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^do^i^cpy ft i THE STRATIGRAPHIC DISTRIBUTION OF THE PALEOZOIC EUMALACOSTRACA
SOME MIDDLE PENNSYLVANIAN HOPLOCARIDA (CRUSTACEA) AND THEIR PHYLOGENETIC SIGNIFICANCE
FREDERICK R. SCHRAM
FIELDIANA: GEOLOGY
VOLUME 12, NUMBERS 13 AND 14 ijrii«ersiiy of Illinois Published by FIELD MUSEUM OF NATURAL HISTORY JAN 28 1970
SEPTEMBER 30, 1969 tkiq
GEOLOGY (JBRAinc
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FIELDIANA: GEOLOGY
* A Continuation of the ^ GEOLOGICAL SERIES oov^^'^S-^^*"^ of FIELD MUSEUM OF NATURAL HISTORY VOLUME 12 FIELD MUSEUM OF NATURAL HISTORY CHICAGO. U.S.A. INDEX Acanthotelson 9, 213 Crustaceans 9, 213-229, 235-286 Adelophthalmus mazonensis 85-100 cydophthalmus, Mazonipterus, n. sp. Aeschronectida, n. ord. 236 85, 101-103 aetherius, Anthrakcris, n. sp. 36-38 Amphibians 9 Deltaic deposition 10, 11 Amphineura 61 Dipnoans 9, 193-209 Annelida 9, 60 Anthracaris 213, 219 edesi, Conchopoma, n. sp. 195 Anthracomedusa, n. gen. 110 Essex Fauna 109, 119, 213 Anthracomedusa turnbidli, n. sp. 110- Eumalacostraca 213-229 112 Eurypterids 85-104 Anthracophausia 213 Anthrakoris, n. gen. 36 Fishes 9, 193-209 Anthrakoris aetherius, n. sp. 36-38 Fossil fauna 8-10 Anthrapalaemon 9, 219 Fossil flora 8-10 Arachnids 9 Fossils, abundance 9 Aratidedhes, n. gen. 251 Fossils, occurrence 4, 8, 9, 15, 121 Aratidedhes johnsoni, n. sp. 252-259 Fossils, preservation 109, 122-127, 137, Archaeostomatopodea, n. subord. 260 155 Arlhropleura 71-76 Francis Creek Shale 3, 10, 11 Arthropleura crislata, n. sp. 79-82 Arthropleurida 9, 71-76, 79-82 Genopterygidae, n. fam. 41-43 Aviculopecten mazonensis 65-66 Genopteryx 43 Glaphyrokoris, n. gen. Bandringa, n. gen. 158 Gl Langford, George 16 Protodonata, 17 Lithoneura 2U Protorthoptera 17, 33-41 Lithoneura carpenieri, n. sp. 24 Liverpool Cyclothem 3 Radula, polyplacophoran 64 linearis 61 Longitubus rayi, Bandringa, n. sp. 160-169 n. 20-24 luciae, Mcluckiepteron, sp. richardsoni, Kallidecthes, n. sp. 238- Lungfishes 9, 193-209 251 rossae, Nacekomia, n. sp. 34-36 219 Mamayocaris Rossites, n. gen. 44 Marine fossils 109-115 11, 59-66, Rossites inopinus, n. sp. 44-45 Mazon Creek 3-5, 8, 9, 15, 16 mazonensis, Adelophthalmus 85-100 Scorpions 171 mazonensis, Aviculopecten 65-66 Serpulidae 60 Mazonia woodiana 172-189 Sharks 59, 157 Mazonipterus, n. gen. 100-101 Spilapteridae 19-24 n. Mazonipterus cyclophthalmns, sp. Strip mines, specimens from 4, 16 101-103 85, Strip mining 3-7 Mcluckiepteron, n. gen. 20 Stomatopoda 260 Mcluckiepteron luciae, n. sp. 20-24 Stylonuridae 100 Medusae 109-115 Syntonopteridae 24-27 Megasecoptera 17, 18 mirabile, Teneopteron 47-52 Teneopteridae, n. fam. 46 mirandus, Glaphyrokoris, n. sp. 38-39 46 Mycterops 85, 103-104 Teneopteron 47-52 Myriapods 9 Teneopteron mirabile theridion, Tyrannophontes, n. sp. 261- 266 Nacekomia, n. gen. 33 Trilobite 59 Nacekomia rossae, n. sp. 34-36 79-82 Nymph, insect 52-54 Trilobitomorpha 71-76, Tullimonstrum gregarium 119-148 n. 110- Octomedusa, n. gen. 112 turnbulli, Anthracomedusa, sp. 112 Octomedusa pieckorum, n. sp. 113- 115 Turnbullia, n. gen. 27 Turnbullia n. 28-31 Opisterostomatopodea, n. subord. 266 priscillae, sp. Ostracodes 9 Tyrannophontes, n. gen. 260 Tyrannophontes theridion, n. sp. 261- 266 Palaeocaris 9, 213 Palaeodictyoptera 17-32 Unnamed crinoid 151 155-156 Unnamed insects 31, 32, 40, 41 Palaeoniscoids 9 Unnamed lungfish 208 Palaeostomatopoda 259 Paleoecology 10-12, 17, 59, 147, Waupecan Sandstone 3 Pelecypods 9, 65-66 whitfieldorum, Howellitubus, n. sp. 60- Perimecturidae 260 61 pieckorum, Octomedusa, n. sp. 113-115 Will County, 111. 3-6 Pleasantview Sandstone 3 woodiana, Mazonia 172-189 Polyplacophora 61 Prestwichianella 9 Xenacanths 9, 59 priscillae, Turnbullia, n. sp. 28-31 Xiphosures 9 TABLE OF CONTENTS PAGE 1. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. Intro- duction. By Eugene S. Richardson, Jr 1 2. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. Insects. By Eugene S. Richardson, Jr 13 3. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. Marine Fauna. By Eugene S. Richardson, Jr 57 4. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. Trilo- bitomorpha, Arthropleurida. By Eugene S. Richardson, Jr 69 5. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. Trilo- bitomorpha, Arthropleurida, II. By Eugene S. Richardson, Jr 77 6. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. Euryp- terida. By Erik N. Kjellesvig-Waering 83 7. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. The Essex Fauna and Medusae. By Ralph Gordon Johnson and Eugene S. Richardson, Jr 107 8. Pennsylvanian Invertebrates of the Mazon Creek Area, Illinois. The Morphology and Affinities of Tullimonstrum. By Ralph Gordon Johnson and Eugene S. Richardson, Jr 119 9. A Crinoid from the Pennsylvanian Essex Fauna of Illinois. By N. Gary Lane 151 10. Bandringa rayi, a New Ctenacanthoid Shark from the Pennsylvanian Essex Fauna of Illinois. By Rainer Zangerl 157 11. Scorpionida: the Holotype of Mazonia woodiana Meek and Worthen, 1868. By Erik N. Kjellesvig-Waering 171 12. New Pennsylvanian Lungfishes from Illinois. By Robert H. Denison . 193 13. The Stratigraphic Distribution of the Paleozoic Eumalacostraca. By Frederick R. Schram 213 14. Some Middle Pennsylvanian Hoplocarida (Crustacea) and Their Phylo- genetic Significance. By Frederick R. Schram 235 THE STRATIGRAPHIC DISTRIBUTION OF THE PALEOZOIC EUMALACOSTRACA SOME MIDDLE PENNSYLVANIAN HOPLOCARIDA (CRUSTACEA) AND THEIR PHYLOGENETIC SIGNIFICANCE FREDERICK R. SCHRAM FIELDIANA: GEOLOGY VOLUME 12, NUMBERS 13, AND 14 Published by FIELD MUSEUM OF NATURAL HISTORY SEPTEMBER 30, 1969 CONTENTS PACE The Stratigraphic Distribution of the Paleozoic Eumalacostraca. by Frederick R. Schram 213 Some Middle Pennsylvanian Hoplocarida (Crustacea) and Their Phylogenetic Significance, by Frederick R. Schram 235 FIELDIANA . GEOLOGY Volume 12, No. 13 September 30, 1969 Publication 1079 The Stratigraphic Distribution of the Paleozoic Eumalacostraca Frederick R. Schram Walker Museum University of Chicago Department of Zoology, Eastern Illinois University INTRODUCTION Johnson and Richardson (1966) deHneated two Middle Pennsyl- vanian faunas within the Mazon Creek area in Grundy, Will, and Kankakee counties in northeastern Illinois —the Braidwood fauna and the Essex fauna. The fossils of these assemblages are found in the ironstone concretions of the Francis Creek Shale, Carbondale Formation, directly above the coal of the Liverpool Cyclothem. These beds are presently classified as being in the Kewanee Group of the Illinois Pennsylvanian (Kosanke et al., 1960). The Braidwood fauna is found along Mazon Creek and in the abandoned strip mines north of the towns of Coal City and Braidwood. The Essex fauna is found in the recently excavated strip mines near the town of Essex. These faunas are considered to represent a marginal deltaic regime, the Braidwood being the freshwater facies while the Essex is a ma- rine one. Crustacea are found in both of these faunas. Five species are de- scribed from the Braidwood localities: Acanthotelson stimpsoni Meek and Worthen, 1865; Palaeocaris typus Meek and Worthen, 1865; Belotelsoji magister Packard, 1886; Anthracaris gracilis (Meek and Worthen, 1865); and Anthracophausia strongi Brooks, 1962c. The Essex fauna contains some 20 recognizable forms, including the above. The proportions of occuiTence of species are quite different between the two faunas. The Braidwood Crustacea are mostly syncarids; the Essex Crustacea are mostly eocarids and hoplocarids. The study of this late Paleozoic fauna, comparable in preservation to that of the Burgess shale in the Middle Cambrian or the Solen- hofen limestone in the Upper Jurassic, is of great import toward un- derstanding the earlj'^ history of the Eumalacostraca. Until now the Library of Congress Catalog Card Number: 73-9291 7 213 214 FIELDIANA: GEOLOGY, VOLUME 12 higher Crustacea have had a record of isolated occurrences through- out the latter half of the Paleozoic. The Paleozoic Eumalacostraca are listed below (Table 1) along with their stratigraphic ranges. H. K. Brooks is at present revising the European forms. Considerable taxonomic confusion exists at the specific level in the European material. No attempt is made to for- mally express synonymies in this paper except in instances where the work is done and in press, where such has been indicated in private conversations with Brooks, or where it has been obvious to me. Stratigraphic correlations between continents and across oceans are difficult. The approximate correlations implied in the table were arrived at using the standard texts of Moore (1958) and Gignoux (1955). Until Brooks (1962c) the Paleozoic Eumalacostraca were very unsatisfactorily classified into the recent orders. The pygocephalo- morphs were referred to the lophogastrid Mysidacea or the "macru- rous decapods." Anthracophausia and Crangopsis were placed in the euphausian schizopods. Palaeomysis and Anthracomysis were con- sidered mysids. Brooks took a number of these Paleozoic problem- atica and classified them together into a new superorder, the Eocarida. This group is characterized by possessing caudal spines and furcae; the protopod of the thoracic appendages as a sympod; and oostegites, seminal receptacles, or both. Although this arrangement is not en- tirely satisfactory it provides a more logical basis upon which to study the Paleozoic Eumalacostraca than the old system of referring them to modern groups. STRATIGRAPHIC DISTRIBUTION Devonian Eocarida The earliest eumalacostracan, Eocaris oervigi Brooks, 1962b, is found in the Middle Devonian of the Rhenish Massif of West Ger- many. The earliest North American eumalacostracan is Devonocaris cuylerensis (Wells, 1957) from the Windom Member of the Moscow Formation, Madison County, New York. The range of the latter genus was extended into the Famennian of the Upper Devonian of Luxembourg province, Belgium, with the assignment by Brooks (1962b) of Palaeocaris destinezi Van Straelen, 1943 to Devonocaris. Palaeopalaemon newberryi Whitfield, 1880 is found in the Ohio Shale, Upper Devonian, and the New Providence Shale, lowermost Mississippian, Kinderhookian, in Lake County, Ohio, and Boyle SCHRAM: PALEOZOIC EUMALACOSTRACA 215 County, Kentucky. Another species, P. iowensis Walter, 1917 oc- curs in the Lower Mississippian Kinderhook Shale near Burlington, Iowa. The eocarids are considered to be the most primitive group of the "caridoid" eumalacostracans and may have been ancestral to some of the higher eumalacostracan groups. They are not only the earliest known but occur throughout the rest of the Paleozoic into the latter part of the Permian and are apparently the last of these early gi'oups to disappear. MiSSISSIPPIAN-LOWER CARBONIFEROUS EOCARIDA The diversity, not only in numbers of species but more signifi- cantly in terms of higher taxa present, markedly increased in the Mississippian-Lower Carboniferous. The Eocarida evidently expe- rienced a notable radiation at that time. Eocaridacea: The genus Anthracophaiisia Peach, 1908 is known in the Lower Carboniferous of Scotland. A. dunsiana Peach, 1908 occurs at the base of the Cementstone group in Berwickshire. Brooks (in press) transferred a number of species assigned by Peach (1908) to the genus Crangopsis into Anthracophausia. They are: A. rhodesi in the Ce- mentstone Group of Northumberland, A. couttsi in the Lower Lime- stone of Lanarkshire, A. magna in the Carboniferous Sandstone Series of Dumfriesshire, A. robusta and A. minuta in the Carboniferous Sandstone Series of Northumberland, and A. hastata in the Cement- stone Group of Northumberland. In addition, the genus Palaeomy- sis Peach, 1908 has also been synonymized by Brooks (in press) with Anthracophausia, embracing the species P. dunlopi of the Soft Band Ironstone in the Lower Coal Measures of Lanarkshire, P. couttsi in the Lower Limestone of the Carboniferous Limestone series in Lanark- shire, and P. tenuis in the Calciferous Sandstone Series of Dumfries- shire. A. traquairi (Peach, 1882) in the Scorpion Bed of Dumfriesshire apparently is the earliest representative of the genus Belotelson Pack- ard, 1886, in view of the more complete picture of the morphology of B. magister now available from the Essex fauna (Brooks, personal communication). The upper limit of the range of Anthracophausia is established by A. strongi Brooks, 1962c from the Middle Pennsyl- vanian Francis Creek Shale of Illinois. The genus Crangopsis Salter, 1863 is apparently restricted to the Lower Carboniferous. Four species are in the literature: C. socialis Salter, 1861 in the Abden Limestone of East Fife, C. eskdalensis 216 FIELDIANA: GEOLOGY, VOLUME 12 (Peach, 1883) of the Calcareous Sandstone Series, Dumfriesshire, C. elegans (Peach, 1883), probably synonymous with C. eskdalensis (Brooks, personal communication) in the Cementstone Group near Duns, and C. huxleyi (Woodward, 1867) in the Lower Coal Measures of Paisley. Pygocephalomorpha : The Order Pygocephalomorpha Beurlen, 1930 appeared in the Lower Carboniferous. These creatures represent an early "crab-like" radiation and were formerly thought to be macrurous decapods or primitive schizopods. The family Tealliocarididae is exclusively Lower Carboniferous. Tealliocaris Peach, 1908 is restricted to the Calciferous Sandstone Series. Several species are currently recog- nized in the literature: T. loudensis Peach, 1908 and T. woodwardi (Etheridge, 1877) in the Wardie Shale, Oil Shale Group, Haddington- shire, Roxburghshire, and Northumberland; T. robusta Peach, 1908 in the Lower Limestone Group of Lanarkshire; T. tarrassiana Peach, 1908 in the Cementstone Group of I^anarkshire, Dumfriesshire, and Berwickshire; T. etheridgei Peach, 1908 in the Calciferous Sandstone Series of Dumfriesshire; and T. formosa (Peach, 1882) in the Dum- friesshire Scorpion Bed. Three species of the pygocephalomorph genus Pseudogalathea Peach, 1883 have been placed in the literature: P. macconochei (Eth- eridge, 1879) in the Cementstone Group of Roxburghshire, P. rotunda Peach, 1883 in the Calciferous Sandstone Series of Northumberland, and P. ornatissima (Peach, 1882) in the Cementstone Group and Scorpion Bed of Northumberland and Dumfriesshire. MississiPPiAN Syncarida The first representatives of the Superorder Syncarida Packard, 1885 are in the Mississippian of North America. These species are classified in the Order Palaeocaridacea Brooks, 1962a. Palaeosyn- caris dakotensis Brooks, 1962c occurs in the uppermost Mississippian in the Heath Shale, Chesterian, Stark County, North Dakota. Squil- lites spinosus Scott, 1938 is also found in the Heath Shale in Fergus County, Montana. Mississippian-Lower Carboniferous Hoplocarida Brooks (1962c) established the Order Palaeostomatopoda to in- clude the earliest hoplocarid genera, Archaeocaris and Perimecturus, as primitive precursors of the stomatopods. Archaeocaris vermi- SCHRAM: PALEOZOIC EUMALACOSTRACA 217 species genera families orders 40r 35 - 30 - 25 - 20 - 15 - 10 - 5 - Lower Upper Carboniferous Carboniferous Fig. 102. Graphic distribution of the species, genera, families, and orders of Paleozoic Eumalacostraca. The upper solid line is the number of species described in the literature; the lower solid line is the number of species after probable syn- onyms are eliminated. formis Meek, 1872 occurs in the New Providence Formation, Kinder- hookian, of Boyle County, Kentucky. A. graffhami Brooks, 1962c is found in the Chaney Shale, Chesterian, of Pontotoc County, Okla- homa. The genus Perimecturus Peach, 1908 extends from the Lower Carboniferous to the Upper Carboniferous of Europe. P. parki (Peach, 1882) was first described from the Cementstone Group and Scorpion Bed of Dumfriesshire. In 1908 Peach described five more species: P. stocki from the Wardie Shale of Midlothian, P. elegans 218 FIELDIANA: GEOLOGY, VOLUME 12 and P. communis of the Dumfriesshire Scorpion Bed, P. ensifer in the Cementstone Group of Langholm, and P. pattoni of the Lower Limestone in the Lanarkshire Carboniferous Limestone Series. Van Straelen (1922) described two species from the Lower Westphahan near Li^ge, Belgium : P. fraiponti and Anthracomysis rostrata. Brooks (in press) has identified the holotype of A. rostrata as the tail fan of P. fraiponti. MississippiAN-Lower Carboniferous Peracarida (?) Two problematic Paleozoic crustacean species are classified as the Order Anthracocaridacea Brooks, 1962c. This order was placed by Brooks tentatively in the Peracarida. Anthracocaris scotica (Peach, 1882) in the Calciferous Sandstone Series of Dumfriesshire was orig- inally described as a species of Palaeocaris but its separate position was recognized by Caiman (1932b). Palaeocaris lansbouroughi Peach, 1908 from the Coal Measures of Ayrshire is probably a member of the genus Acadiocaris Brooks, 1962c. Acadiocaris novascotica (Cope- land, 1957a) was discovered in the Horton Group, Osagean, Lower Mississippian, of Inverness County, Nova Scotia. The known speci- mens of Anthracocaris and Acadiocaris do not display the ventral surfaces which might carry the taxonomically definitive oostegites. Brooks (1962a, c) has offered the alternative explanations that the Anthracocaridacea either are true peracarids, resembling the Tanai- dacea, or they are transitional to the Syncarida, marking the depar- ture from the general caridoid ancestor. No satisfactory solution can be adduced for the present. Pennsylvanian-Upper Carboniferous Eocarida The Pennsylvanian-Upper Carboniferous appears to have been the time of greatest diversity of the Paleozoic Eumalacostraca. This impression may simply derive from the fact that the rich Mazon Creek faunas happen to have been preserved from the Pennsylvanian rather than some other period. Eocaridacea: As was mentioned above, two eocaridaceans have been described from the Francis Creek Shale, Belotelson magister and Anthracophau- sia strongi. The former species has been previously known from only three specimens, all found a century ago along the banks of Mazon Creek. It is the most abundant, well-preserved eocarid in the Essex fauna. Its morphology may now be reconstructed in great detail and SCHRAM: PALEOZOIC EUMALACOSTRACA 219 may require separating this species from the Anthracophausiidae where Brooks has tentatively placed it. There are a few other cari- doid forms in the Essex fauna which appear to be eocaridacean. Pygocephalomorpha : The Pennsylvanian pygocephalomorphs belong to the family Py- gocephalidae Brooks, 1962c. Members of the genus Pygocephalus Huxley, 1857 have had a varied taxonomic history (Brooks, 1962c and in press). Specimens have been assigned to the notostracan genus Apus by Prestwich (1840) and have been refeiTed to under four other generic names now synonymized with Pygocephalus: An- thrapalaemon Salter, 1861, Palaeocarabus Salter, 1861, Diplostylus Salter, 1863, and Necroscilla Woodward, 1879. There is also consid- erable confusion as to how many species there are. Van der Heide (1951) recognizes three: P. dubius (Milne-Edwards, 1840) of the Westphalian of Scotland, Continental Europe, and Canada; P. gros- sarti (Salter, 1861) in the Namurian of Scotland; and P. holti (Wood- ward, 1911) of the Middle Coal Measures of Rochdale, Lancashire. The other pygocephalids have not had such a confused history. Anthracaris gracilis (Meek and Worthen, 1865) of the Francis Creek Shale was originally referred to as Anthrapalaemon. Brooks, 1962c established its separate position. A. gracilis occurs principally in the Braidwood fauna and only very rarely in the Essex. A more abun- dant Essex pygocephalid is a new species of the genus Mamayocaris Brooks, 1962c, a genus previously known only from the Lower Per- mian. Brooksocaris aisenvergi Birshtein, 1966 is found in the Lower Stephanian of the Don Basin in the Soviet Union. Pseudotealliocaris caudafimbriata (Copeland, 1957b) occurs in the Canso Group, West- phalian, of Nova Scotia. Tealliocaris belli and T. barathrata Cope- land, 1957b have been suppressed as synonyms of P. caudafimbriata by Brooks (1962c). Pennsylvanian-Upper Carboniferous Syncarida The syncarids were first discovered in the Mazon Creek concre- tions some 30 years before the living forms began to be described. Acanthotelson stimpsoni Meek and Worthen, 1865 formed the basis for the superorder Syncarida (Packard, 1885). Besides being found in the Carbondale Formation of Illinois it also occurs in the lower Shelburn Formation of Indiana in strip mines south of TeiTe Haute. Palaeocaris typus Meek and Worthen, 1865 is found in both the Mazon Creek faunas and at Terre Haute. Other legitimate species 220 FIELDIANA: GEOLOGY, VOLUME 12 of Palaeocaris are P. canadensis Brooks, 1962c in the Riverdale and Pictou Groups, Westphalian, of Nova Scotia; P. burnetti Woodward, 1881 in the Coal Measures of the Irwell Valley of Lancashire; and P. praecursor (Woodward, 1908) occurring in the clay ironstone con- cretions of the Coal Measures of Ilkeston, Derbyshire. Some species of more questionable validity are P. lohesti Van Straelen, 1922 from the base of the Chokier, Westphalian, near Li^ge; and P. vander- grachti Pruvost, 1922 from the same strata, Woensdrecht, The Neth- erlands. Finally, there is Pleurocaris annulatus Caiman, 1911a from the Coal Measures near Cosely, England. Pennsylvanian Hoplocarida The Essex fauna contains some new hoplocarids. One abundant species forms the basis of a new order which represents a swimming radiation of this superorder (Schram, 1967). It is related to another species referred to by Zangerl and Richardson (1963) from the humu- lite facies of the Logan Quarry Shale. The Essex fauna also contains the earliest stomatopod, which forms a morphologic link between the palaeostomatopods and the later stomatopods. (In addition to the other Crustacea mentioned above, the Essex fauna also contains a few non-eumalacostracans. There are two spe- cies of ostracodes and three of branchipods. Leptostraca are rare. They are represented by a species of Dithyrocaris, perhaps a Ceratio- caris, and a new form which appears to be allied with Sairocaris (Peach, 1883). There are some strange forms which may be Lepto- straca or may even constitute an entirely new group.) Permian Eocarida In the Permian the Paleozoic Eumalacostraca were in decline. Of the Eocarida only the pygocephalomorphs remain. Mamayocaris jepseni Brooks, 1962c in the Vale and Opeche Formations of Okla- homa and South Dakota marks the upper range of this genus. The family Notocarididae Brooks, 1962c is confined to the southern hemi- sphere. Notocaris tapscotti (Woods, 1923) is found in the Upper Dwyka Shale outside of Kimberley, South Africa. Pygaspis gins- hurgi Fabre and Huard, 1967, in the Upper Dwyka near Lainsburg, South Africa, is surely a member of the genus Notocaris and is prob- ably synonymous with N. tapscotti. The genus Paulocaris Clarke, 1920 (=Liocaris Beurlen, 1931, = Pygaspis Beurlen, 1934) is the South American notocarid. P. pachecoi Clarke, 1920 occurs in the Upper Permian Irati Formation of Sao Paulo state, Brazil. Beurlen has SCHRAM: PALEOZOIC EUMALACOSTRACA 221 placed a number of species, anatomically similar to P. pachecoi, in the literature from these same Irati beds in Sao Paulo, Parafia, and Rio Grande do Sul: P. heuni and P. angusta 1931, P. brasilicus and P. quadrata, 1935, and P. clarkei and P. marianoi, 1953. Permian Syncarida The syncarid genus Palaeocaris, familiar in the Pennsylvanian, is also found in the Lower Permian: P. krejcii (Fritsch, 1875) of the Gaskohle and Kalkstein near Pilsen, Bohemia; and P. rochei (Brocchi, 1880) of the Autun Shale in the Autun basin in France. The earliest stygocaridacean syncarid, Clarkecaris brazilicus (Clarke, 1920) of the Irati Shale of Brazil, was recognized as a distinct genus by Messalira (1952). Uronectes fimbriatus Jordan, 1847 occurs in the Steinkohl of Saarbriicken and the Lebach of Bohemia. Permian Eucarida The Permian also contains the roots of the Mesozoic and Ceno- zoic radiations. Palaeopemphix sosiensis, P. affinis, and P. meyeri Gemmellaro, 1890 were described from the Lower Permian of Sicily. They have been considered primitive decapods. Birshtein (1958) described an erymaid decapod, Protoclytiopsis antiqua, from the Up- per Permian deposits of western Siberia. Permian Peracarida The earliest definite peracarids appear in the Permian. Palaeo- phreatoicus sojanensis Birshtein, 1962 from the Kazan Formation near Archangel formed the basis for a new family, the Palaeophrea- toicidae. At that time Birshtein also placed in this family Palaeo- crangon problematicus (Schlotheim, 1820) which is found in the Zech- stein of Gluckbrunn, Germany. It had been removed from the trilobites by Schauroth (1854) where it had been originally placed. Glaessner and Malzahn (1962) described some peracarids from bore- holes into the Zechstein in the lower Rhine region. One was another palaeophreatoicid, Protamphisopus reichelti. A second, an apseudid tanaidacean, Ophthalmapseudes rhenanus, was a redescription of a supposed decapod discussed by Malzahn (1958). Finally, they men- tioned an isolated, pyritized thorax which bore certain similarities to the modern cumacean, Nannastacus, but due to insufficient material it was tentatively identified as belonging to a nebaliacean in the Zechstein fauna. Ph s c 5 S O .2 ^ ~ s: vi ^ " 1^ c C« "^ o s 2 -a i: m X E I ^ §. •r o 3 s o c^ cc C C EC 'c 0, cu Q. a; cC a,' a; r,: rt; O o S aj a^ g "I G, CU O O ^ I c a> o O 4-* 02 cS Eh 225 i I ^ ec CO CO ;£ B S o 2 ?s -§ *^ X f5 - a; a; q: q; Q,' a; c: q; CO 1 SCHRAM: PALEOZOIC EUMALACOSTRACA 229 COMMENTS The problem of synonymies is a very real one in evaluating the record of the Paleozoic Eumalacostraca. Much of the descriptive work was done in the last century when the typological concept of systematics was in vogue. This is especially evident in the work of Peach who, while making reliable observations, employed variations to justify the erection of new species. In Peach's work the problem can be easily perceived. Other early taxonomic assessments cannot be as easily judged since the original descriptions are not detailed enough to assess without locating the original material and examin- ing it anew. This problem is not restricted to the early workers. The errors of typology have persisted down to the present day. Some of this confusion has been rectified at least in regard to the genera and some suites of species. It seems improbable to me that there would be ten true species of Anthracophausia in the Calciferous Sand- stone Series of Scotland or seven species of Paulocaris all in the same stratigraphic horizon. The broad definition of a species based on the recognition of variability within populations is opposed to the narrow typologic definitions of the above. Pygocephalus dubius or Palaeo- caris typus, where detailed anatomical studies have been made, have been shown to have existed over a period of time virtually unchanged. An approach taking variations into account is to be considered in- dicative of a more natural taxonomy. The ranges of the various taxonomic categories are presented in Table 1. Although the greatest number of registered species is in the Mississippian-Lower Carboniferous, the Pennsylvanian-Upper Carboniferous is more diverse in terms of families and orders present. Figure 102 graphically presents distribution. The orders, families, and genera of Paleozoic Eumalacostraca are presented for the De- vonian, Carboniferous, and Permian periods. A steady increase is registered with a peak reached in the Pennsylvanian-Upper Carbo- niferous after which the Permian decline is noted. The described species, upper solid line, reach a peak in the Mississippian-Lower Carboniferous. If probable synonymies are taken into consideration, lower solid line, the effect of taxonomic "splitting" can be seen. The Mazon Creek crustacean faunas occur at the peak of this Paleozoic radiation. Succeeding papers in this series will focus on various aspects of this history which, hopefully, will result in a clear understanding of this radiation, i.e., its relation to what went before and what followed, including the development of the modern fauna. 230 FIELDIANA: GEOLOGY, VOLUME 12 ACKNOWLEDGEMENTS The author wishes to thank all the people who have read the manuscript and offered comment, especially Drs. R. G. Johnson, A. M. Ziegler, and J. R. Bolt of the University of Chicago and Dr. E. S. Richardson, Jr. of Field Museum of Natural History. Work was supported in part by National Science Foundation grant GB 5772. REFERENCES Beurlen, K. 1930. Vergleichende Stammesgeschichte Grundlagen Methoden, Probleme un- ter besonderer Buriicksichtigung der hoheren Krebse. Fortschr. Geol. Paleo., 8, pp. 317-464. 1931. Crustaceenreste aus den Mesosaurierschichten (Unterperm) von Bra- silien (Sao Paulo). Paleo. Z., 13, pp. 35-50, 7 figs. 1934. 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