous pedicellariae present in large numbers around the peristome with simple fistulate (with a round foramen and an arc of small teeth) occuring in small numbers on the oral ambulacra. Narrow-valved and spatulate tridentate pedicellariae present on the oral surface with a terminal-toothed form present adapically. Holotype: NHM 2008.618 Natural History Museum, London. Paratype: NHM 2008.619 Natural History Museum, London. Etymology: This new genus has characters of both Brissopsis and Metalia which is reflected in the name by combining ‘Briss’ and ‘alius’ (masculine of the nominative ‘ alia ’, with ‘ alius ’ meaning other or changed). The species is named after Henk van Noordenburg who brought this new species to my attention. Type locality: Off Siquijor Island in the Philippines (depth 200 m). Description: The test is of medium size (at the widest points the holotype measures 38 mm (length) by 32 mm (width) by 19 mm (height at vertex), the plating thin and delicate. The outline is broadly-ovate (Fig. 1 A), with slightly indented anterior lateral ambulacra and sunken anterior ambulacrum forming a shallow frontal notch (anterior sulcus). The broadest point of the test is halfway along the anterior petals (20 % of the way along the length of the test) with the greatest height (vertex) halfway along the posterior petals. The posterior is truncated with an almost oblique vertical face, rounding adorally, protruding at the midpoint of the subanal fasciole (Fig. 1 C & D). The periproct is small and ellipitical (Fig. 1 F) being partially visible from above (Fig. 1 A), but not from below (Fig. 1 B). The anterior of the test rounds to a short, slightly blunt face. The oral surface is convex and steeply angled forming a distinct keel (Fig. 1 F); while the sides of the test are regularly arched aborally towards the sunken petal area (Fig. 1 E & F). The anterior ambulacrum (Fig. 1 A) is narrow, non-petaloid and sunken. The pore-pairs are differentiated with lobed tube feet present aborally. Adapically the pore-pairs are uniserial with a perradial granular zone. The anterior and posterior paired ambulacra are petaloid (Fig. 1 A), narrow, of equal length and closed distally. The anterior pair slowly flex outwards for two-thirds of their length distally, the outer series at an angle of 25 degrees, increasing to 60 degrees for the last third of their length. The inner series are more greatly curved with only eight pore-pairs developed distally. The posterior pair being confluent for two thirds of their length proximally, the last third flexing outwards, with only the six distal pore-pairs of the inner series developed. The pores are elongated and non-conjugate, reducing in size towards the apical system. The outer pores in the inner series of the anterior petals are larger than inner pores, being almost equal in the outer series. The pores in both series of the posterior petals are approximately equal in size. A few irregular miliary tubercles are present along the proximal edge of the distal ambulacral plates of the posterior petals (Fig. 1 A). All other ambulacral plates within the pore-zones are smooth. Outside the pore-zones the anterior ambulacral plates have miliary tubercles within the plates and the occasional small primary tubercle adradially. The aboral plates of interambulacra 1–4 are each raised centrally, forming a faint keel (Fig. 1 A). The posterior interambulacrum 5 is slightly concave sloping gently towards the posterior edge. Tuberculation on interambulacra 1–4 is predominantly uniform aborally with all tubercles perforate and weakly crenulate and set in horizontal series. In interambulacrum 5 a naked zone is present down the midline to the periproct with only miliary tubercles present. Larger primary tubercles are present within the peripetalous fasciole, particularly on the third and fourth plates of the posterior interambulacrum and around the surrounding plates of the front edge of the anterior petals. On the oral surface (Fig. 1 B) and around the periproct (Fig. 1 F), the primary tubercles are enlarged but less abundant. The sternum (Fig. 1 F & G) is densely tuberculated in radiating series, the tubercles decreasing in size towards a central ridge which terminates in a posterior protrusion (Fig. 1 C & D). The edges of the sternal system, the ambulacra and the area around the peristome are broadly naked (Fig. 2 B). The plastron (Figs 1 B & 2 B) is amphisternous, the labrum having a narrow posterior prolongation reaching the end of the first adjoining ambulacral plate. The anterior edge is smooth forming a very slightly protruding lip. The peristome is situated anteriorly, has moderately developed surrounding phyllodes and is crescentshaped. The apical system is situated centrally (Fig. 1 A), has four genital pores and is ethmolytic with genital plate 2 elongate separating both the posterior genital plates (G 4 & G 1) and posterior ocular plates (OcV & OcI). The anterior genital pores are smaller and closer together than the proximal pores, which are situated on a ridge that connects interambulacra 1 and 4. The peripetalous fasciole crosses distally the non-petaloid ambulacrum but stays close to the petals on the other ambulacra. On the interambulacra the peripetalous fasciole curves only gently inwards, not following the direct line of the petals. The subanal fasciole is shield-shaped (Figs 1 F & G, 2 F & G) with typically four pore-pairs in each side but no distinct radiating furrows. The band of clavulae is thinnest at the lower part, becoming broader towards the periproct. A relatively broad (compared to other Brissopsis) band of clavulae forms an anal fasciole (Figs 1 A & 2 A) connecting the subanal fasciole with the peripetalous fasciole. The denuded test and spines are cream/white, while in life the dermis is typically beige/light-brown. Many of the specialised tube feet and pedicellariae have a black skin, visible as black flecks on the test. Primary spines on the aboral surface are slender, curved (both laterally and dorso-ventrally), widening and flattening slightly dorso-ventrally distally (Fig. 9 Ki & Kii). The spines have smooth longitudinal ridges which increase in width distally, terminating in slightly rounded tips. The neck of the spine is straight (not pinched) while the collar is prominent, sloping slightly upwards towards the dorsal surface of the spine. Primary spines within the peripetalous fasciole are shorter, with less curved shafts and broader, more flattened distal regions (Fig. 9 Ji & Jii). Primary spines on the oral surface (Fig. 9 Ni & Nii) are similar to those aborally, but some have recurved shafts (curving in different directions proximally and distally) with more flattened and angular distal regions. Primary spines on the plastron (see Fig. 2 B, D & F) are distinctly spatulate (spoon-shaped), with very sloping collars towards the dorsal surface. Miliary spines on both the oral and aboral surface are only slightly curved dorso-ventrally (Fig. 9 Li & Lii), the tips only very slightly widened and flattened. Those within the peripetalous fasciole have a relatively broad shaft that increases in width distally forming a globular head (Fig. 9 Ii). The shaft typically has six to eight serrated ridges which increase in height distally. Clavula structure is similar in the peripetalous and subanal fascioles with a long narrow shaft and a distinct ‘crown’ distally (Fig. 9 O). The degree of development of the crown varies slightly with some clavulae having a more simplified structure (Fig. 9 Iii). Penicillate tube feet are present around the peristome (see Fig. 2 B & G where they are visible as dark patches around the mouth). The suction discs of the anterior ambulacral aboral tube feet are formed of four to six lobes, while within the peripetalous and subanal fascioles (Fig. 9 Pi & Pii) they are formed of six to ten lobes. The sphaeridia are slightly elongate-round, occurring adorally not in grooves or pits. These have no ornamentation and thus offer no species-specific characters. This species has a particularly diverse array of highly localised pedicellariae (Fig. 9 A–H) all of which have three valves. Three distinct forms of globiferous pedicellaria are present. These can be grouped into two types, fistulate (tubular) and open-bladed, which only occur on the oral surface. Fanged open-bladed globiferous pedicellariae (Fig. 9 Di–Diii) are forceps-like having straight, narrow, predominantly open valves (neck of blade is closed, gradually opening distally), with each valve terminating in two long, inward-facing fangs. The valves only meet where the fangs interlock, with one valve typically being shorter than the other two. The proximal region is small compared to the length of the valve; while the neck is short and attached to a very short simple, unornamented, stalk (Fig. 9 Diii). The valves are covered in a black skin which is thick and glandular in appearance around the fangs. This is the likely source of venom as there are no internal glands or stalk glands present. This form occurs in large numbers on the posterior of the oral ambulacra (Fig. 2 G; visible as dark ‘dots’ on the posterior of the ambulacra). Fanged fistulate globiferous pedicellariae (Fig. 9 Ei–Eiv) have large curved valves that are closed and tubular with each valve having an inverted triangular opening distally (Fig. 9 Ei), terminating in two long inward-facing fangs (Fig. 9 Eii). The proximal regions of the valves are proportionally large, with large adductor muscle insertion points. The neck of the valve is muscular, with muscle strands entering through the outside of the proximal region through transverse slits into the adductor muscle insertion points. The stalk is very short with vertical projecting rods around its base (Fig. 9 Eiv). A large glandular mound occurs on the distal region of each valve (Fig. 9 Eiv), encasing all but the tips of the fangs and is the likely source of venom. This makes them easily visible on the test, where they occur in large numbers around the anterior of the oral ambulacra and around the peristome (Fig. 2 B). Simple fistulate pedicellariae (Fig. 9 Ci–Civ) occur in very small numbers on the anterior oral ambulacra. This form has highly curved tubular valves that terminate in a circular foramen with an arc (no tooth on the apex of the lower lip) of eight to ten small teeth. The proximal region of each valve is proportionally large, with large adductor muscle insertion points. The stalk is simple and very short, and is attached to the valves by a short muscular neck. The valves are covered in a thick black skin (Fig. 9 Cii), which is evenly distributed. Tridentate pedicellariae with straight valves occur as three forms: narrow-valved (Fig. 9 Giii & Giv), spatulate (Fig. 9 Gi & Gii) and terminal-toothed (Fig. 9 Fi–Fiii). The narrow-valved form occurs only on the anterior of the oral ambulacra. This form has narrow open blades that taper distally, with small peripheral teeth along the edges of the blades, which interlock along their entire length. The ossicles of the blades consist of an open lattice distally, becoming tighter proximally with a Y-shaped apophysis at the neck of the blade (Fig. 9 Giii). The proximal region is proportionally small, with small adductor muscle insertion points. The neck of the pedicellaria is short and muscular, attached to a short stalk (less than the length of the valves). The spatulate form (Fig. 9 Gi) is similar to the narrow-valved form but with an expanded blade distally and is found posterior of the peristome on the oral ambulacra. This form typically has larger proximal regions than the narrowvalved form; however, intermediates between the two are present. The third tridentate form has narrow valves with each valve terminating in a proportionally large distal tooth (Fig. 9 Fi–Fiii). This form only occurs on the aboral surface within the peripetalous fasciole. Small peripheral teeth are present down half to three quarters of the length of the edges of the blades, which interlock distally. The proximal regions are proportionally large and triangular. The neck is thin (Fig. 9 Fiii) and attached to a narrow short stalk (shorter than the length of the valves). A single form of rostrate pedicellaria is present (Fig. 9 Hi–Hiii), typical of the spatangoid type. This only occurs in very small numbers on the oral ambulacra. The valves are highly curved and narrow (Fig. 9 Hii), the sides of the valves being non-denticulate with overturned lips giving a partially enclosed appearance (Fig. 9 Hi). The distal region of the blade has a slightly overturned fan of small teeth (Fig. 9 Hi). These overlap when the three valves are together, being the only point of contact for the blades. The neck is short and narrow, connected to a long stalk. Triphyllous pedicellariae (Fig. 9 Ai & Aii) occur orally and aborally with a short muscular neck on a midlength stalk (twice the length of the valves). The blades are rounded, tapering distally. Teeth are present along the edges of the valves, which interlock with those of the other blades along their entire length. Ophicephalous pedicellariae with large proximal handles occur as a single form (Fig. 9 Bi–Biv), present in large numbers on the oral ambulacra and in the subanal fasciole. The blades are highly constricted (Fig. 9 Bi, Bii & Biv) resulting in an inverted triangular blade. The edges of the blades are denticulate with an overhanging lip distally. The teeth of the blades interlock both laterally and distally forming the jaw-set. This sits directly on the stalk, which measures four to five times the length of the valves.Published as part of Coppard, Simon E., 2008, A comparative analysis of the spatangoid echinoid genera Brissopsis and Metalia: a new genus and species of spatangoid (Echinodermata: Echinoidea: Brissop- sidae) from the Philippines and the reassignment of Brissopsis persica to Metalia, pp. 1-23 in Zootaxa 1760 on pages 3-6, DOI: 10.5281/zenodo.29398

A new genus of mellitid sand dollar (Echinoidea: Mellitidae) from the eastern Pacific coast of the Americas

Coppard, Simon E. (2016): A new genus of mellitid sand dollar (Echinoidea: Mellitidae) from the eastern Pacific coast of the Americas. Zootaxa 4111 (2): 158-166, DOI: 10.11646/zootaxa.4111.2.

Brissopsis Agassiz 1840

Brissopsis Agassiz, 1840 Catalogus systematicus Ectyporum Echinodermatum fossilium Musei Neocomiensis, O. Petitpierre, Neuchâtel. Jent and Gassmann, Solothurn, p. 3 and 16. Type species: Brissopsis lyrifera Forbes, 1841, by original designation. Assigned species: Mortensen (1951 a) lists seventeen Recent species and two varieties (Brissopsis alta Mortensen, 1907; B. atlantica Mortensen, 1907; B. atlantica var. mediterranea Mortensen, 1913; B. bengalensis Koehler, 1914; B. columbaris Agassiz, 1898; B. elongata Mortensen, 1907; B. evanescens Mortensen, 1950; B. jarlii Mortensen, 1951 b; B. luzonica (Gray, 1851); B. lyrifera (Forbes, 1841); B. micropetala Mortensen, 1948; B. obliqua Mortensen, 1948; B. oldhami Alcock, 1893; B. pacifica (Agassiz, 1898); B. parallela Koehler, 1914; B. persica Mortensen, 1940; B. persica var. elevata Mortensen, 1940; B. similis Mortensen, 1948; B. zealandiae Mortensen, 1921) and a further 75 fossil species in this genus. Distribution: Eocene to Recent, Indo-Pacific, Atlantic, Mediterranean.Published as part of Coppard, Simon E., 2008, A comparative analysis of the spatangoid echinoid genera Brissopsis and Metalia: a new genus and species of spatangoid (Echinodermata: Echinoidea: Brissop- sidae) from the Philippines and the reassignment of Brissopsis persica to Metalia, pp. 1-23 in Zootaxa 1760 on pages 2-3, DOI: 10.5281/zenodo.29398

Lissocidaris xanthe Coppard & Noordenburg, 2007, sp. nov.

<i>Lissocidaris xanthe</i> sp. nov. (gender: feminine) <p>Figs 1, 2, 4, 5 and 6.</p> <p> <b>Diagnosis:</b> This new species has aboral primary interambulacral spines that are typically cream in colour (occasionally with faint, pink banding) and slightly curved, measuring more than twice the test’s horizontal diameter (h.d.) at the ambitus. The spines have a relatively smooth surface formed by dense anastomosing cortical hairs that coalesce to form a continuous crust between low, longitudinal ridges. These ridges typically number twenty-two and are more apparent when the spine is viewed in transverse section. The collar of the spine is short and dark brown, while the neck is typically white and unconstricted. Oral primary interambulacral spines have serrated edges, while scrobicular tubercles are slightly adpressed forming a loose mail around primary tubercles. The large globiferous pedicellariae have moderately narrow valves, each valve with a narrow longitudinally oval opening beneath a large terminal tooth. Small globiferous pedicellariae have proximal regions with rounded edges and a longitudinally oval opening beneath the terminal tooth. Crenulations are well defined on the adapical side of tubercles above the ambitus. The colour of the primary ambulacral spines, secondary spines, miliary spines and the test epithelium is yellow-orange/light-brown.</p> <p> <i>Holotype</i>: NHM 2007.4 Natural History Museum, London.</p> <p> <i>Paratype</i>: NHM 2007.5 Natural History Museum, London.</p> <p> <b>Etymology:</b> Named after the light, fair colour (yellow-orange/light-brown) of the test epithelium, primary ambulacral spines, secondary spines and miliary spines.</p> <p> <b>Description:</b> The test is relatively thick, subpentagonal in outline and inflated, with flattened adapical and oral surfaces (Figs 2 C & D). The denuded test (Figs (2A–E) is whitish-cream (the epithelium is yelloworange/light-brown in life) with a h.d. of 47 mm and a vertical diameter (v.d.) of 34 mm (paratype). The apical disc is monocyclic, with all ocular plates narrowly insert (Fig. 2 A), measuring 40 % of the test’s h.d. (19 mm in the paratype). The genital plates are all of equal size, widening towards the periproct. The genital and ocular plates are whitish-cream, with a brown, raised, central region. This fades into pink beneath the gonopores. The madreporite is dark brown, convex, covering most of genital plate 2. All apical plates are densely covered with granules, which increase in size towards their outer edge. The genital pores are proportionally small, slightly elevated and occur near to the outer third of the genital plates. The periproctal plates are brownishpink, the five at the corners of the pentamerous periproct, elegantly pointing between the genital plates, being contiguous with the ocular plates.</p> <p>The ambulacra are slightly sinuate (Fig. 2 D), 25 % the width of the interambulacra (5.5 mm in the paratype) at the ambitus. The interporiferous zones are approximately twice the width of a pore-zone, with each adradial plate to the marginal tubercle having a large outer secondary tubercle, with two smaller secondary tubercles placed one above the other. These form regular vertical series, with each plate also having miliary granules. No naked or sunken median line is present. The pore-zones are slightly sunken, pores nonconjugate, approximately equal in size, with each pore separated by a broad wall (Fig. 2 H).</p> <p>The interambulacra (Fig. 2 C) are formed of two series of eight or nine primary tubercle bearing plates. Tubercles have areoles of moderate size and depth (Fig. 2 I), which are separated by at least two rows of scrobicular tubercles aborally, the last two sometimes being confluent adorally. Areoles on the oral surface are distinctly transverse-oval, while adapical areoles are circular. Primary tubercles are of moderate size, perforate, with distinct traces of crenulation on the adapical sides of the tubercles above the ambitus. Scrobicular tubercles are of moderate size. Outside the scrobicular ring the plates are densely covered with secondary tubercles which decrease gradually in size, becoming granular towards the slightly sunken median line. Horizontal lines are distinct, particularly adapically, but neither the median line nor the horizontal lines are naked.</p> <p>The peristome measures 38 % of the test’s h.d. (Fig. 2 B) (18 mm in the paratype) with twelve ambulacral imbricate plates in each column and pores-pairs uniserially arranged. Six to seven non-ambulacral plates form a regular interradial series reaching the mouth.</p> <p> <b>FIGURE 6.</b> Pedicellariae from A, holotype of <i>L. xanthe</i> <b>sp. nov.</b>, B, holotype of <i>L. fusca</i>, C, holotype of <i>Calocidaris micans</i> and D, <i>Compsocidaris pyrsacantha</i> (S.E. Coppard private collection); i & ii, large form of globiferous pedicellaria; iii (oral) & iv (aboral), small form of globiferous pedicellaria; v, tridentate pedicellariae; vi (side view) & vii (internal view) individual valves of large globiferous pedicellariae; viii (side view) & ix (internal view) individual valves of the small form of globiferous pedicellaria; x (side view) & ix (internal view) individual valves of tridentate pedicellariae; Axi, individual valves of triphylous pedicellariae from <i>L. xanthe</i> <b>sp. nov.</b>; Cxii individual valve of the large form of tridentate pedicellaria of <i>Calocidaris micans</i>.</p> <p>The Aristotle’s lantern (Figs 2 F & G) and perignathic girdle are typical of the cidaroid form, with small epiphyses that do not project and relatively high apophyses (Fig. 2 E).</p> <p>Aboral primary interambulacral spines (Figs 4 Avii & Aviii) are long, slender and tapering distally, typically cream in colour (occasionally with faint, pink banding), and slightly curved (see Fig. 4 Avii). The longest measures 85 mm on the holotype (Figs 1 A–C) (x 2.4 the test’s h.d.) with a diameter of 3 mm. These are cylindrical in transverse section adapically (Figs 5 Ai & Aii), becoming slightly flattened beneath the ambitus. The spines are relatively smooth and typically have a glossy sheen (Fig. 4 Ai) (particularly on smaller specimens, the glossy sheen being absent on the aboral primary spines of some larger specimens) with approximately twenty-two longitudinal ridges faintly visible on primary spines at the ambitus (Fig. 4 Avii). This is due to the presence of dense, anastomosing cortical hairs that coalesce to form a continuous crust between longitudinal ridges (Fig. 5 Aii). These ridges are more noticeable beneath the ambitus due to the reduced depth of the cortical hairs. In smaller specimens the ridges appear to be completely absent but are visible when the spine is viewed in transverse section. The collar of the spine (Fig. 4 Aiii) is short (typically 2 mm), dark brown and increases in thickness towards the milled ring. The neck of the spine (Fig. 4 Aiii) is long, straight and completely smooth with no cortical hairs, or ridges in the cortex (Fig. 5 Ai). The ridges begin above the neck of the spine and continue to the spine’s tip. When viewed in transverse section (Fig. 5 Ai & 5Aii) the medulla is seen to be formed of reticulated stereom which is slightly darker in colour than the surrounding stereom. The tips of the primary spines gently taper and flatten dorso-ventrally to a blunt tip (Fig. 4 Aii), or in small specimens to a small crown; in both cases the longitudinal ridges are more apparent due to the absence of cortical hairs.</p> <p>Oral primary interambulacral spines are white (Figs 1 B & 4Axii), distinctly flattened, slightly curved and moderately broad, with serrated edges, these being more apparent in smaller specimens. These have slightly rounded blunt tips, with those that are near the mouth tapering distally. They have an orange-brown collar, a smooth white neck, and longitudinal ridges on both the upper and lower surfaces. The third and fourth oral primaries (Fig. 1 B) are transitional to the aboral primaries.</p> <p>Primary ambulacral spines are undifferentiated orally and aborally (Fig. 4 Av). These typically measure 3– 4 mm in length, are slender, slightly flattened and curved, with longitudinal ridges. These spines taper distally and have slightly rounded tips.</p> <p>Scrobicular spines are slightly adpressed and ridged (Figs 4 Aviii & Aix), widening above the base up one quarter of their length, where they taper to a blunt tip. They typically measure 5 mm in length and do not form a very close mail around the base of the primaries (Fig. 1 A). Secondary spines on the ambulacra and interambulacra are slender (Fig. 4 Av), 2–3 mm in length, slightly flattened and curved, with longitudinal ridges. These spines also taper distally and have slightly rounded tips. Miliary spines (Fig. 4 Aiv) are shorter (1.5 mm in length) but proportionally broader than the secondary spines, and are distinctly flattened with longitudinal ridges. They are broadest at their base, tapering to a rounded tip. Oral peristomal spines densely cover the peristomal plates. These are highly curved and distinctly club-shaped, with serrated longitudinal ridges (Fig. 4 Axi).</p> <p>Large globiferous pedicellariae are abundant aborally, particularly down the interambulacra, with all pedicellariae examined having three valves. They have short necks and stalks (Figs 6Ai & Aii), with large valves that gradually narrow above the adductor muscle insertion points towards a large and sharply pointed terminal tooth (Figs 6Avi & Avii). The opening beneath the terminal tooth (Fig. 6Avii) is longitudinally oval in shape distally, becoming U-shaped above the lip. The oval region is surrounded by narrow, pointed, peripheral accessory teeth. These decrease in size around the U-shaped region. The distal region beneath the opening is narrow, increasing in width towards the adductor muscle insertion regions. These muscle insertion regions are longer than wide, and measure just over half the length of the valves.</p> <p>The small form of globiferous pedicellaria is very abundant both orally (Fig. 6Aiii) and aborally (Fig. 6Aiv), particularly down the ambulacra. Pedicellariae of this form have relatively long stalks, particularly on the oral surface, with short necks. Each valve has a proportionally large terminal tooth (Figs 6Aviii & Aix) typical of the Cidarina. Peripheral teeth are present along the edges of the valves, being larger around the longitudinally oval opening, which measure 22 % of the length of the valve. The lip beneath the opening is denticulate. The distal region of the valve beneath the lip and the proximal adductor insertion regions are broad, while the edges of the valves are curved.</p> <p>Tridentate pedicellariae occur only as one form (Figs 6Ax & Axi), typical of the genus. They have relatively long stalks with a short neck. The valves are slender, have a terminal tooth, and increase in width proximally, flaring above the proximal region. These flaring regions are visible as crests when viewed from the side (Fig. 6Ax). The valves slightly constrict beneath these crests, directly above the adductor muscle insertion regions. The proximal regions are slightly longer than they are wide with a relatively small keel.</p> <p>A single pedicellaria representing a third class was found on the paratype, but not on the holotype or any of the other nine specimens examined. This has a long stalk with a short neck and very short conical shaped valves (Fig. 6Axii). The adductor muscle insertion regions are fairly open, indicating a very limited grasping pressure. This single pedicellaria is clearly not representative of this species and appears to be an anomaly.</p> <p>Spicules in the tube feet occur as smooth rods. These are typically straight but occasionally bifurcate at one end.</p>Published as part of <i>Coppard, Simon E. & Noordenburg, Henk Van, 2007, A new species of Lissocidaris (Echinodermata: Echinoidea: Cidaridae) from the Philippines: convergent evolution among smooth-spined cidaroids, pp. 53-65 in Zootaxa 1493</i> on pages 55-63, DOI: <a href="http://zenodo.org/record/176981">10.5281/zenodo.176981</a&gt

Lissocidaris Mortensen 1939

Lissocidaris Mortensen, 1939 The John Murray Expedition (to the Indian Ocean), 1933 – 34: scientific reports. Report on the Echinoidea of the Murray Expedition vol. 6 (1), p. 11–14, pls. 3, 5 & 6. British Museum (Natural History). Type species: Lissocidaris fusca Mortensen, 1939 by monotypy. Assigned species: only the type speciesPublished as part of Coppard, Simon E. & Noordenburg, Henk Van, 2007, A new species of Lissocidaris (Echinodermata: Echinoidea: Cidaridae) from the Philippines: convergent evolution among smooth-spined cidaroids, pp. 53-65 in Zootaxa 1493 on page 54, DOI: 10.5281/zenodo.17698

Coelopleurus L. Agassiz 1840

Coelopleurus L. Agassiz, 1840 Catalogus systematicus Ectyporum Echinodermatum fossilium Musei Neocomensis. p. 12 and p. 19. Type species: Cidaris coronalis Leske, 1778 (= Coelopleurus equis L. Agassiz, 1840) by monotypy. Assigned species: see Mortensen (1935) for the additional 11 species/subspeciesPublished as part of Coppard, Simon E. & Schultz, Heinke A. G., 2006, A new species of Coelopleurus (Echinodermata: Echinoidea: Arbaciidae) from New Caledonia, pp. 1-19 in Zootaxa 1281 on page 4, DOI: 10.5281/zenodo.17339

Coelopleurus exquisitus Coppard & Schultz, 2006, sp. nov.

Coelopleurus exquisitus sp. nov. Figs. 1–5, table 1 a. Diagnosis: Long, highly curved primary spines that are banded red and pale­green for three quarters of their length distally (see Figs 1 A–C and 3 D–F). For the basal quarter of their length the spines blend from pale­green to lavender mid­way through the spine’s collar. The collar measures 18 % of the ambital primary spine’s length and has a prominent longitudinal dorsal ridge. Smaller longitudinal ridges are present on both the collar’s dorsal (Fig. 3 G) and ventral (Fig. 3 H) surface with granules between dorsal ridges. Secondary spines (Fig. 3 K) are either cream or olive green blending into red distally and tapering to a point. The test has large, straight edged, naked median interambulacral regions (Figs. 1 A and C, 2 A and F; 3 C) that start from the genital plates, and comprise 65 % of the width of the interambulacra measured at the ambitus. These naked median regions are purple in contrast to the test’s olive/light brown epithelium, with each median region having a pinkish/lavender undulating line that continues to the lower element of the fifth plate from above. The peristome is large (Figs. 1 B and 2 B), measuring 56 % of the test’s horizontal diameter. The auricles are robust, with moderately high processes (Fig. 2 E). The aboral ophicephalous pedicellariae have stalks that are not distinctly swollen or ‘fleshy’ (Fig. 3 L). The distal and proximal regions of the ophicephalous valves are equal in length and are slightly constricted above the adductor muscle insertion points (Fig. 4 H and I). Holotype: EcEh 1281, MNHN­Paris­Echinoderms, N. O. “Vauban” MUSORSTOM 4, St. DW 181 350 m, 18 ° 57 ’S 163 ° 22 ’E, New Caledonia, C. Vadon Coll. 18 th September, 1985. Paratypes: fifteen specimens; fourteen specimens in MNHN­Paris­Echinoderms (EcEh 1282), N. O. “Vauban” MUSORSTOM 4, St. DW 181 350 m, 18 ° 57 ’S 163 ° 22 ’E, New Caledonia, C. Vadon Coll. 18 th September, 1985. One specimen in the Natural History Museum, London, registration number (NHM 2006.599). Other material: Five specimens (unregistered) MNHN­Paris­Echinoderms, N. O. “Jean­Charcot” BIOCAL, St. DW 50 240–260 m, 23 ° 07’ South, 167 ° 54 ’ East, New Caledonia, Guille and Menau Collection, 31 th August 1985; Registration No. J 20052 Sydney Museum collection,, Coreolus Expedition, 23 ° 06’ South, 167 ° 05’ East, South of Isles of Pines, New Caledonia, at a depth of 520 m. Etymology: After the exquisite coloured markings on the test and spines. Description: The test is subcircular (Figs. 1 A, 2 A and E) (not distinctly pentagonal as in C. maillardi see tables 1 a and 1 b) with a horizontal diameter of 32.5 mm (holotype) and is moderately inflated adapically with a vertical diameter of 17.8 mm. The ambulacra are slightly raised aborally (Fig. 2 C) while the interambulacra are correspondingly slightly sunken (Fig. 2 D). The base colour of the epithelium of the test is olive/light brown (Fig. 1 A), these regions being white on the denuded test (see Fig. 2 A–E). The apical system is small (9.0 mm) and dicyclic (see Figs. 2 A and 3 A) with all ocular plates exsert. The genital plates are proportionally large, with ophicephalous pedicellariae on their inner edge (Fig. 3 A). The attachment points for the ophicephalous pedicellariae are small granules on the denuded test (Fig. 2 A). The genital plates and ocular plates are brightly coloured. The points of the ocular and genital plates are pink with a lavender central region in contrast to the orange terminal plates of the outer series of the interambulacra and the olive/light brown (white on naked test) of the ambulacra. The genital pores are small and are encompassed on the genital plates by a purple U­shaped region that forms the top of the purple naked region of each interambulacrum. The periproct on the holotype measures 4.8 mm (approximately 15 % of the test’s horizontal diameter) and has four equally sized, valve­like, triangular plates. These are cream in colour, and fill the periproct (Fig. 3 A). The ambulacra are slightly inflated, the adradial edges undulating (Fig. 3 B) as the ambulacra expand towards the ambitus and decreasing in size towards the peristome. Plating on the ambulacra is formed of typical arbaciid triads (compound plates formed of three plates), each triad possessing a large primary tubercle with an imperforate mamelon, the areole forming a raised platform on the plate, with two series of primary tubercles in each ambulacrum. The uppermost ambulacral plates bear a primary tubercle only on one side (i.e. in one series). On the plate immediately beneath the ambitus the primary tubercles on the ambulacra and interambulacra are of equal size (Fig. 2 C and D). Adapically the areoles are confluent, but from the compound plate directly above the ambitus to the peristome there is a narrow median region which has a series of secondary tubercles, interspaced with miliary tubercles (Fig. 3 B). These increase in number adorally. There are no secondary or miliary tubercles in the pore­zones above the ambitus, while miliary tubercles are present in the pore­zones beneath the ambitus. The ambulacra are olive/light brown (Fig. 1 A), while on the naked test the ambulacra are white with orange median lines (Figs. 2 C and 3 B), which begin from the fourth primary tubercle and bisect the median region continuing to the apex of the test adorally. The interambulacra are broader than the ambulacra at the ambitus (Figs. 3 B and C) with a ratio of 1.5:1.0. The interambulacra are characterized by large naked median regions (Figs. 2 D and 3 C). These are purple and sharply defined both in colour (bordered by olive lines with red dots on the adradial edge of the lower corner of each plate) and by small straight edged adradial ridges. This naked region comprises 65 % of the width of the interambulacra measured at the ambitus. A pinkish/lavender undulating line proceeds down the centre of each naked median region, which increases in the width of the undulation towards the ambitus. The purple colouration starts from the genital plates and is clearly defined to the sixth plate from above (Fig. 3 C). The purple colour faintly continues beyond the naked region towards the peristome, but not in the midline between the two series of tubercles beneath the ambitus (Fig. 3 C). There are no primary tubercles in the naked median regions on the first four plates adapically. One or two secondary tubercles are present on the adambital and lower adradial edges of the fourth plates. On the fifth plate there are several secondary tubercles and a small primary tubercle, which lacks a large areole. The sixth plate (at or just below the ambitus depending on which series is being observed) has a large primary tubercle, equal in size and structure to those in the same region on the ambulacra. There are 6–7 primary tubercles in each of two main series of primary tubercles, which gradually decrease in size towards the peristome. The median region between these two series is relatively narrow with a single series of secondary tubercles and miliary tubercles on either side. Three distinctive red ‘dots’ are present in the suture line between the central elements of plates 5 and 6 in one series and plate 5 in the second series (Fig. 3 C). These correspond to the positions of three red secondary spines. An outer series of tubercles extends from the peristome to the apical disc, which become increasingly oblique and reduced in number adapically. Such tubercles on the naked test appear white in the upper element of each compound plate and red in the lower element appearing as red patches in olive lines that border the naked median regions. These red tubercles increase in number with a single tubercle in the lower corner of plate 1, 2 in the lower corner of plate 2, with a maximum of three in an oblique series in the lower corner of plate 3. The peristome is subcircular (Fig. 2 B and E) and on the holotype measures 17.4 mm in diameter, 56 % of the test horizontal diameter. The peristomial membrane is dark brown with five pairs of buccal tube feet (Fig. 1 B). Each pair being distinctly separated from the next and closely surrounded by large numbers of ophicephalous pedicellariae. A few scattered ophicephalous pedicellariae occur across the peristomial membrane, which is otherwise naked. The buccal notches are shallow but distinct while the gill­tags proceed to the midpoint of the fourth ambulacral primary tubercle (Fig. 2 B). The auricles have moderately high processes and are robust in appearance (Fig. 2 E). Primary spines occur in three forms (see Figs. 3 D­J). The most adapical large primary spines (the first large tubercles on the ambulacra) are moderately curved and almost cylindrical except for the laterally compressed ridge, which proceeds up one third of the spine’s dorsal length (Fig. 3 F). These spines are banded red and pale­green for three quarters of their length distally. For the basal quarter of their length the spines blend from pale­green to lavender. This colouration occurs on all sides of the spine. The collar of the spine does not distinctly end, as ridges and furrows occur along the spine’s total length and are clearly visible when seen in cross­section (Fig. 4 B). Granules occur between the ridges, most noticeably proximally, but also along the spine’s length both dorsally and ventrally. The tips of these spines are slightly rounded, but without an obvious hyalinecap. The spines are subcircular when viewed in cross­section with distinct ridges and furrows, which proceed longitudinally down the spine’s length. The internal structure (Fig. 4 B) consists of a small central cavity (which comprises 27 % of the spine’s diameter), a large region of dense stereom and an outer dermis. The central cavity is comprised of an axial cavity, which is filled with loosely reticulated stereom and 12–14 radiating solid wedges. These project into and through the dense stereom. The dermis is thin and rough in texture, but non­verticillate. The second form of primary spine start on the fifth ambulacral tubercle and the second interambulacral tubercle beneath the naked median region and continue down to the apex of the test. These primary aboral spines are highly curved (Fig. 3 D and E), and flattened both laterally and ventrally above the collar. On their upper surface these spines have the same colouration as the adapical primaries, however, on their underside the spines are typically white (Fig. 1 B), with occasionally a faint impression of the pattern observed on the dorsal surface. The longest primary spines in this species are of this form and on the holotype the longest spine measures 76.2 mm (2.3 times the test’s horizontal diameter) but does not have the tip present. The collar is well defined and typically measures 18 % of the spine’s total length with distinct longitudinal furrows and ridges on the underside (Fig. 3 H), and granules and a few ridges on the dorsal surface (Fig. 3 G). In cross­section the spines are triangular with a convex ventral surface. Their internal structure consists of a small central cavity, which comprises 23 % of the spine’s horizontal diameter, a large, dense region of stereom and a relatively thick epidermis. The central zone is composed of an axial cavity, which is filled with loosely reticulated stereom and 12–14 radiating solid wedges. These project into the dense stereom region. The epidermis is smooth continuing from the spine’s tip to the spine’s collar. The third form of primary spine is present on the oral surface. They are short (typically not longer than 15 mm) and dorso­ventrally flattened (Figs. 3 I and J; 4 C). On their dorsal surface they are light green with a central ridge (Figs. 3 I and 4 C), while their ventral surface (Fig. 3 J) is white and smooth. The collar is well defined, measuring 18 % of the spine’s length, with distinct ridges dorsally and ventrally. These spines are almost diamond­shaped when viewed in cross­section (Fig. 4 C). Their internal structure is similar to the ambital spines but compressed dorso­ventrally. However, the axial cavity is proportionally larger (33 % of the spine’s diameter), filled with loosely reticulated stereom and has short radiating solid wedges which typically number from 12–14. The dense stereom is well developed and continues between the dorsal and ventral solid wedges. The epidermis is thick and smooth and continues down the spine’s length to the collar. Secondary spines (Fig. 3 K and 4 D) are slender and pointed (not club­shaped) either red proximally blending into green distally or cream with longitudinal ridges. These spines are circular in cross­section with a large central cavity (relative to the primary spines) measuring 55 % of the spine’s diameter, which consists of a small axial cavity filled with loosely reticulated stereom and 12–14 solid wedges. The region of dense stereom is proportionally smaller than in the primary spines, while the epidermis is rough (but nonverticillate) and thin. Miliary spines are similar in colour to secondary spines but lack the longitudinal ridges. Ophicephalous pedicellariae (Figs. 3 L and 4 E, H and I) are abundant all over the test surface. These are particularly noticeable around the periproct (Fig. 3 A) and along the adradial edges of the ambulacra. The stalks of the ophicephalous pedicellariae are relatively long (typically 1.5 mm), the skin of the stalk being only very slightly swollen proximally (Figs. 3 L and 4 E). The neck of each pedicellaria is short but is not distinctly swollen, indicating the absence of glandular tissue in this region. The distal and proximal regions are of approximately equal length, with peripheral teeth present both along the edges of the valves and along the edges of the apophyses (Figs. 4 H and I). These interlock when the valves are closed. The distal regions are constricted, the degree of constriction varying from being very constricted (Fig. 4 H) to moderately constricted (Fig. 4 I) on a single sea urchin. Both specimens illustrated (Figs. 4 H and I) were removed from the aboral surface of the holotype. Ophicephalous pedicellariae are also abundant on the oral surface. These are particularly noticeable around the peristome, the valves of which are only slightly constricted. All ophicephalous pedicellariae in this species exhibit very developed proximal handles, which increase grasping pressure (Mortensen, 1935) and are typical of this class of pedicellaria. The tridentate pedicellariae have long narrow valves (Figs. 3 M, 4 F and J) which meet for their entire length. Peripheral teeth are present along the edges of the valves, which interlock. The neck of each pedicellaria is narrow and relatively short, and is attached to a relatively long (approximately 1.5 mm) and narrow stalk. Tridentate pedicellariae are abundant on the aboral surface particularly around the adapical region of the ambulacra, and are typical of the genus. Triphyllous pedicellariae are less numerous than the other two classes in this species, and have a small head supported by a relative long, broad neck on a proportionally (in relation to the size of the valves) long stalk (Figs. 3 N and 4 G). The valves are broad and spoon­shaped with small peripheral teeth along the edges of the valves which interlock when the head of the pedicellaria is closed. This form of pedicellaria is distributed all over the test in small numbers but provide no species­specific characters.Published as part of Coppard, Simon E. & Schultz, Heinke A. G., 2006, A new species of Coelopleurus (Echinodermata: Echinoidea: Arbaciidae) from New Caledonia, pp. 1-19 in Zootaxa 1281 on pages 4-12, DOI: 10.5281/zenodo.17339