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18 Iii. Additions To TULANE STUDIES IN GEOLOGY Volume 2, Number 1 December 30, 1963 CONTENTS THE EVOLUTION OF THE EUCARIDA, (CRUSTACEA, EUMALACOSTRACA), IN RELATION TO THE FOSSIL RECORD MARTIN D. BURKENROAD /)/:) P.lH'l'JlEl\"'l' 01" Gb'O /,oar, 'l'L'LAXE UN lYEIU::il'LT ana ,ll.USEO .i.Y .\CI07Y.JL DH P . \. N.d.11l.cl Page AB sTRACT -------------------------------------- -· --------------------------·------- .. --------------------------------------- 3 I. INTROD u CTI 0 N---- ------------------------------------------------------------------------------------------------- 3 II. THE FOSSIL RECORD ____________________________________________________ ____ -------------------------------------- 4 A. PALEOZOIC 1. Devonian and Carboniferous ------------------------------------------------------------------------- 4 2. Permi an ----------------------------------------- _________________________ ---------------------------------------- 8 B. MESOZOIC 1. Triassic _____ ------------------------------··-------- ---------------------------- ·--------------------------- 9 2. Jurassic ____________ ·-----------------------------------------------------··-------------------------------------··- 11 III. PROBABLE CHARACTERISTICS OF FOSSILS OF THE HYPOTHETICAL PALEOZOIC STEMS OF MA ]OR EUCARID T AXA ---------------------------------------------· ___ ·--··- 14 ACKNOWLEDGMENTS ______________ ______ _______ ____ _ __ -------------------- ------ ·----------------------- 16 LITERATURE CITED ____ --------------------------------------------------------------------- ------------------------- 16 ILLUSTRATIONS TEXT FIGURE 1. Dendrogram of Eucarid evolution________________________________________________ 2 ADDITIONS TO A CATALOGUE OF THE DESCRIBED RECENT AND TERTIARY SPECIES OF ACEST A AND PLICACEST A HAROLD E. VOKES PHOPESNOR UF GEOLOG1' '1TLA1\"E / '}.'1 r' ENSITY I. INTRODUCTION _ _ _____ __ ___ ____ ___ _ __ _ _______ ·------------- __________ --------------------------------- 18 II. ADDITIONS TO THE CATALOGUE OF ACEST A --------------------------------------------------------- 18 Recent species _____ ________________________ ---------------------------------------------------------------- 18 Tertiary species ____________________________________________________________ ·-------------------------------· ___ 18 Species incorrectly referred to Acesta _____________________________________________________ ---·- 19 III. ADDITIONS TO THE CATALOGUE OF PLICACEST A --- -- -- --- -- --------------------------- 20 LITERATURE CITED ___ ------ ---- .. __ - ---- ·--·---------- ·---------- - 20 Jura. \ I I \ I I I I \ 15 I I \ \ I \ I I // ,,/ ' \ I I // / / / II ,- 12 \ I I / / · . / \ I I / /// \ I I \ I I \ I I \ \ I Trias. \ \ I. Superorder EU CARIDA 8 9 10 \ \ 2. Order EUPHAUSIACEA I 3. Order DECAPODA \ \ 0/ 4 . Suborder DENDROBRANCHIATA \ I I I \ I I I // 5. Suborder PLEOCYEMATA, n. 6. Supersection NATANTIA \ I\ ,, p 7. Supersec tion REPTANTIA \ \ I / 8. Section EUKYPHIDA 6 / / 7 \ \ 9. Sec t ion STENOPODIDA \ // I I 10. Section MACRURA \ \ I 11. Section ANOMALA \ \ I 12. Section BRACHYURA \ I 5 13. Subsection THALASSINIDEA \ I Perm. 14. Subsection (GLYPHEIDEA) \ 4 \ I Subsec tion HOMARIDEA \ I I 15. ,1 16. Subsection PALINURIDEA \ (? Palaeopemphix} I \ I ,, \ I ,, \ 13 2 \ I \ I Penn. \I \ Figure 1. Dendrogram of Eucarid Evolution. l 1 \ 2 THE EVOLUTION OF THE EUCARIDA, (CRUSTACEA, EUMALACOSTRACA), IN RELATION TO THE FOSSIL RECORD MARTIN D. BURKENROAD DEPAR'l'JJ1ENT OF GEOLOGY, TULANE UNIVERSITY and JIIUSEO NACIONAL DE PANAMA ABSTRACT I. INTRODUCTION Eumalacostracan fossils from the Missis­ A new consideration of the adult and the sippian indicate the beginnings of Recent developmental characteristics of Recent Eu­ superorders other than the Eucarida; which carid Crustacea indicates that all previously­ latter probably also differentiated around proposed systems of classification of the this time, from a primitive shrimp with members of the order Decapoda are in vari­ carapace sculpture like that of Palaeopalae­ ous degrees polyphyletic (in the sense of mon, in a marine form not yet discovered grouping taxa together some of which are There is no valid evidence tO suggest Eu­ more closely related to members of other carid polyphyly. The only Paleozoic record groups given equal rank; cf. Simpson, 1961, of the su perorder seems to be the peculiar p. 120 ff.). This conclusion (for which the form Palaeopemphix from the Permian, evidence is given in detail in a forthcoming which is certainly not a Glypheid but may treatment of the Recent Eucarida) necessi­ represent an early, calcified offshoot of the tates a critical examination of paleonto­ stem-form of the order Decapoda (the fam­ logical findings, since the primary test of ily P ALEOPEMPHICIDAE, n.). In the any phylogenetic hypothesis is its compati­ early Triassic, not only are the known Eu­ bility with the fossil record. The present carids definitive Decapods, but this order paper is, then, a review of current paleonto­ was already differentiated into the suborders logical ideas from the new point of view, Dendrobranchiata and PLEOCYEMATA and is offered in advance of derailed evi­ ( n.), which were themselves already sub­ dence from the Recent in order to secure divided (e.g., the Peneidae were presumably comment on its weaknesses from specialists already separated from the AEGERIDAE, in fossils. n. fam.). The available Mesozoic represen­ The Recent evidence indicates that the tation undoubtedly includes a dispropor­ following more or less widely-accepted major tionate frequency of forms specialized by taxa are monophyletic: ( 1) the superorder calcification; and this record seems tao late, Eucarida Calman (carapace fused with all fragmentary and non-consecutive to supply thoracic tergites, probably in relation to the crucial evidence either for or against the habit of jumping backward; appendix mas­ present view of Decapod phylogeny founded culina on male second pleopod) ; ( 2 ) the on evidence from Recent forms. The habits Eucarid order Euphausiacea Boas (body-gills and characteristics tO be expected of Pale­ entirely lost but podobranchs including that ozoic fossils representing the hypothetical of the eighth thoracic appendage retained; stems of Eucarid groups are outlined. specializations for sperm-transfer and for EDITORIAL COMMITTEE FOR THIS PAPER: H. K. BROOKS, Department of Geology, University of Florida, Gainesville, Florida ]OHN S. GARTH, Hancock Foundation, University of Southern California, Los An­ geles, California MARTIN F. GLAESSNER, Department of Geology, University of Adelaide, Adelaide, Australia HENRY B. ROBERTS, Division of Marine Invertebrates, United States National Mu­ seum, Washington, D. C. 3 4 Titlane Studies in Geology Vol. 2 pelagic life; ( 3) the Eucar~d order. De~a­ but does seem to show that the conflicting poda Latreille (loss of the eighth epi podite previous deductions from th.e known fo~sils and podobranch, enclosure of all gills in the are not required by the eviclence. A f mal branchial chamber and enlargement of the section of the paper outlines the probable maxillary pump with correlated reduction of characteristics of Paleozoic Eucarid fossils the muscular basal part of the pereiopodal ex­ which (according to the present view) are podites; attenuation of the endopod of the to be hoped for in the gap between the first thoracic appendage and permanent flex­ early, generalized Eumalacostra~a. kn0wn ure of that of the second one) ; ( 4) the from the Devonian and the speciaLzed De­ Decapod supersection Reptantia Boas (loss capod genera ( Peneine, Glypheid and Ho­ of all pleurobranchs anterior to the fifth maridean) so far recorded from the Permo­ thoracic somite; specializations for ben­ Trias and the early Triassic. thonic life) . The Recent evidence unequivocally indi­ II. THE FOSSIL RECORD cates that the "suborder Natantia" of Boas A. Paleozoic: is polyphyletic. The Peneids represent one 1. Devonian and Carboniferous: The of two major branches of the order Deca­ Paleozoic Eumalacostraca have recently been poda and require separation as the sub­ extensively revised by Brooks ( 1962): A~­ order Dendrobranchiata (Bate, redefined). cording to him, the fossil record begms rn They retained dendrobranchiae and a non­ the Devonian with three shrimplike genera incub::uory habit with naupliar eclosion from believed all to have had a more or less en­ the generalized Eumalacostracan that also larged antennular peduncle, undifferentiated gave rise to the marsupial superorder Pera­ biramous thoracic limbs with an undivided carida · but differentiated from the ancestral ' sympod, furcal plates on the telson (how­ Decapod by a few unique specializations, as ever, cf. l.c., Plate 52, fig. 2; and also How­ well as by developing chelae on the fourth ell, 1957, fig. 1) and a well-developed to sixth thoracic legs. The other major carapace, shorter than the pleon but .cover­ branch of the Decapoda, for which the new ing although not fused to all thoraoc ter­ suborder PLEOCYEMATA is required, lost gites. Oostegites have not been se~r:' al­ the secondary rami of the gills and developed though this might be "due to a deficiency pleopodal incubation with zoeal eclosion, but of the fossils" ( l.c., p. 224). apparently continued at first to be acbelat~. One of these Devonian genera ( P alaeo­ This incubatory branch appears to have di­ palaemon) extended into the Low~r .~is­ vided early into two stems. The f irst-dif­ sissippian; and during the later Mississip­ ferentiated of these two, which was especial­ pian
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