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Fig. 24 in New insights into the origin and relationships of blastoid echinoderms
Fig. 24. Oral surface of the coronate Stephanocrinus angulatus Conrad, 1842, early Silurian (Wenlock), New York, USA, showing similarity to blastoid oral surface. Five erect ambulacra (A–E) arise from the trunk mounting plates, which are homologues of the lancets of blastoids. Food grooves are covered by paired ambulacral cover plates (cp). The mouth frame is composed of five homologues of blastoid deltoids and dichoporite lateral plates (L1–L5), and covered by primary oral cover plates (1–5). G, gonopore; PD, posterior deltoid. For plate homologies see Table 2. Redrawn and relabelled from Brett et al. 1983: 638, fig. 4A.
Fig. 21 in New insights into the origin and relationships of blastoid echinoderms
Fig. 21. Oral area of the hemicosmitoid Hemicosmites pyriformis Buch, 1846, Middle Ordovician, Russia. Three ambulacra, interpreted as A, shared BC, and shared DE, converge on the mouth (M) and are surrounded by a circlet of nine radial plates (R1–R9) outside which is another circlet of nine lateral plates (L1–L9). Three wedge plates occur immediately clockwise of radial plates R3, R6, and R9. The hydropore (not shown) occurs in the adoral part of radial 9. In life, biserial cover plates would have roofed over the food grooves. Redrawn and relabelled from Bockelie 1979: 375, fig. 8b.
Fig. 18 in New insights into the origin and relationships of blastoid echinoderms
Fig. 18. Ambulacra and brachioles in the callocystitid Pseudocrinites pyriformis Paul, 1967a, Ludlow (Silurian), England, UK. The mouth and food grooves are hidden by cover plates (shaded). Both ambulacra start with two brachiole facets on the left. These are interpreted as BC1 and C1 in the C ambulacrum and DE1 and E1 in the E ambulacrum (left). F, facet; H, hydropore; M, mouth. Redrawn from Paul 1967a: 329, fig. 21.
Fig. 17 in New insights into the origin and relationships of blastoid echinoderms
Fig. 17. Oral and ambulacral plating of the callocystitid Lipsanocystis traversensis Ehlers and Leighley, 1922 (UMMP 33311), Middle Devonian, Michigan, USA, illustrating the alternative "B–E the same" pattern of primary brachioles in which all four ambulacra have the first brachiole facet to the left. A1, oral view of showing the four ambulacra and double hydropore; A2, the same with interpretation of oral plates superimposed; A3, interpretation of oral and ambulacral plating. Ambulacrum A stopped developing after the first plate formed. In ambulacra B–E the first brachiole is to the left and the second to the right. B–E, ambulacra; A1a–E1a, first plates in corresponding ambulacra; b second plates in ambulacra B–E; G, gonopore; H, hydropore; M, mouth; P, periproct; 6, 7, orals 6 and 7 under the conventional interpretation. Dashed line outlines of ambulacra and brachiole facets. Only the first two ambulacral plates are shaded.
Fig. 16 in New insights into the origin and relationships of blastoid echinoderms
Fig. 16. Ontogenetic development of ambulacra in the callocystitid Lepadocystis moorei (Meek, 1871), Late Ordovician, Ohio, USA, showing addition of oral and ambulacral plates and development of the "BD different" pattern of primary brachioles. A. Triradiate stage with the first pair of ambulacral plates (A1a, A1b, and so on) interpreted as A, shared BC and shared DE. Arrows indicate the positions where the radial water vessels of the shared ambulacra branch to produce a pentaradiate rhombiferan, assuming subterminal branching. B. Pentaradiate stage with a single brachiole per ambulacrum. The two new brachioles are interpreted as the first brachioles in separate ambulacra C and E. Arrows indicate positions where the radial water vessels of all five ambulacra branch to produce second brachioles. C. Pentaradiate stage with two brachioles per ambulacrum. The second brachioles in the shared ambulacra are interpreted as B1 and D1. Note that the subterminal branching has produced the "BD different" pattern of primary brachioles in which the first two brachioles in ambulacra B and D branch to the left as viewed in the growth direction, whereas in ambulacra A, C and E only the first brachiole is on the left. In addition, the second brachioles exhibit a pattern which is A right, B left, C right, D left, and E right, which equals Lovén's Law as restated by Paul and Hotchkiss (2020). M, mouth; O6 and O7, orals 6 and 7 in the conventional interpretation of the oral circlet. Solid lines, plate sutures; dashed lines, outline of ambulacra and brachiole facets. Redrawn from Paul and Hotchkiss 2020: 1096, fig. 8.
Fig. 20 in New insights into the origin and relationships of blastoid echinoderms
Fig. 20. Oral plating of two blastozoans showing similarities in arrangement of facet-bearing plates. A. Rhombifera bohemica Barrande, 1867, Late Ordovician, Czech Republic. B. Lysocystites sculptus Miller, 1889, early Silurian, Indiana, USA. In both the mouth (M) is encircled by only five plates (O1–O5 in A), which alternate with five radial plates (R1–R5) that bear ambulacral facets (F) and are connected to the mouth by short food grooves. In Lysocystites (B) the facet-bearing plates are the homologues of eublastoid lancet plates. A–E, ambulacra A–E; An, anus; d, additional anal deltoids; L1–L5, lateral plates. For plate homologies see Table 2. A after Kesling 1962: 283, fig. 2; B after Sprinkle 1973: 141, fig. 35.
Fig. 15 in New insights into the origin and relationships of blastoid echinoderms
Fig. 15. Brachiole and main ambulacral plating in hemicosmitoid Caryocrinites ornatus Say, 1825, Wenlock (Silurian), New York, USA. The ambulacra are erect. Each brachiole arises from a pair of main ambulacral plates, the adoral of which are shaded. The aboral plates are more than twice as wide. The first brachiolar plate (1a) is adoral to the second (2) and supports the idea that main ambulacral plates are first brachiolar plates modified to form the ambulacral axis. Redrawn from Sprinkle 1975: 1067, fig. 2.
Fig. 23 in New insights into the origin and relationships of blastoid echinoderms
Fig. 23. Oral surface of the blastoid Codaster acutus McCoy, 1849, early Carboniferous, England, UK, showing interpretation of oral plating. Five ambulacra (A–E) are recumbent on the lancet plates (La) with the floor plates omitted for clarity. Five deltoid plates (d) form the mouth (M) frame and share the adoral parts of the hydrospire slits (H), the aboral parts of which are in the radial plates (R). An, anus. For plate homologies see Table 2.
Fig. 13 in New insights into the origin and relationships of blastoid echinoderms
Fig. 13. Thecal plates of the Silurian blastozoan Lysocystites sculptus Miller, 1889, Indiana, USA, showing homologies with blastoids, coronates, and glyptocystitoids. A–E, Carpenter's (1884, 1891) ambulacra. Unknown erect ambulacra arose from facets (F) on ambulacral plates (RR), homologues of blastoid lancet plates and the radial plates of Rhombifera Barrande, 1867 (Fig. 19A3). The mouth frame is composed of five deltoids (LL, homologues of the lateral plates of Rhombifera). The periproct (P) lies in the CD interambulacrum above additional deltoid plates (white) and shares the gonopore (G). The main body of the cup is formed of five infralateral plates (ILL, homologues of blastoid radials) and three basal plates (BB). Compare with the oral view, Fig. 20B. For plate homologies see Table 2. Redrawn from Sprinkle 1973:140, fig. 34.
Fig. 11 in New insights into the origin and relationships of blastoid echinoderms
Fig. 11. Thecal plates of blastoids. A–E, Carpenter's (1884, 1891) ambulacra; BB, basals; DD deltoids; La, lancets; P, periproct; RR, radials; white plate below periproct in CD interambulacrum is one of the extra deltoids. Dashed lines are radial sinuses for ambulacra. For plate homologies see Table 2.
Fig. 12 in New insights into the origin and relationships of blastoid echinoderms
Fig. 12. Thecal plates of coronates showing homologies with blastoid plates. A–E, Carpenter's (1884, 1891) ambulacra; BB, basal plates; DD, deltoid plates; P, periproct; RR, radial plates; TMP, "trunk mounting plates", which bear facets for erect ambulacra and are homologues of blastoid lancet plates and glyptocystitoid radial plates. White plate above the periproct is the extra deltoid plate and small black oval above that is the gonopore. For plate homologies see Table 2.
Fig. 22 in New insights into the origin and relationships of blastoid echinoderms
Fig. 22. Oral area of the hemicosmitoid Thomacystis tuberculata Paul, 1969, Late Ordovician, Wales, UK. Ambulacra are interpreted as B, C, D, and E and neither ambulacrum A nor plate O3, the oral associated with ambulacrum A, is developed. Facets (F) are shared by ambulacral plates the homologies of which are uncertain, and lateral plates (L1–L9). Primary ambulacral plates (O1, O2, O4, O5). Plate L4 of hemicosmitids is missing in Thomacystis and caryocrinitids (Paul 1984: 144, fig. 87). An, anus; G, gonopore; M, mouth; N, pore for presumed nerves. Redrawn and relabelled from Paul 1969: 193, fig. 2.
Fig. 14 in New insights into the origin and relationships of blastoid echinoderms
Fig. 14. Subterminal branching in the ambulacra of the callocystitid Callocystites fresti Paul, 2015a (SM A.85652), Wenlock (Silurian), Indiana, USA. Open arrow points to the place where the next lateral branch of the radial water vessel would have taken place, but the tip of the radial water vessel was at the top of the brachiole (Br) that arose from the final facet (solid arrow). Thus, branching of the radial water vessel in brachiole-bearing echinoderms was always subterminal. Modified from Paul and Hotchkiss 2020: fig. 7.
Fig. 10 in New insights into the origin and relationships of blastoid echinoderms
Fig. 10. Standard thecal plate diagram for the hemicosmitoid Hemicosmites spp. The theca consists of four circlets, basals (B, BB), infra-laterals (ILL), laterals (LL), and radials (RR). In Hemicosmites spp. there are four BB, six ILL, nine LL and nine RR. Hemicosmites spp. have only three erect ambulacra (A, shared BC, and shared DE) on facets developed on plates R4:R5, R7:R8, and R1:R2, respectively. P, periproct (surrounding anus). Compare with Figs. 6G2 and 9. For plate homologies see Table 2. Redrawn and relabelled from Bockelie 1979: 371, fig. 4a.
Fig. 7 in New insights into the origin and relationships of blastoid echinoderms
Fig. 7. Structure of the erect ambulacra of the coronate Stephanocrinus angulatus Conrad, 1842, Wenlock (Silurian), New York, USA. Biserial brachioles (1–7) are numbered in order of formation and the first two plates (1a, 1b, and so on) are modified to form the main ambulacral trunk. Note that because this is an aboral view, brachiole 1 is anatomically left but on the right of the figure. Redrawn from Brett et al. 1983: 640, fig. 8.
Fig. 9 in New insights into the origin and relationships of blastoid echinoderms
Fig. 9. Thecal plating in the callocystitid Lepadocystis moorei (Meek, 1871), Late Ordovician, Ohio, USA, illustrating typical extraxial plate arrangement in callocystitid and glyptocystitoid rhombiferans. Plates are arranged in four circlets, basals (B, BB), infra-laterals (IL, ILL), laterals (L, LL), and radials (R, RR), plus the orals which are largely axial in origin (see Fig. 5). All glyptocystitoids have four BB, five ILL and LL, and originally six RR. In all callocystitids R5 is absent. In Lepadocystis L5 interrupts the radial circlet and the periproct (P) is surrounded by four plates, IL4, IL5, L4, and L5. Lepadocystis has five, short ambulacra (dashed outlines below letters A–E) on the thecal plates and five pectinirhombs (Rh) across plate sutures B2:IL2, R1:L2, R3:L3, R3:L4, and R6:L1. Currents entered slits in the first mentioned plate and exited from the other plate. For plate homologies see Table 2. Redrawn and relabelled from Kesling 1967: S207, fig. 106.2c.
Fig. 5 in New insights into the origin and relationships of blastoid echinoderms
Fig. 5. Plating in the oral area and in ambulacrum A of the glyptocystitoid Lepadocystis moorei (Meek, 1871), Late Ordovician, Ohio, USA, showing interpretation of ambulacral growth and plate homologies. A–E, Carpenter ( 1884, 1891) ambulacra. Brachiole facets (F) are numbered (A1–A9) in ambulacrum A in the order in which they formed during growth, which cannot be changed after initial development. Each brachiole facet is developed on a pair of floor plates, the adoral of which is shaded pale grey. These are the first brachiolar plates to form during growth and become modified as floor plates. O1–O5, first and 1–5, second oral plates sharing the very first brachiole facet in each ambulacrum. Thus, these are also first brachiolar plates modified as orals. The first facets in each ambulacrum branch to the left as viewed in the growth direction (away from the mouth), but the second (black, F) obey Lovén's Law as restated by Paul and Hotchkiss (2020), AR, BL, CR, DL, ER, where A–E represent Carpenter's ambulacra and L and R represent left and right of the ambulacrum. This is because the ambulacra exhibit the " BD different" pattern of primary brachioles in which ambulacra B and D have the first two brachioles on the left, whereas ambulacra A, C and E have only the first brachiole left. Six primary oral cover plates (dark grey) and the two additional orals, O6 and O7, are identified. O1 and O7 always share the gonopore (G) and hydropore (H) in glyptocystitoid rhombiferans. Redrawn from Paul and Hotchkiss 2020: fig. 6.1 and Kesling and Mintz 1961: fig. 1.
Fig. 6 in New insights into the origin and relationships of blastoid echinoderms
Fig. 6. Examples of pan-dichoporite blastozoans. A. The callocystitid Pseudocrinites bifasciatus Pearce, 1843 (BMNH 40189), Wenlock (Silurian), West Midlands, UK, showing the lenticular theca (above) with peripheral ambulacra, the rapidly tapering proximal stem and more cylindrical distal stem (below) terminating in a root structure. B. The rhombiferid Rhombifera sp. (MPZ 2020/590), Late Ordovician, Zaragoza, Spain, showing the stem attached to the theca, which includes the basal, infralateral and base of the lateral plates (above) and one extensive pectinirhomb. C, D. The fissiculate blastoid Codaster acutus McCoy, 1849, early Carboniferous, Yorkshire, UK. C. Oral view of SM E5400 showing the central mouth, five radiating ambulacra, pectinirhomb-like hydrospires, and circular anus (below). D. Polished lateral view of SM E5398 showing basal and radial plates, anus (A) and very thin hydrospire folds (H). E. One branched ambulacrum of the callocystitid Callocystites jewetti Hall, 1852 (ROM 991N), Wenlock (Silurian), Ontario, Canada, showing the ambulacral structure. The mouth is to the left. Brachiole facets (lower branch) are each on a smaller adoral and larger aboral floor plate. Biserial brachioles alternate along the ambulacrum (Br), so main ambulacral axis has double biserial floor plates. F. Stereo images of the coronate Cupulocorona rugosa Donovan and Paul, 1985 (NMW 2013.8G.65i), Late Ordovician, Carmarthen, Wales, showing theca shape and extensive coronal processes (arrow). Top of the small basal plate is displaced. The rest of the theca up to the tip of the coronal process is formed by the "radial" plates. Erect ambulacra arose within the valleys between the coronal processes. G. The hemicosmitoid Hemicosmites extraneus Eichwald, 1840 (GIT 398–898), Late Ordovician, Madise, Estonia. Lateral view (G1) showing, from below, basal, infralateral, lateral, and radial plate circlets. Cryptorhomb pores can be seen in the infralateral and lateral plates. Oral view (G2) showing three large facets for erect ambulacra.
Fig. 8 in New insights into the origin and relationships of blastoid echinoderms
Fig. 8. Structure of the ambulacra and brachioles in the callocystitid Pseudocrinites bifasciatus Pearce, 1843 (SM A.10192), Wenlock (Silurian), Dudley, UK. First plate (1st) in a brachiole lies directly above the adoral floor plates (grey). This suggests floor plates are the first pair of brachiolar plates modified to form the ambulacral axis. Second and subsequent brachiolar plates white. In callocystitids, floor plates are developed on the thecal surface. A. Side view showing the high profile of the ambulacral axis and rapidly tapering brachiole. B. Impressions of ambulacral floor plates on the thecal plates. Arrows point to the mouth. Redrawn and relabelled from Paul 1967a: 326, figs. 17, 18.
Fig. 4 in New insights into the origin and relationships of blastoid echinoderms
Fig. 4. Origin of Lovén's Law in echinoderms illustrated by the oral plating in edrioasteroid Walcottidiscus Bassler, 1935 (USNM 376690), early Cambrian, locality unknown. A–E, five ambulacra arranged in a 2-1-2 pattern so that three radiate from the mouth, shared DE, A, and shared BC. A1, BC1, and DE1 represent the first ambulacral plates in the three primary ambulacra. A2, BC2, and DE2 represent the second ambulacral plates. When the shared ambulacra bifurcated to give five there was no other position in which the new ambulacral plates could be added than in the acute angle between the dividing primary rays. Thus, the new plates were B2, C2, D2, and E2. Once all five ambulacra reached the stage of having two floor plates, the positions of the first plates obey Lovén's Law as restated by Paul and Hotchkiss (2020), i.e., AR, BL, CR, DL, ER, that is the plates labelled A1, BC1, BC2, DE1, and DE2. Redrawn and modified from Paul and Smith 1984: 454, fig. 7A.
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