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Echinodermata THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS ECHINODERMATA ARTICLE 6 Pages 1-16, Plates 1-4, Text-figures 1-8 A LIVING SOMASTEROID, PLATASTERIAS LATIRADIATA GRAY By H. BARRACLOUGH FELL UNIVERSITY OF KANSAS PUBLICATIONS JULY 9, 1962 THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS Echinodermata, Article 6, pages 1-16, Plates 1-4, Text-figures 1-8 A LIVING SOMASTEROID, PLAT ASTERIAS LATIRADIATA GRAY By H. BARRACLOUGH FELL Department of Zoology, Victoria University of Wellington, New Zealand CONTENTS PAGE PAGE ABSTRACT 4 PINNATE FASCIOLAR GROOVES 13 INTRODUCTION 4 NUTRITION 14 MATERIAL EXAMINED 5 ABORAL SKELETON 14 BODY SHAPE 5 SYSTEMATIC POSITION 15 ARM STRUCTURE 6 REFERENCES METAPINNULES 11 15 BUCCAL SKELETON 12 ADDENDUM 16 ILLUSTRATIONS PLATE FACING PAGE FIGURE 1. Platasterias latiradiata GRAY. Aboral aspect of ventral skeleton after exposure. Specimen from disc and one arm. Specimen from Corinto, Corinto, Nicaragua 12 4 Nicaragua 4. Platasterias latiradiata GRAY. Oblique ventral 2. Platasterias latiradiata GRAY. Adorai aspect of aspect of part of a regenerating arm, partially disc and one arm of specimen illustrated in dissected to expose the skeleton. Holotype, from Plate 1 5 Tehuantepec, Southern Mexico, now in British 3. Platasterias latiradiata GRAY. External aspect of Museum of Natural History, London 13 FIGURE PAGE FIG URE 1. Chinianaster levyi THORAL (Chinianasteridae) 5 6. Platasterias latiradiata: dissection of buccal skeleton and arm-base 10 2. Villebrunaster thorali SPENCER (Chinianasteri- dae) 6 7. Platasterias latiradiata: A, furrow aspect of fur- row-wall; B, block-diagram of virgalia and fas- SPENCER (Archegon- 3. Archegonaster pentagona ciolar channels; C, transverse section of arm, 7 asteridae) near tip, in regenerating region (holotype) 11 4. Platasterias latiradiata GRAY (Platasteriidae): 8. Paxillae: A, Platasterias latiradiata GRAY (Plata- A, virgalia in ventral oblique aspect; B, trans- steriidae, Somasteroidea); B, Psilaster acumina- verse section of base of arm 8 tus SLADEN (Astropectinidae, Asteroidea); C, 5. Platasterias latiradiata: internal aspect of ven- Luidia neozelanica MORTENSEN (Luidiidae, tral skeleton of basal region of arm 9 Asteroidea) 15 ABSTRACT Platasterias latiradiata GRAY, hitherto supposed to be posed evidence of external ampullae in fossil somasteroids an aberrant luidiid asteroid, is shown to have funda- is considered to have been a misinterpretation. SPENCER ' S mentally the same structure as the Cambro-Ordovician inferred homologies between somasteroid virgalia and Somasteroidea, and must accordingly be referred to that asteroid adambulacral and marginal plates are confirmed, group of Asterozoa. The pinnate character of the skele- and the superambulacral plate is also shown to be an ton of the somasteroid arm, first elucidated by SPENCER occluded virgalium. A series of adductor muscles be- (1951), is shown to extend also to the external alimentary- tween the ambulacral plates permits partial, temporary respiratory system, and to have been the basis from which erection of a shallow furrow, approximately of asteroid the asteroid morphological features arose, by successive type; the triangular depressions in which these muscles changes in axial gradients during development. Features are inserted are represented by corresponding structures not conserved in fossil somasteroids include paired erectile in Chinianaster, and accordingly it is considered that a webs on either side of the inter-pinnular fasciolar grooves, shallow, temporary furrow must have been erected during incorporating serially arranged cover-plates, closely simu- feeding activity in at least some Paleozoic Chinianasteri- lating those of crinoids. The buccal skeleton, though ap- dae. The family Platasteriidae Caso is redefined. Possible proximating that of asteroids, is demonstrably of pinnate derivation of all astroradiate extant echinoderms from structure, which is elucidated. Enlarged external cupules crinoids is postulated, on the basis of the marked resem- accommodate the tube-feet when retracted, and the sup- blances to crinoids now evident in somasteroids. INTRODUCTION As recently announced, Platasterias latiradiata is thence to the small, central mouth. The ambulacral a surviving member of the Somasteroidea, an archaic ossicles are not erected to form an inverted-V, as in group of star-shaped echinoderms hitherto thought asteroids, but are placed in a more or less recumbent to have become extinct early in the Ordovician Period attitude; thus the radial furrow is not invaginated (FELL, 1961). A brief account of the analytical into the arm, but is merely a shallow, narrow groove method by which the affinities of the animal were first underlying the ambulacral ossicles. As can now be realized has already been given (FELL, 1962). This inferred from the living example, a Y-shaped depres- contribution is limited to brief discussion of the major sion on the median aspect of the ambulacral ossicles features of somasteroid anatomy, as illustrated by houses a transverse adductor muscle, capable of pro- Platasterias, with the stress mainly upon endoskeletal ducing a partial, temporary elevation of the median structures of paleontological significance. A more de- ventral surface of the arm, during selective detrital tailed discussion of axial gradients and their theoreti- feeding; the presence of similar Y-shaped depressions cal implications will be presented elsewhere. in at least Chinianaster, among the lower Paleozoic Somasteroids differ from all other echinoderms in members, suggests that this muscle, and the mode of having an extremely flattened, star-shaped body, in feeding it facilitates, may be a more generalized char- which the skeleton of the arms resembles that of bi- acter of early somasteroids. As we can also infer from serial arms of crinoids, the former being built of the living example, somasteroids are characterized by elongate rods (called virgalia) arranged in obliquely having simple tube-feet, without suckers, and a blind transverse rows on either side of the axial series of gut resembling that of ophiuroids and luidiid aster- ambulacral ossicles. The rows of virgalia in the more oids. SPENCER (1951) regarded the somasteroids as unspecialized somasteroids are arranged like the pin- ancestral to both asteroids and ophiuroids, and ranked nules of a biserial crinoid arm, and form the adjacent them as a subclass of Asterozoa. This view is sup- walls of intervening grooves, which apparently con- ported, but Platasterias is clearly seen to lie near the vey water-currents to the radial food-groove, and line of descent which led to luidiid asteroids. EXPLANATION OF PLATE 1 FIGURE 1. Platasterias latiradiata GRAY. Aboral aspect of disc and one arm. Specimen from Corinto, Nicaragua, Allan Hancock Foundation, Velero III Station 962-39, 2-5 m, 4 May 1939. R, 64 mm., r, 11 mm. Photo M. D. KING. UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS ECHINODERMATA, ARTICLE 6 PLATE 1 FELL - Living Somasteroid UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS PLATE 2 ECHINODERMATA, ARTICLE 6 FELL - Living Somasteroid LIVING SOMASTEROID 5 Photographs accompanying this paper were made Museum, and of Miss AILSA M. CLARK, the curator by M. D. KING, Victoria University of Wellington. of the Department of Echinoderms. Subsequently, Text-figures (except Figures 1-3) were drawn by me. five more or less complete specimens were received from the Allan Hancock Foundation, University of Southern California, by courtesy of Captain F. C. ZIESENHENNE. This latter material, all in dried condi- tion, was obtained off Corinto, Nicaragua, in shallow water. The presence of fine sand-grains in the fascio- lar grooves of both the Tehuantepec and Corinto material suggests that the species normally frequents soft, sandy botom, the recorded depth being 2 to 5 m. Although very little museum material exists, and most of it derives from one Velcro Ill station, it seems likely that the species is local, rather than rare. BODY SHAPE The highly characteristic outline of the body in Platasterias (Pls. 1, 2) is due to the petaloid form of the arms, which are narrow basally, broaden rapidly, and then narrow again distally; as a consequence, the bases of the arms are separated by deep, narrow inter- radial fissures, the disc being confined to a relatively small central area. Whereas no asteroid is known to have a body of this form, it is characteristic of the chinianasterid somasteroids (Figs. 1, 2) and is evi- dently an original feature of somasteroids, since the Chinianasteridae are the least specialized members of the group. Among other echinoderms, only crinoids possess petaloid arms. In the more specialized soma- steroids, for example Archegonaster (Fig. 3), the body assumes a pentagonal outline, and the virgalia are partly stabilized as adambulacral, intermediate and marginal elements. Platasterias exhibits stabilization of the virgalia, but in other respects is evidently much FIG. 1. Chinianaster levyi THORAL (Chinianasteridae), more primitive than Archegonaster, from which it Lower Ordovician, France, x4.5 (after Spencer, 1951). differs widely in general appearance. In dorsal aspect each arm presents a mid-radial MATERIAL EXAMINED ridge (Pl. 1) caused, not by any underlying ambulac- ral furrow, but by the massive capitula of the ambu- Platasterias latiradiata GRAY, 1871, was described lacral ossicles themselves. The older descriptions on the basis of a single specimen from Tehuantepec, (GRAY, 1871, et auctt.) refer to a longitudinal
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