t\vs HARVARD UNIVERSITY Library of the Museum of Comparative Zoology us ISSN 0027.4100 bulletin OF THE Museum of Comparative Zoology A Review of the North American Fossil Amiid Fishes JOHN R. BORESKE, JR. HARVARD UNIVERSITY CAMBRIDGE, AAASSACHUSEHS, U.S.A. VOLUME 146, NUMBER 1 18 JANUARY 1974 PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Bulletin 1863- Breviora 1952- Memoibs 1864-1938 JoHNSONiA, Department of Molliisks, 1941- OccAsioNAL Papers on Mollusks, 1945- Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of tlie Gulf of Maine. Reprint. Bnies, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- malian Hibernation. Peters'. Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae (Mollusca: Bivalvia). Whittington, H. B., and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolu- tion of Crustacea. Proceedings of the New England Zoological Club 1899-1948. (Complete sets only.) Publications of the Boston Society of Natural Histor>'. Authors preparing manuscripts for the Bulletin of the Museum of Comparative Zoology or Breviora should send for the current Information and Instruction Sheet, available from Editor, Publications OflBce, Museum of Comparative Zoology, Harvaid University, Cambridge, Massachusetts 02138, U.S.A. (g) The President and Fellow* of Harvard Colleee 1974 A REVIEW OF THE NORTH AMERICAN FOSSIL AMIID FISHES JOHN R. BORESKE, JR.i CONTENTS Abstract 1 Introduction 2 Acknowledgments 2 Abbreviations 3 A7?Jia calva 3 Nomenclature 3 Ecology 4 Geographic Distribution 4 Pleistocene Occurrences 4 Diagnosis 5 Morphometries 5 Methods 6 General Proportions and Growth 8 Comparisons with Fossil Forms 10 Discussion 17 Meristics 18 Supravertebral Scale Rows 18 Branchiostegal Rays 20 Fin Rays 20 Vertebral Elements 25 Vertebral Column of Amia calva 28 Vertebral Features 28 Vertebral Dimensions 33 Valid North American Fossil Genera and Species 37 Amia fragosa 37 Amia uintaensis 47 Amia cf. uintaensis 64 Amia scutata 66 Amia cf. scutata 70 Amia cf. calva 72 Amiidae incertae sedis 72 Specimens Removed from the Amiidae 74 Summary and Conclusions 75 Literature Cited 81 Plates 84 Abstract. North American amiid fishes range from Cretaceous ( Albian ) to Recent. Amiids are 1 Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 Bull. Mus. Comp. Zool, 146(1): 1-87, January, 1974 common fossils in Late Cretaceous and Tertiary freshwater deposits and apparently occupied a habitat much like that of the Recent species Amia calva. Morphometric, meristic, and cranial char- acters of articulated specimens from the Fort Union Fonnation (Paleocene), Green River For- mation (Eocene), Florissant Fonnation (Oligo- cene). Pawnee Creek Formation (Miocene), and a Recent A. calva sample from Wisconsin have been used here in an attempt to revise the taxon- omy and evolutionary history of the group. Whereas seven genera and twenty-three species of fossil amiids have been described on the basis of disarticulated, often isolated elements, only three taxa have heen described from complete or partially complete material. Amia fragosa (Late Cretaceous to Middle Eocene), A. uintaensis (Pal- eocene to Early Oligocene), A. scutata (Early to Middle Oligocene), and A. calva (Middle Plio- cene to Recent) are here considered the only valid taxa. Amiid remains are first known in the North American fossil record from the Early Cre- taceous (Albian) Paluxy Formation of Texas. This disarticulated material shows resemblances both to A»ii« and to the Late Mesozoic European genera Uroclcs and Arniopsis. Paramiatus gurleyi (Romer and Fryxell, 1928) from the Green River Formation of Wyoming is a synonym of A. frag- osa. Tlie differences between Amia and the large Early Cenozoic form Protamia are insufficient for recognition of Protamia as a genus distinct from Amia. The Eocene and Oligocene forms Protamia media, Pappichthys medius, P. plicatus, P. scler- ops, P. laevis, P. symphysis, P. corsonii, Amia whiteavesiana, and A. macrospondyla are s\monyins of A. uintaensis; they were based on undiagnostic cranial and vertebral characters. Morphometric and meristic similarities indicate that little evi- dence exists for maintaining separate Oligocene species A7nia scutata and A. dictyocephala. Amia exilis is a synonym of Amia scutata; it was based on undiagnostic vertebral characters. A. scutata is morphometrically distinguishable from A. calva only on the basis of a slightly larger head/stan- 1 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 dard-length ratio. The Eocene ta.xa Amia de- pressus, A. newherrianus, A. gracilis, and Htjpamia elegans are nomina diibia. Comparison of the fossil forms with the Recent Amia calva suggests the following ta.xonomic and possible phylogenetic relationships: ( 1 ) Amia frag- osa survived until the Middle or Late Eocene, with no phylogenetic affinities with the modern form; (2) Amia iiintaensis appears to be closer than Amia fragosa to the ancestry of Amia calva, which evolved through an intermediate fonn such as Amia scutata; (3) establishment of the Recent species Amia calva had begun at least by the be- ginning of the Pliocene; and (4) diere are simi- larities in the Paleocene and Eocene amiid fossil record of North America and Europe. INTRODUCTION Aviia is a genus of freshwater fishes that includes one of two extant representatives of the holostean level of organization. It includes a number of species of which only Amia calva exists today; other forms of Amia are found in the fossil record, and extend from the Late Cretaceous to ap- proximately the Middle Pliocene. This study is an attempt to determine the taxon- omic and phylogenetic relationships among the various species of Amia. It is established on osteology as well as on morphometric and meristic data from both Recent and fossil forms. This data is used to compare the available features of the fossil forms with Recent Amia calva and to detennine the validity of previous descriptions based on various osteological, morphometric, or meristic character-states. Until recently, a major difficulty in inter- preting the taxonomy of fossil amiids has been the paucity of articulated specimens. Five genera and twenty-one species of fossil forms have been described from disarticu- lated, often isolated, elements (Table 19); only two taxa have been described from articulated specimens: Paramiatus ii^iirleyi (Romer and Fryxell, 1928) and Amia scutata (Osborn et al., 1875). Recent works by Estes (1964) and Estes and Berberian ( 1969 ) , based on disarticulated elements from the Late Cretaceous Lance and Hell Creek formations, are the only published studies of Amia fragosa, although O'Brien (1969) completed an M.A. thesis on the osteology of A. frap.osa, describing articu- lated specimens from the Late Cretaceous Edmonton Formation of Alberta. Much more articulated material is now available and provides more detailed in- formation on the cranial and postcranial anatomy of amiids. These specimens have been useful in this revision of the taxonomy as well as in the determination of possible relationships to European and Asian forms. In an attempt to understand the evolution and interrelationships of the fossil and Re- cent amiids, a growth-series study has been made on a Recent A. calva sample from Wisconsin, and is compared moi"phometri- cally and meristically with the fossil forms. A great number of fossil specimens, includ- ing the holotypes and paratypes of all North American amiid species, have been exam- ined. Several European taxa have been studied at the British Museum ( Natural History), London; Museum National d'His- toire Naturelle, Paris; and the Institut Royal des Sciences de Belgique, Brussels. ACKNOWLEDGMENTS I am especially grateful to Professor Richard Estes (University of California at San Diego) for his advice and criticism in the preparation of this manuscript. Cecile Janot- Poplin and Sylvie Wenz ( Museum National d'Histoire Naturelle, Paris), and Karel Liem ( Museum of Comparative Zo- ology) read the manuscript and offered criticisms that substantially improved the text. Additional thanks are due to Donald Baird ( Princeton University ) , Henry Booke and Bany Cameron (Boston University), William J. Hlavin (Cleveland Museum of Natural History), Farish A. Jenkins, Jr. (Museum of Comparative Zoology), Charles Meehan ( Chamberlayne College), Robert R. Miller (University of Michigan), David Pariis ( New Jersey State Muse- um), Colin Patterson (British Museum of Natural History), Clayton Ray (National Museum of Natural History), Bobb Schaef- fer (American Musevun of Natural History), Hans-Peter Schultze ( Geologisch-Paleon- I Fossil Amiids • Borcske tologisches Institiit der Gcorg-Aiigust-Uni- vcrsitiit, Gottingen), Keith Thomson (Yale University), and Hainer Zangerl (Field Museum of Natural History) for their help- ful suggestions. I am also grateful to Leslie Whone for preparation of tables, and to Siri Falck-Pedersen Boreske, Laszlo Meszoly, and Charles Chamberlain for illustrations. This study was supported by grants from Sigma Xi, Marsh Fund, and the Albion Foundation. ABBREVIATIONS AMNH — American Museum of Natural History, New York, New York. ANSl^ — Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania. BMNH — British Museum (Natural History), London, England. CM — Carnegie Museum, Pittsburgh, Pennsylvania. F: AM — Frick- American Museum Collection, New York, New York. FHKSM— Fort Hays Kansas State Museum, Hays, Kansas. FMNH — Field Museum of Natural History, Chicago, Illinois. FSM — Florida State Museum, Gainesville, Florida. MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. MNHN — Museum National d'Histoire Naturelle, Paris, France. NMC — National Museum of Canada, Ottawa, Canada. PU — Museum of Natural History, Princeton University, Princeton, New Jersey. ROM — Royal Ontario Museum, Toronto, Canada. SMUSMP— Shuler Museum of Paleontol- ogy, Southern Methodist University, Dallas, Texas. UA — University of Alberta Museum, Edmonton, Canada. UCMP — Museum of Paleontology, University of California, Berkeley, California. UMM — West Texas Museum, University of Texas, El Paso, Texas. UMMP — l^ni\ersity of Michigan Museum ot Paleontology, Ann Arbor. Michigan. UMMZ — University of Michigan Museum of Zoology, Ann Arbor, Michigan. USNM— National Museum of Natmal History, Wa.shington, D.C. YPM — I'eabody Museum of Natural History. Yale University, New Haven, Connecticut. AMI A CALVA LINNAEUS, 1766 Amid calvii is the only extant species of the family Amiidae. It is a predaceous fish that exclusively inhabits fresh waters of the eastern LTnited States. Except for the gar, Lepisosteus, Amia calva is the onl\' other living representative of the holo.stean fishes. Its common name, "bowfin," refers to the long dorsal fin that arches in a bow over most of the length of the fish's back. Amia calva has previously been known as the dogfish, marshfish, mudfish, grindle, or lawyer. The osteology of Amia calva has been extensively described and discussed by Schufeldt (18S5), Bridge (1S77), Allis (1889, 1897), and Goodrich (1930). The following discussion is limited only to the nomenclatural problems, ecology, geo- graphic distribution, and character-states of Amia calva that are relevant to study of the fossil forms. Nomenclature Jordan and Evermann (1896) noted that although Linnaeus (1766) had applied the binomial name Amia calva to the genus, Gronow (1763) had earlier used Amia as a nonbinomial name for fishes presently classified as Apoiion Lacepede. They fur- ther suggested that should Gronow's earlier ipplication of the name be given prece- dence and transferred to Apof^on, then Ainiatus Rafinesque (1815) should replace Amia Linnaeus. Jordon (1906) stated that this transfer of names was a necessary com- pliance with the rules of nomenclature, but later (1919), although citing Opinion 20 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 ( 1910 ) of the International Commission on Zoological Nomenclature which favored Gronovv's priority, Jordan found tlie trans- fer of names inconvenient, for most autliors had rejected Gronow's names as pre- Linnaean. In 1925, Jordan recommended to the Commission that certain names of Gronow supported by Opinion 20 be re- jected in favor of the more accepted Linnaean terminology. The Commission's Opinion 89 ( 1925 ) resolved ( among others ) the nomenclatural problem of Amia, by con- curring with Jordan's recommendation that ". . . Amia Gronow be set aside in favor of Amia Linnaeus, even if other names of Gronow are allowed." Rafinesque's name Amiatus is then a junior synonym of Aynia Linnaeus. Some later workers seem to have been unaware of Opinion 89. Thus Hussakof ( 1932 ) accepted the validity of the transfer of the name A7nia Gronow to the percoid teleost Apo^on. Romer and Fryxell (1928) named their fossil amiid from the Eocene Green River Formation Paramiatus instead of Paramia, and Whitley ( 1954 ) changed the name of Lehman's (1951) fossil amiid from the Eocene of Spitzbergen from Pseu- damia to Pseudamiatus. The latter is invalid as Pseudamia was a valid name in itself and Pseudamiatus is its junior synonym regard- less of the Amia- Amiatus controversy. Ecology Aside from notes regarding breeding, diet, and zoogeographical occurrences, little has been written in the past 50 years about the ecology of Amia calva. Dean (1898) and Reighard ( 1903 ) have made the only ex- tensive published investigations of the habits and habitat of the fish. A thorough study of the biology of A. calva throughout its range is long overdue. Geographic Distribution The distributional map of Amia calva ( Fig. 1 ) is based on information drawn from Hubbs and Lagler (1967), and Blair et al. ( 1968 ) , and from examinations of unpublished records at the Ohio State Uni- versity Museum of Zoology, Museum of Comparative Zoology, and the University of Michigan Museum of Zoology. The dis- tribution limit is a flexible boundary allow- ing for seasonal occurrences and other natural variations. The known northern limit of A. calva extends from the Missis- sippi drainage system in Minnesota south of Duluth, eastward through Lake Nipissing and the Ottawa River to the St. Lawrence- Champlain basin ( encompassing Quebec as far north as Quebec City, and Vermont). A. calva is distributed throughout the Great Lakes region, but is not found in the Lake Superior drainage basin, except in its outlet, the St. Mary's River. Southward, it has been recorded from the Hudson River to western Connecticut ( recorded as the result of introduction; Hubbs and Lagler, 1967); Harrisburg, Pennsylvania, to the Susque- hanna River; and along the Atlantic slope to the Carolinas and Florida. Westward, A. calva occurs along the Gulf Coast to southern Texas as far south as Brownsville, and northward, through eastern Texas, southeastern Oklahoma, northwestern Ar- kansas, eastern Missouri, and approximately 50 miles west of the Mississippi River to Brainard, Minnesota. Pleistocene Occurrences Amia calva has been reported from only three Pleistocene localities: (1) Chicago, Illinois, (2) Vero Beach, Florida, and (3) Itchtucknee River deposits, Columbia County, Florida (MCZ 9524, 9529, 9542). Hay (1911: 552) reported "Professor Frank Baker (Chicago Academy of Science) has shown me a considerable part of the skele- ton and scales of a bowfin which he found in the Pleistocene clay near Chicago." A thorough search of the Chicago Academy of Science collecrions failed to produce this specimen. Hay (1917, 1923) listed Amia calva among the fossil vertebrate remains found in the Pleistocene sands at Vero Beach, Florida. Swift ( 1968 ) , in his review of fossil fishes of Florida, figured Hay's Aiiiia specimens (left dentary and a gular plate; FSM collections) and concluded that Fossil Amiids • Borcskc / /«°«'m;;; L.._ ; , /■ \ r — ■'• ..—■•J^ ! \ : NORTH DAKOTA ; ''w , L . I \ I \minnesota '""^GO- ^^ > • to ' ■» ^^^^^^■>.-,. / /o* ^1 ! I liJUH—^ m o fe'" oo ^— i / / 'l-^-.- i '^ V / ° /'^OlORAdS"— ^ — 1 •(__ \ / /■ j. VMlSbl) I \ /' ; ; KANSAS ' ' "^ \ / J ! \ \ L .' ^ I o , \ .j^^'^oZ. f. ! I \ I ; "^w MEx-6 T-«- -I V; • Q *• .OKLAHOMA L T / -TEXAS ^ 7; > /• I ^ .. ^ / i '. Fossil forms of "••v^ ' ! . © /i3/77/(7 Sp. 'N • A.fragosa \ s'"~-. ° A.uintaensis ''■^ '\ a A.c\.uintaensis \ ' A. scuta to *• » /J.cf. scuta fa \ ■ Amiidae incertae sedis X Pleistocene location of A.calva Fig. 1. Distribution of km\a calva. Fossil occurrences of Amia spp. ore represented by symbols explained in the legend. A. calva was probably very common in the Pleistocene fresh waters of the United States. The pancity of Pleistocene material does not necessarily mean the fish was not common in the Pleistocene, but does indi- cate that A77ua remains have not been extensively collected or identified in exist- iii'j; museum Pleistocene collections. Diagnosis Vertebral meristics similar to A. sctitata, but head/standard-length proportion (0.271 mean) is smaller than in the fossil forms. Extrascapular strap-shaped and relatively wide at midlitie, as in A. scutata, but pos- terior edge is curved so that it is proximally convex, then concave toward the distal corner, which results in a posterolateral projection. Pterotic borders frontal pos- teriorly rather than laterally; anterior end is as wide as posterior end. Orbital excava- tion is shallower than in other species, with a mean depth-to-length ratio of 0.100. In- fraorbital 4 is smaller than infraorbital 5, less robust than in fossil Amia. Preopercu- lum as wide dorsally as ventrally. Symphy- seal incurx'ing of dentary relatively less than ill A. fraii^osa, ])ut greater than in A. scutata and A. uintacnsis: little or no overlapping of dorsal coronoid articulation surface on ventral siuface of ramus; deep Meckelian groov^e. Ventropostcrior process of cleith- rum less sculptured than in other species of Amia. V^omerine teeth shaip, conical, num- bering between 15-27, more anteriorly placed than in A. uintacnsis or A. fraii.osa. Bones less ossified than in fossil Amia. Greatest known standard-length 650 mm. MORPHOMETRICS Comparison of morphometric and meris- tic data of Recent and fossil Amia has facilitated an e\'aluation of the taxonomy as well as clarified anatomical trends. Many 6 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 generic or specific character-states for Amia ''dicfyocepJmla,'^ Amia scutata, "Paramiatus gurleiji,^' and Amia fragosa have been pre- viously estabhslied on osteological data based primarily on gross anatomical propor- tions (head/standard-lcngth ratio and posi- tions of insertion of pelvic and anal fins/ standard-length ratios) and skull propor- tions (parietal/frontal and operculum width/length ratios). Meristic character- states have also been used for A. "dictijo- cephala" and A. scutata. Altliough an age-growth analysis on Amia calva was done by Cartier and Magnin (1967), no moiphometric investigation of a growth-series of Recent A. colva has yet been completed or used for comparison with fossil forms. Estes and Berberian (1969: 10) suggest that knowledge of the growth-series of A. calva would be of con- siderable importance in tracing the ancestry of the modern species. Hammett and Hammett (1939) made a moi-phometric study of the Recent Lepisos- teus platijrhinciis, taking length dimensions of a sample of live fish from Florida. Since Lepisosteus, like Amia, is one of the two extant holosteans, their analysis is poten- tially useful in providing information on the ancient species. However, they did not actually compare the live material or data with any fossil material. According to Imbrie (1956), Simpson et al. (1960), and Gould (1966), growth studies offer excellent means with which to clarify evolutionary and taxonomic prob- lems in the fossil record. An interesting model utilizing morphometric data for synonymy of fossil forms was made by Thomson and Hahn ( 1968 ) on the growth- series patterns of Devonian rhipidistian fishes, in which they showed that Thiirsius clappi was actually a juvenile form of Eusthenopteron foordi. In studying fossil material, as Thomson and Hahn (196S: 201) indicate, there is a problem in deter- mining the age, sexual matmity, and envi- ronmental regime of the animal. Also, of course, it is necessary to have sufficient fos- sil material with which to erect an adequate growth-series. This present analysis is undertaken (1) to determine w hether skull and axial skele- tal proportions of amiids are isometric or allometric with increasing size, (2) to establish the variation in meristic charac- ters of Recent A. calva, and ( 3 ) to compare moiphometrics and meristics of Recent A. calva with those of the fossil forms. This study utilizes a small sample of 18 Recent A. calva specimens from the St. Croix River, Wisconsin. Measurements were taken from a growth series that includes the size range of most of the articulated fossil forms. The largest A. calva specimen, from St. Joseph County, Michigan (UMMZ 197683), was analyzed to see whether the large specimen would agree with the anatomical propor- tions and meristic characters of the Wiscon- sin specimens. Three smaller specimens from Pewaukee, Wisconsin (MCZ 8970), were also included. The fossil sample con- tains six complete and ten partially com- plete amiid specimens ranging in age from Late Cretaceous to Late Miocene which, although moiphometrically similar in vary- ing degree, are too few to warrant conclu- sions in themselves. Methods Measurements chosen for this study ( Fig. 2) are those of Hubbs and Lagler (1967: 20). In fossil forms, because of the lack of preservation of internal soft anatomy as well as the impossibility of determining their interbreeding potential, these particular measurements necessarily assume an in- creased taxonomic significance, since they often provide the only viable parameters with which to designate genera and species. Measurements taken on A. calva are limited to those also represented in the fossil speci- mens. Each of the A. calva measured was X-rayed, except for three small specimens, which were cleared and stained. The range of error for aj] measmements taken on Re- cent and fossil material is ±0.04 mm. The range of error for the ratios is ±0.08 mm; Fossil Aaiiids • Boreske Fig. 2. Index to the measurements used, superimposed upon an outline drawing of Amia. Key for body measurements: TL = Total-Length SL =z Standard-Length H zi: Head-Length C ^ Caudal-Length Pf =3 Insertion of Pelvic Fin P = Insertion of Anal Fin HL =: Standard-Length minus Head-Length ML := Standard-Length minus Mandible-Length Key for abbreviations of cranial elements used in morphometric study: M = Mandible G Gular |5 Infraorbital F = Frontal Par = Parietal O zi: Operculum Table 1. Length dimensions of 22 specimens of Amia calva L.: 21 from Wisconsin (MCZ 8970'), 1 from Michigan ( UMMZ 197683)'* Measurements in mm Specimen Class Range Code Total Length No. TL SL ML H HL Pf P c 1* 80.0 1 80.0 70.5 57.0 22.0 48.5 32.5 12.5 10.5 2* 95.0-105.0 2 100.0 85.0 70.0 25.0 60.0 39.5 16.0 15.0 3 207.0-212.0 6 210.0 175.0 145.0 50.5 124.5 80.9 35.0 35.0 4 227.9-232.0 4 230.0 193.0 161.0 54.6 138.4 88.5 35.5 36.0 5 241.0 241.0 199.0 165.9 56.8 142.2 93.5 38.0 42.0 6 291.0 291.0 237.0 197.0 64.0 173.0 115.0 52.5 54.0 7 310.0 310.0 248.0 207.0 68.5 179.5 112.0 46.0 62.0 8 339.0 339.0 274.0 230.0 73.0 202.0 125.0 51.0 64.0 9 385.0 385.0 313.0 259.0 82.0 231.0 142.0 52.0 72.0 10 433.0 4.33.0 349.0 293.0 91.0 258.0 170.0 71.0 84.0 11 475.0 475.0 .399.0 335.0 103.0 296.0 181.0 82.0 76.0 12 507.0 507.0 423.0 359.5 109.0 317.0 192.0 93.0 81.0 13»» 756.0 756.0 648.0 545.3 164.0 480.0 299.0 138.0 102.0 See Figure 2 for abbreviations. 8 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 this margin of error is graphically inconse- quential in this study. Specimens of A. calva whose total length was between 207 mm and 507 mm were selected because this range of A. calva would provide the best information for comparison with the fossil species. Twenty-two specimens of A. calva were measured (Table 1). Eighteen of these are from the St. Croix River, Wiscon- sin. These 18 specimens of A. calva fall into ten categories arranged here by ap- proximately 20-30-mm class range incre- ments in total-length. Although these categories represent arbitrary rather than biological growth stages, they provide suflBcient information on the morphologic size changes of A. calva. Three smaller specimens (MCZ 8970, also from Wiscon- sin) witli a size range of 80-105 mm total length (TL) were included to de- termine whether they would follow the predicted allometric effect on the growth- series Hne, since, as Thomson and Hahn (1968: 205) note, it is a common feature for the early stages of juvenile animals to have heads proportionately larger than the bodies. Hay (1895) notes that an 80-mm A. calva is beyond the embryonic stage and is an early juvenile with most of its bones at least partially ossified. The 80-mm speci- men has a proportionately larger head to standard-length ratio than the other mem- bers of the growth-series (Table 3). Al- though this ratio decreases slightly with increasing size, the head /standard-length ratio of 0.312 for the 80-mm specimen does not deviate far from the growth-series line (Figs. 3-4). The largest specimen (UMMZ 197683) was used as a size limit for the other end of the growth-series continuum. It may be assumed that this fish had already reached the size or point of maturity at which fish normally begin to decrease their rate of growth. This specimen still retains the morphological proportions of the smaller specimens (Figs. 3-4) and, like them, falls remarkably close to the constant relative size-growth lines of the various proportions. Although from Michigan, this specimen does not appear to deviate from the growth- series line established by the Wisconsin specimens of A. calva. The Michigan speci- men of A. calva, since it agrees with the growth-series continuum established by the Wisconsin specimens, is helpful in extend- ing comparison to the larger fossil amiids: "Paratniatus gurleiji" (FMNH 2201), Amia fra^osa (MCZ 5341), and Amia uintaensis (PU 13865), which are outside the size range of the Wisconsin sample. General Proportions and Growth Allometric growth, according to Gould (1966: 595), describes geometrically pro- gressive change in shape or proportions with size, and is generally represented by a curvilinear line or, in certain cases, by a straight line in which the Y-intercept is significantly different from 0. For the Amia calva growth series dis- cussed here, the ordered pairs correspond- ing to the proportions in each series have been plotted on a graph, as well as the straight line corresponding to the equation y = a + bx (of the best fit computed ac- Table 2. Length DrMENSIONS ; OF 6 ARTICULATED FOSSIL AMIIDS Measurements in mm TL SL ML H HL Pf P C A. scutata PU 10172 '■«yo4.o 339.0 276.5 106.0 233.0 159.0 73.0 e«t65.0 A. scutata UMMP V-57431 — 388.0 313.8 121.0 267.0 183.0 83.0 A. kehreri BMNH P33480 249.0 191.0 160.8 59.2 131.8 89.0 38.5 58.0 "Paramiatus gurleiji" FMNH 2201 702.0 510.0 430.0 157.0 353.0 <'«f265.0 78.0 192.0 A. fragosa MCZ 5341 575.0 455.0 383.0 142.0 313.0 210.0 75.0 115.0 A. uintaensis FV 13865 848.0 664.0 — 214.0 450.0 288.0 116.0 160.0 See Figure 2 for abbreviations. Fossil Amhds • Boreske 9 320- 280- 240- 200- 160- 120- 80- 40- I 40 —I 1 I I I 120 200 280 STANDARD I I I 360 LENGTH mm 680 440 520 600 Fig. 3. Relative growth-lines of head-length (H), pelvic fin insertion (Pf), and ana! fin insertion (P) plotted arith- metically against standard-length, for 18 specimens of Recent Ami'o calva (A = MCZ 8970 and H = UMMZ 197683 are included for comparison). cording to the method of least squares); whicli nearly passes through the origin of the results of these calculations appear in the graph. The coefficient of correlation is Figures 3-4. Practically all the ratios in almost equal to 1.0 in each case, an indica- Figure 3 fall onto straight lines, each of tion that the computed straight line provides 10 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 Table 3. Comparisox of length proportions in 22 specimens of Amio calva with fossil amiids Specimen Code H/SL Pf/SL P/SL 1 0.312 0.461 0.177 2 0.294 0.464 0.188 3 0.289 0.462 0.200 4 0.283 0.459 0.184 5 0.285 0.470 0.191 6 0.270 0.485 0.222 7 0.276 0.452 0.185 8 0.266 0.456 0.186 9 0.262 0.455 0.198 10 0.261 0.487 0.203 11 0.258 0.454 0.206 12 0.258 0.454 0.220 13 0.259 0.461 0.213 0.258-0.289'' 0.452-0.487* 0.184-0.222* mean = (0.271)* mean = (0.463)' mean = (0.199)* Oligocene A. scutata PU 10172 0.313 0.469 0.215 A. scutata UMMP V-57431 0.312 0.472 0.214 Eocene A. kehreri BMNH P33480 "Paramiatus gurleyi" FMNH 2201 A. fragosa MCZ 5341 A. uintaensis PU 13865 0.310 0.466 0.201 0.308 *'sto.520 0.153 0.312 0.462 0.165 0.322 0.434 0.175 Range and mean exclude Specimen Codes 1 & 2 (MCZ 8970) and 13 (UMMZ 197683; a very good fit for the ratio series, and that the relative growth of these three dimen- sions is essentially isometric rather than allometric. The Wisconsin specimens (in- cluding the 80-105-mm specimens ) and the larger Michigan specimen all fall close to the line calculated for each of the three ratios (Fig. 3). The proportions of head- length/standard-length, insertion of pelvic fins/standard-length, and insertion of anal fins /standard-length are shown in Table 3. The head/standard-length ratio shows a slight decrease with increasing size, but this ratio series nonetheless has a very high coefficient of correlation for the strength of the linear relationship (Fig. 4). The lengths of the mandible, parietal, frontal, and operculum in Recent A. calva appear in Table 4, and the proportional ratios in Table 6. The relative growth rate of each of these proportions is constant with X and Y-intercepts of the straight line close to the origin. The coefficient of correlation for the variables in each of the proportions is 0.997, 0.975, and 0.997, respectively ( Fig. 5). Combined, these two factors indicate constant and therefore isometric relative size-growth of the compared skull element. Comparisons with Fossil Forms Six articulated fossil specimens were available for measurement and calculation of head /standard-length and positions of insertion of pelvic and anal fins/ standard- length (Tables 2-3). The measurements taken from the fossil forms are as exact as conditions allow, although it must be stressed that varying degrees of crushing and distortion have occurred in fossilization, and evaluation of the morphometries should be qualified with this in mind. Head /standard-length ratios (Fig. 4). The fossil forms all show a slightly greater head/ standard-length ratio than does the Recent species (Table 3; Fig. 4). A. uintaensis (PU 13865) is the largest of Fossil Amiids • Boreske 11 Table 4. Length dimensions of Mandible (M), Gular (C), Frontal (F), Parietal (Par), Infraorbital ^(I''), and Operculum (O) in 22 specimens of A. calva Measurements in mm Operci ilum Specimen Dors. -Vent. Ant.-Post. Code M G F Par 18 (OL) (OD) 1* 13.5 8.0 10.0 5.0 5.0 8.1 7.5 2* 14.9 9.4 11.4 6.1 5.8 9.0 8.4 3 30.0 19.0 18.0 9.7 12.0 14.0 12.9 4 32.0 21.0 20.0 9.9 13.5 15.0 14.2 5 33.5 20.5 19.0 10.0 14.2 16.0 14.9 6 39.0 26.0 25.0 11.0 17.0 16.9 16.5 7 42.0 28.0 25.5 11.5 18.0 18.1 17.5 8 45.0 27.0 26.2 13.5 21.0 19.8 18.5 9 53.0 31.0 31.0 14.7 22.5 22.8 21.8 10 56.0 32.5 32.2 17.0 26.5 22.6 22.5 11 63.0 39.0 38.4 18.5 30.5 28.1 27.8 12 66.5 42.5 39.0 19.5 31.5 27.8 28.5 13** 102.7 — 60.7 30.0 — — o MCZ 8970. «<» UMMZ 197683. all the fossil specimens, but nonetheless has a greater head/ standard-length ratio than any of the others. The head of this form is so much more elongated than the head in A. fra^oso (MCZ 5341), A. kehreri (BMNH P33480), and "Paramiatus gurleiji" (FMNH 2201) that it offsets the fact that its vertebral column includes approximately 20 more vertebrae than do these three forms (Table 9). Thus, although A. idntaensis Table 5. Length dimensions of Head (H), Mandiiu^ (M), Gular (G), Frontal (F), Parietal (Par), Infraorbital ^'{V'), and Operculum (O) in fossil amiids Measurements in mm Operculum Dors.-Vent. Ant.-Post. H M G F Par F (OL) (OD) A. cf. scutata UCMP 38222 — 65.2 — 46.0 23.0 35.0 — — A. scutata PU 10172 106.0 62.5 31.2 35.0 16.0 — 29.0 28.0 A. scutata UMMP V-57431 121.0 74.2 44.3 20.0 29.1 27.9** A. "dictt/ocephala" AMNfl 2802 111.5 68.0 38.0 17.0 29.1 32.0 30.0 A. kchrcri BMNH P33480 59.2 30.2 — 20.0 8.4 15.3 20.5 19.0 "Paramiatus fiurletji" FMNH 2201 157.0 80.0 — 58.0 23.6 25.0 40.0 37.0 A. fra^osa MCZ 5341 142.0 72.0 68.7 56.0 22.8 39.0 36.2 A. fragosa MCZ 9264 80.0 40.0 — 26.0 10.5 18.5 — — A. uintaensis PU 13865 214.0 — — . 88.0 34.0 — 55.0 51.0 "Protamia" mongoliensis AMNH 6372 — — 81.0 — — — 54.0 52.0 A. uintaensis PU 16236 315.0** 220.0 158.0 160.0 60.0 — — 95.0 A. fragosa MCZ 9291 — ■ — — — — — 27.0 25.0 A. fragosa AMNH 9315 — — — — 29.0** 27.0 A. fragosa UA 5450* — — 26.0 10.0 — — — A. fragosa UA 5458* — — 30.0 12.0 — — A. fragosa UA 5480* — — — 20.0 26.0 24.0 •Data from O'Brien (1969). •• Est. 12 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 240- 220- 200- EQUATiONof STRAIGHT LINE- M= 7.268 + (0,234) (SLl u y leo- COEFFICIENT ol CORRELATION = 0.999 ISO- MEAN ^ -. 0.271 STANDARD DEVIATION 5^ = 0.0117 COEFFICIENT of VARIATION =4.33% E E 140. X ; 120- a S 100- I ^^ 60- xX' SO' /^ 40' /^ 20- ^x 0. 320 400 460 STANDARD LENGTH mm y EQUATION of STRAIGHT LINE - PF = 1 438 + (0.454)(SL) ■^ 288 ' 240- COEFFICIENT of CORRELATION .0.995 MEAN f^ '- 0.463 STANDARD DEVIATION |^ -■ 00131 COEFFICIENT of VARIATION ■ 2 83% y '" 0 y y y y y y 192- y 144. 96- y 48. X 0. X 240 J20 400 STANDARD LENGTH mm E E ■z- 120' EQUATION of STRAIGHT LINE ■■ P = -5.171 +(O.E27)(SL) COEFFICIENT of CORRELATION = 0.987 MEAN ^ = 0.199 STANDARD DEVIATION^ .0.0135 COEFFICIENT ot VARIATION '6.78 % 240 320 400 STANDARD LENGTH mm Fossil Ami ids • Boreske 13 has significantly more vertebrae than these other forms, this feature is not reflected in a comparison of head/ standard-length ra- tios ( Table 3 ) . This is also true, to a lesser extent, in both A. scutata specimens (PU 10172, UMMP V-57431) from the Ohgocene Florissant Formation; these specimens fall into the head/ standard-length range of the three fore-mentioned forms, but like A. uintaensis, possess vertebral columns having nearly the same number of centra as those in A. calva. Thus in themselves the head/ standard-length ratios are of little help in comparing the fossil forms, but when coupled with the corresponding lengths of the vertebral column (based on number of centra) they are informative. A. uintaen- sis (PU 13865) and A. scutata (PU 10172, UMMP V-57431) have relatively elongated heads; A. kehreri (BMNH P33480), A. fragosa (MCZ 5341), and "Paramiatiis g,ur- leyi" (FMNH 2201) have relatively shorter heads, since the head/standard-length ratio is less than might otherwise be expected considering the smaller total-number of centra (only two-thirds the number of centra of A. uintaensis, A. scutata, and A. calva ) . A tentative growth-line ( also calcu- lated by the best-fit method ) was included for A. jragosa on the basis of three speci- mens (Fig. 4). In comparison with the growth-line of the Recent species (0.271 mean), it reflects the larger head/ standard- length ratio of A. fragosa (0.310 mean). The growth-line computed for A. jragosa is linear and falls near the origin, indicating that increase in head size/ standard -length was isometric, as in A. calva. Fin relations] lips. In the smaller fossil forms, the ratio of the point of insertion of the pelvic fin/ standard-length shows little deviation from the modern species (Table 3; Fig. 4) except for two Eocene specimens, "Paramiatus gurleyi" (FMNH 2201) and A. uintaensis (PU 13865), which fall out- side of the range on either side of tlie size- growth line. The greater ratio for "Para- miatus gurleyi," however, is probably the result of distortion in its preservation. The length of the pelvic fin insertion/ standard- length does not appear to be a satisfactory taxonomic index, distinguishing neither the fossil forms from one another nor the fossil forms from the Recent A. calva. "Paramiatus gurleyi" (FMNH 2201), A. uintaensis (PU 13865), and A. jragosa (MCZ 5341) have a relatively shorter dimension between the anal fin and the end of the vertebral column than do A. calva, A. scutata, and A. kehreri (Fig. 31). Any attempt to inteipret the fossil data for this ratio is complicated by the fact that con- siderable overlap with the Recent species occurs. Both long-bodied (A. scutata) and short-bodied (A. kehreri) forms fall within the range of A. calva, while other long- bodied (A. uintaensis) and short-bodied (A. jragosa, including "Paramiatus gur- leyi") fonns fall below the range of the Recent species (Table 3). Although the ratio of anal fin/ standard-length may pos- sibly be useful in distinguishing A. jragosa (including "Paramiatus gurleyi") from A. calva, A. scutata, and A. kehreri, it is not useful in distinguishing either of the two fossil fonns from one another or from A. calva. The smaller dimension indicated by the low ratios (0.153, 0.165) of A. jragosa is doubtless a reflection of its shorter axial column. The relatively small (0.175) ratio for A. uintaensis is probably in part the result of its longer head, wliich increases its standard-length in relation to the other forms; at any rate, the difference between the A. uintaensis ratio and the range for Recent A. calva is not very sig- nificant. Mandible I head ratios. A comparison of Fig. 4. Relative growth-lines (broken-solid lines) of head-length, pelvic fin insertion, and anal fin insertion plotted arithmetically against standard-length for Recent Amia calva (A = MCZ 8970 and ■ = U/AMZ 197683 are included for comparison) with compared fossil forms: fl = A. hagosa (A. kehreri) BMNH P33480; f- = A. fragosa (Pararr^iafus gurleyi) FMNH 2201; f'-^ = A. fragosa MCZ 5341; s^ = A. scufafa PU 10172; s^ = A. scutofo UMMP V-57431; u = A. uintaensis PU 13865. The broken-dotted line is the "best fit" line for available specimens of A. fragosa. 14 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 no 100 90^ tOuATiON o( STRAIGHT LINE ■ M ■-! 010 + (0-636HM) COEFFICIENT of CORRELATION ^ 0. 997 MEAN ^ ' 0 609 STANDARD DEVIATION ^ ■ 0 0171 COEFFICIENT of VARIATION -■ 2.81 % 55- 50. / 45- EQUATION 0) STRAIGHT LINE ^ PAR = 0 278 + (0 479HF) '^ y COEFFICIENT ol CORRELATION • 0.975 /^ ^ 40- E MEAN ^ . 0.495 / /"^ E ± 35- STANDARD deviation""- 0 0333 ^^ / COEFFICIENT of VARIATION- 6 73% / / "^ V O / / X UJ 30- f' y ~t / / < / / UJ 25- / < V A S / /•' 20- 1!- 10. /^' 5. 0. / 30 40 50 60 SO 90 rOO 110 120 FRONTAL LENGTH mm o-»r EQUATION of STRAIGHT LINE : OD ■ - 1.559 + (l.079)(&LI COEFFICIENT of CORRELATION - 0.997 MEAN g2 -0.964 STANDARD DEVIATION ^ - 0.0326 COEFFICIENT of VARIATION- 3.40% 20 30 40 OPERCULUIM LENGTH mni Fossil Amiids • Boreske 15 Table 6. Cranial proportions in 22 specimens OF A. calva Specimen Code M/H Par/F OD/OL 1 0.614 0.500 0.926 2 0.596 0.535 0.933 3 0.594 0.539 0.921 4 0.586 0.495 0.947 5 0.590 0.526 0.931 6 0.609 0.440 0.976 7 0.613 0.451 0.967 8 0.616 0.515 0.934 9 0.646 0.474 0.956 10 0.615 0.528 0.996 11 0.611 0.482 0.989 12 0.610 0.500 1.025 13 0.626 0.497 0.586- 0.440- 0.921- 0.646* 0.539" 1.025" mean mean mean = (0.609)" = (0.495)" = (0.964)" ■* Range and mean exclude Sjiecimen Codes 1 & 2 (MCZ 8970) and 13 (UMMZ 197683). the mandible/head ratios of Recent A. calva with those of the fossil forms ( Table 7; Fig. 5) indicates that the A. scutata and A. ''dictyocephala" (AMNH 2802) ratios are very close to those of A. calva. The A. fra^osa .specimens (including "Taramiatus ^urleijr FMNH 2201 and A. kehreri BMNH P33480) have a mean mandible/ head ratio of 0.507, which, when compared to the A. calva mean ratio of 0.609, indicates a relatively smaller mandible to head size (Table 7). Unfortunately, A. uintaensis (PU 13865) cannot be used in this com- parison, since the mandibles are buried in matrix. A reconstruction of a disarticulated A. uintaensis (PU 16236) specimen from the Late Pal eocene has been made, and its ratio is approximately 0.693. Thus man- dible/head proportions may be valid for distinguishing specimens of A. fra^usa and A. uintaensis from one another as well as from A. calva and A. scutata. This ratio, however, caimot be used as a valid criterion for distinguishing A. scutata from A. calva. The 0.693 mandible/ head ratio of A. uintaeims indicates that this form has the largest mouth gape of the four valid species. A tentative growth-line for the mandible/ head-length proportion of A. fra additional Ohgocene specimen and compared them with my sample of A. calva, which showed between seven and eight pehic lepido- trichia (Table 8). Although Osborn et al. ( 1878) counted ten pelvic lepidotrichia, my recount of their A. scutata specimen (PU 10172) showed only seven (Plate 4). The bifurcation of the fin rays might have been inadvertently included in their original count. The holotype of A. "dictyocepliala" (USNM 3992) (Fig. 27) showed seven rather than the six lepidotrichia that Cope (1875) had diagnosed. A specimen of A. scutata (UMMP V-57431) (Fig. 27A) also has seven lepidotrichia; both of these are within the range of Recent A. calva. Of the remaining fossil forms, A. fraii^osa and "Paramiatus ^urleyi" have eight, and A. uintaensis nine, A. uintacnsis being the only fossil form not to fall within the range of Recent A. calva. This difference is insuf- ficient to demonstrate any taxonomic value, however, at least until more A. uintaensis specimens are known. Anal fin. Anal fin lepidotrichia have been included in the diagnoses of A. "dic- tyocephala" and A. scutata (Cope, 1875), and also in the description of A. .scutata (Osborn et al, 1878). Each of the original counts of nine anal rays for each specimen concurs with my recount and also falls within the range of eight to eleven for Re- cent A. calva (Table 8). A. jra 11 ^n ^li ^5 2r~ Caudal Fin Lepidotrichia no. TT Fig. 9. Number of caudal lepidotrichia in 20 speci- mens of Recent Ami'a calva. arrive at an accurate count of the total number of hypurals or to verify whether this one-to-tvvo relationship exists in all the amiid fossil forms. The only available fossil form in which this one-to-two hypural- lepidotrichia coiTCspondence in the ventral caudal region can clearly be seen is in A. scutata (YPM 6241; Fig. 8). Vertebral Elements Two regions of the vertebral column, the trunk and the caudal regions, are defined by their relationships to the ribs, neural arches, and haemal arches. The trunk region con- sists of monospondylous vertebrae that pos- sess paired basapophyses having gradually changing angles, dorsal neural facets, and ventral aortal facets. The number of tnmk vertebrae in my sample of Ainia calva varies from 36 to 38. The caudal region consists of three types of vertebrae, listed from an- terior to posterior: regular monospondylous centra bearing neural and haemal arches, diplospondylous centra bearing neither neural nor haemal arches (neural and haemal facets still present ) , and ural centra. Since the neural and haemal facets are still present in the diplospondylous centra, there is no way to differentiate the latter from the monospondylous type in a disarticulated state. In my sample of A. calva, the number of regular caudal monospondylous centra (24-26) fluctuates by two centra, that of the diplospondylous caudal centra ( 14-17 ) by three (Table 9). The posterior caudal region of A. calva consists of two types of urals: centra with hypurals attached by a layer of cartilage (free urals), and centra that are fused directly onto the hypurals, often lacking the neural arches (fused urals). When dis- articulated, the fused urals can often be distinguished from the free urals, since part of the hypural usually remains fused to the ural, extending the posterior articular surface downward. The nonfused (free) urals cannot be distinguished in a disarticu- lated state from the monospondylous or diplospondylous caudal centra. The num- ber of urals with fused hypurals is readily counted, since they are distinguishable from the remainder of the vertebrae. In order to identify a free ural, it is necessary to observe the relationship between the ural and its conesponding hypural and lepidotrichia. It is often difficult to make this distinction between free and fused urals, since the caudal region is seldom complete in articulated fossil forms. In A. calva the number of urals with ankylosed hypurals ranges between seven and nine. There are approximately seven principal urals fused to hypurals, followed by one or two small additional urals that do not ar- ticulate with the preceding vertebrae but lie dorsal to the upturned portion of the vertebral column. Because it is difficult to discern these urals in smaller specimens of A. calva, the count may be slightly biased, and a comparison of the fossil forms with the range established for A. calva must be made with this consideration in mind. I counted the number of centra between the anterior dorsal fin pterygiophore and the posterior anal fin pterygiophore, since Cope (1875) used the number of central elements between these points as a specific character for A. "dictyocephala" (USNM 26 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 Fig. 10. Am'ia calva (648 mm SL) caudal: A, Whitehouse (1910) and Lund's (1967) definition of first ural; B, Nybelin's (1963) definition of first ural; C, first fused ural. 3992 ) . The range for the number of centra in this region of Recent A. calva is 33 to 37. There is considerable variation in total number of centi'a {i.e., segments) in Recent A. calva (81-90), which may pose a prob- lem in comparing specific vertebrae. Thus in two A. calva, for example, the eightieth vertebral segment of one individual might not correspond to the same position in the vertebral column or even type of centrum as the eightieth segment of the second indi- vidual. This should be considered in any comparisons of several A. calva individuals, as well as in comparisons of the fossil forms, which share this variation in vertebral seg- ments (Table 9). Also, fusion of vertebral elements may occur in Recent A. calva. In some specimens, as many as five centra were found fused together at points throughout the vertebral column; this con- dition was present to a lesser degree or absent in other specimens (Tables 10-12). These fused centra also occur in the fossil forms, as in A. uintaensis (YPM 6244). The actual number of such fused centra can often be established only by counting ex- ternal features such as basapophyses, neural facets, aortal facets, or haemal facets. Romer and Fryxell's ( 1928 ) study of "Paramiatus gurleyi" is the only pubHshed description of a complete articulated fossil amiid. They distinguished this form from the Recent species by the supposed pres- ence of a deeper body, and also noted that the number of centra was considerably less than in A. calva. The vertebral column is completely preserved, so that it is possible to obtain an accurate count of the vertebrae (Plate IB). "Paramiatus p,iirleyi" has 67 vertebral segments in contrast to the mean of 86 in A. calva (Table 9). Osborn et al. (1878) described A. sciitata (PU 10172) on the basis of a specimen lacking a caudal fin (Plate 4). Since the specimen is other- wise complete, they were able to estimate that their specimen had 82 vertebral seg- ments. Cope (1875) described A. "clictyoce- Fossil Amiids • Boreske 27 Table 9. Comparison of VERTEBRAL CHARACTERS IN RECENT AND FOSSIL AMUDS Number of Centra between Anterior Number Number Number Dorsal-Fin of Mono- of Diplo- of Ural Pterygiophore Total Number of spondylous spondylous Centra and Posterior Number of Trunk Caudal Caudal with Fused Anal-Fin Centra"" Centra Centra Centra Hypurals Pterygiophore" Recent Amia calva (20) Wis. & Mich. 81-90 36-38 24-26 14-17 7-9 33-37 mean mean mean mean mean mean = 85.8 = 37.3 = 25.2 = 16.2 = 8.3 = 35.5 Oligocene A. scutata PU 10172 83*** 36 25 15 yooo 35 A. scutata UMMP V-57431 81*** 36 24*** 15*** Y* 00 37 A. "dicttjoccphala" USNM 3992* — — 35 Eocene "Paramiatus ^urlcyi •* FMNH 2201* 67 26 19 16 6 26 Amia uintaensis PU 13865 85 31 26 21 7 36 Amia uintaensis AMNH 785 25 20 7 — A. fragosa MCZ 5341 65 25 18 15 7 25 A. kehreri BMNH P33480 62*** 24 16 16 6*** 24 types. o -- "" ^ including diplospondylous units (as one), "o" Est. pluild" from a specimen (USNM 3992) in which only the mid-body region was pre- served. He felt that the number of ver- tebrae between the anterior dorsal fin pterygiophore and the posterior anal fin pterygiophore had ta.xonomic significance. A comparison of this specimen with Recent A. calva showed that the v'ertebral count of this region is essentially the same in both species. This character is therefore not useful in distinguishing this species from the Recent form or in characterizing it as a specific taxon. The specimens of A. scutata are within the range of A. calva in total number of vertebrae as well as in the number of vertebrae in the vimous cate- gories (Table 9) . Based on the similarity of number of vertebrae in A. scutata to that of A. calva, it appears that the amiid vertebral column has not changed meristically from Oligocene to Recent. Additional data from five undescribed fossil amiid specimens with relatively com- plete axial skeletons has been of consider- able help in estimating vertebral counts of the fossil forms. A complete specimen of A. uintaensis from the Green River For- mation (PU 13865) has a complete axial skeleton (Plate 3). Interestingly, the total number of centra (85) does not differ from that of A. scutata or A. calva (Table 9). The only variation is in the number of trunk centra and the number of diplospon- dylous caudal centra. There are fewer trunk centra in this specimen of A. uintaen- sis (31) than in A. scutata, which has a mean of 36, or in A. calva, \\'hose trunk centra are a mean of 37. A partially com- 28 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 plete A. uintaensis specimen (AMNH 785), also from the Green River Formation, shows almost the same number of diplospondylous caudal centra as PU 13865 (20-21 respec- tively). The lesser number of trunk centra in both specimens of A. uintaensis is thus offset by a greater number of diplospondy- lous caudal centra. In comparing the verte- bral column of A. uintaensis with that of A. calva, A. scutata, and A. fragosa, it ap- pears that although A. uintaensis shares the same total number of vertebral segments with A. scutata and A. calva, it does not conform to their proportional division of the column into trunk and caudal regions. A. uintaensis has a trunk/ total-number ver- tebral ratio of 0.365, while A. fragosa has a ratio of 0.300 as compared to the A. calva ratio of 0.440. Three complete specimens referred to here as A. fragosa ("Paramiatus gurleyi" FMNH 2022, A. kehreri BMNH P33480, and A. fragosa MCZ 5341) have vertebral columns that differ proportion- ately and meristically from A. calva, A. scutata, and A. uintaensis. A. fragosa has significantly fewer centra than the other fossil forms, with approximately 12 fewer trunk vertebrae and 8 fewer monospondy- lous caudal centra. It has approximately the same number of diplospondylous caudal centra as A. calva and A. scutata, with the number of fused hypurals also essentially the same (Table 9). Thus A. fragosa and A. uintaensis are meristically distinct from one another and also from A. scutata and A. calva, suggesting that these two earlier forms can be taxonomically separated on vertebral meristic characters. VERTEBRAL COLUMN OF AMIA CALVA The existing taxonomy of many North American fossil amiids is based primarily on vertebral characters. Many of the spe- cies of "Protamia," and the genus itself as described by Leidy (1873a) from the Bridger Formation, have been established solely on height/ width proportions and length (thickness), shape of the neural and aortal facets, and various foramina of isolated vertebrae. Fossil species of Amia from the Bridger and Cypress Hills forma- tions have also been defined on character- states of isolated vertebrae. In order to analyze this usage, variation in vertebral character-states of A. calva has been studied. The axial skeleton of Recent Amia calva is relatively well known. It is one of the few modem forms that have diplospondylous vertebral centra posteriorly, a condition that, according to Schaelfer (1967), func- tionally increases the flexibility of the pos- terior part of the body. Shufeldt (1885) was one of the first to describe the verte- brae of Amia, and Hay's ( 1895 ) well- known work on the vertebral column of Amia provides a relatively complete and informative description of the axial skele- ton, as well as one of the first discussions of intracolumnar variation of the centra. Hay observed some gradual changes in centrum proportions, and in the position of the neural and aortal facets. Vertebral Features Dorsal and ventral facets, basapophyses, foramina, and ridges on the centra have been used as diagnostic characters in the taxonomy of fossil amiids. There are three types of paired facets on the vertebrae: dorsal neural facets for the neural arches, ventral aortal facets for the aortal supports, and haemal facets for the haemal arches. Neural facets. The neural facets are shallow depressions under the neural arch bases, which in life are filled with cartilage. Cartilage is present between the centrum and its associated neural arch. Some speci- mens of A. calva have much deeper facets, with a small ossified ridge built up on the borders. These neural facets occur in pairs on the dorsal surface of both trunk and caudal vertebrae, and between the two facets lies a groove that partially receives the spinal cord. According to Hay (1895: 7-9), there is a marked anteroposterior change in the posi- tion of the neural facets. He contended that at the anteriormost end of the vertebral column the neural arch bases occur be- tween two vertebrae and rest equally on Fossil Amiids • Boraske 29 VENTRAL dUJZ^ odZlL^ 28 35 36 37 38 4II> DORSAL Fig. 11. Configuration of aortal facets (as) and neural facets (ns) on trunk and anterior caudal vertebrae of Amio calva (339 mm SL). both; going posteriorly the.se bases shift gradually backward. He also observed that there is a change in the spacing of the neural iuches; they are close together in the anterior trinik region and more widely spaced posteriorly. Hay is correct in regard to the change in spacing of the neural arches, but he is not altogether correct in his description of the change in position of these arches in relation to the centra. An examination of the Wisconsin A. calva sample showed that, after the first few an- teriormost centra and corresponding neural arches, the middle of the neural arches is situated at the juncture between the centra. This placement continues along the axial column until the first diplospondylous ver- tebra occurs. At this point, the next five to seven neural arches are found aligned to the middle of each of the corresponding centra, after which the arches appear to move forward slightly and correspond ir- regularly to the vertebral bodies. The configuration of the neural facets themselves varies in the trunk region of the vertebral column of A. calva. The neural arches in the anterior trunk region are thicker and wider than those in the more posterior trunk region which have become more flattened and elongated. The shape of the neural facets reflects this trend ( Fig. 11). After the first two ccMitra, the facets assume an hourglass shape, being narrower in the middle and broader at each end. 30 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 This can be related to the fact that the neural arches are situated at the juncture of two centra so that each neural facet sup- ports the anterior and posterior halves of two different neural arches, whose bases are narrow at the extremity and thick in the center. Although the neural facets in any given specimen of A. calva conform to this general trend, the individual configura- tion of the facets varies slightly. Given this variation in shape of the neural facets, it is useless to attempt characterization of the vertebral column of any amiid species based on configuration of neural facets. Aortal and haemal facets. On the ven- tral side of the trunk vertebrae are two thin cartilaginous projections that are located on either side of the dorsal aortal supports. When the skeleton is dried, these projec- tions leave marked depressions, which, like the neural facets, vary gradually from the first anterior vertebra to the last few trunk vertebrae; at this point the aortal facets coalesce with the basapophyses (Fig. 11). The point where these two elements are completely merged marks the termination of the trunk centra, and the next centrum is that of the first caudal vertebra. These structures, which were derived anteriorly from the basapophyses and aortal facets, here become the haemal facets. The first pair of aortal facets is very small and ovoid. The next few centra bear aortal facets that, as Hay (1895) also observed, are circular. The following aortal facets become successively elongated, until, with the tenth or twelfth vertebra, these facets have evolved into a long pair of slits, usu- ally narrower at the midpoint. Posteriorly, these slitlike aortal facets remain basically the same shape until, at the end of the dorsal trunk region, they merge with the basapophyses to form haemal facets. Hay (1895: 54-57) states that the cartilaginous aortal supports penetrate deeply into the centra of younger individuals, while in older specimens they rest superficially on the centra. The aortal facets are deeper and more distinct than the neural facets. Beginning with approximately the tenth or twelfth vertebra, the slit-shaped aortal facets are vertically situated on either side of an indentation that contains the aorta (Fig. 11). The first four centra have thicker and shorter supports with relatively little or no space between them. The aorta lies ventrally under the basioccipital, which bears aortal supports whose facets are of the same shape as the first four centra (Estes and Berberian, 1969, fig. 2B for A. frafi^osa). The aortal facets of the first eight vertebrae are different from all other trunk vertebrae, whose shape, as mentioned above, is basically an elongated slit. These aortal supports are thus helpful in dis- tinguishing the first eight or so vertebrae from the remainder of the trunk centra in disarticulated specimens (Fig. 11). Haemal facets. The haemal facets, which contain a cartilaginous layer be- tween the centrum and the haemal arches, are nearly rectangular-shaped pairs that do not vary along the caudal portion of the vertebral column until the first fused urals. The furrow or indentation that lies between the aortal facets in the trunk centra con- tinues in the caudal region between the paired haemal facets, although it gradually decreases in width and depth. Unlike the neural facets, the haemal facets are out- lined by an ossified border, which can be helpful in distinguishing dorsal from ventrid surfaces in disarticulated caudal vertebrae. Since the ossified walls are tilted 20 degrees posteriorly to accommodate the haemal arches, which articulate with the cartilaginous layer diagonally rather than laterally, those borders are also useful in determining the anteroposterior orientation of the centrum. Basapophyses. Amia trunk centra are distinguished from the caudal vertebrae by their having prominent paired processes, which ha\e been called transverse pro- cesses, parapophyses, or diapophyses. I follow the terminology of Bolk et al. ( 1936), wherein they designate these structures, which are the processes for pleural ribs, as basapophyses ( "basalstiimpfe" ) . The first centrum often lacks these basapophyses Fossil Amiids • Boreske 31 20 44 24 48 28 56 8 32 ])etween the basapophyses is generally still available in 32 Bulletin Museum, of Comparative Zoology, Vol. 146, No. 1 fossil forms, even in those with broken basa- pophyses, it is used here as a basis of comparison between the Recent and fossi! forms. Since the angles steadily decrease posteriorly along the vertebral column (Figs. 12, 14), they are also useful in orienting disarticulated centra to approxi- mate position along the column. Although there is individual variation in these angles (Tables 10-12), they are nevertheless con- sistent enough to help in determining the general position in the column of any single trunk centrum. The range of angles extends from approximately 180 degrees anteriorly to 45 degrees posteriorly. Since the three A. calva specimens studied were of varying sizes ( 193 mm SL, 382 mm SL, and 423 mm SL), it would appear that there is no significant change in the angles with increasing size or age of the fish (Fig. 14). Although this transition is not perfectly linear, the angles are always decreasing posteriorly, and at least in the specimens I measured, there was never an instance of an angle's measurement being greater than that of the preceding centrum. The angle decrease occurs at a fairly constant rate until approximately the thirtieth trunk ver- tebra, at which point the rate of decrease of the angles is much accelerated ( Fig. 14 ) . The angle of the basapophyses is thus a reliable parameter in identifying the gen- eral position of isolated trunk centra. Foramina, bone ridges, and first centrum. The trunk centra of A. calva have lateral foramina that, although lacking the uni- formity of the neural and aortal facets, occur in irregular, distinct paired linear patterns. The foramina of the tnmk and caudal vertebrae transmit numerous small blood vessels. On the lateral surfaces perpendicular to the anterior and posterior articular surfaces of the individual centra are prominent bone ridges. These bone ridges add support to the arch anlagen, and also help unify the anlagen into a sturdy, functional vertebral body (Schaeffer, 1967). Externally, these bone ridges are not as regular as they are internally, although they still lie antero- posteriorly in the lateral and ventral regions and extend vertically along the basapo- physes. They are also quite prominent in the notochordal furrow. Such bone ridges are not a unique feature of A. calva, and are common in teleosts. The centra in A. calva are amphicoelous. The first four to six centra differ from all corresponding centra by having the anterior articular surface more convex than concave. The first centrum in nearly all specimens observed lacked basapophyses, and should therefore be considered a minor taxonomic character since first centra do occasionally occur with very small basapophyses. The Angle of basapophyses Fig. 13. Index to the measurements used, superimposed upon an outline drawing of an Amia calva vertebra. Fossil Amiids • Boreske 33 ^. co/yo verfebrol lengths 1( 24 n <0 eth. Golden Valley Formation, North Dakota: PU 18567, coronoid teeth and vertebrae. Wasatch Formation, Wyoming: PU 13260, tooth plates; PU 13259, cranial fragments and dentaries. Bridger Formation Wyo- B Fig. 15. A, Amia calva. Recent, Wisconsin; above, lateral, and below, dorsal views of skull. B, Amia scutafa. Early and Middle Oligocene; above, lateral, and below, dorsal views of skull (sensory canal system and pit-lines are not known since skull elements are in articulation). C, Amio fragosa, Late Cretaceous to Middle Eocene; above, lateral, and below, dorsal views of skull (sensory canal system and pit-lines after Estes, 1964). D, Amia uinfaensis, Paleocene to Early Oligocene; above, lateral, and below, dorsal views of skull (sensory canal system is only known in the mandible, operculum, nasal, lacrimal, antorbital, extrascapular, and suprascapular, all of which conform with those of A. calva). Abbreviations: a, angular; ao, antorbital; br, branchiostegal rays; d, dentary; ds, dermosphenotic; es, extrascapular; fr, frontal; io, interoperculum; io- io'' io'* io'^', infraorbital series (suborbitals & postorbifals); la, lacrimal; m, maxilla; n, nasal; op, operculum; p, preoperculum; pa, parietal; pt, pterotic; r, rostral (ethmoid); s, suprascapular; so, surangular; sm, supramaxilla; so, suboperculum. Dotted lines indicate the sensory canal system; dashed lines indi- cate pit-lines. 38 Fossil Amiids • Boreske 39 ining: YPM 6245, vomer and cranial frag- mcMiVs; YPM 6246, vertebrae; YPM 6247, dentary; YPM 6248, vertebra and cranial fragments; YPM 6254, verte]:)rae, basioccip- ital, vomer; YPM 6261, left opercnlnm; ANSP 5630, vertebra. Green RivcT Forma- tion, Wyoming: MCZ 5341, FMNH 2201, complete specimens. Known distribution. North Dakota, Wyoming, Montana, and Alberta. Revised diapwsis. Vertebral colimm with significantly fewer total ctMitra (65 mean) than the other species, with approxi- mately twelve fewer trnnk vertebrae (25 mean) and eight fewer monospondylous caudal centra ( 17 mean ) . Distance be- tween anal fin insertion and the end of the vertebral column relatively short, with dorsal fin terminating close to caudal fin. Caudal lepidotrichia 19-20 rather than 23- 27. Ascending processes of parasphenoid perpendicular to the main anteroposterior parasphenoid axis; more posterior place- ment of parasphenoid tooth-patch. Pari- etals squared in outline. Marginal teeth simple pointed cones, palatal teeth usually stout styliform crushers. Supraorbital sen- sory canal not entering parietal. Excava- tion of orbital notch in frontal relatively larger. Dentary with additional horizontal shelf of coronoid articulation surface adja- cent to lingual border of alveolar ridge; coronoid articulation surface extensive, overlapping ventral half of ramus; dentary with pronoimced arch rather than gradual curve in ventral outline. Greatest known standard-length 510 mm. Introduction Jordan (1927) described Kindleia fra^osa as a new genus of cichlid fish from the Late Cretaceous Edmonton Formation of Al- berta. This tentative placement of Kindleia within the Cichlidae was largely the result of his misinteipreting the splenial tooth plates for fused lower pharyngeal bones (Estes, 1964). One month later, Russell ( 1928a) independently published a descrip- tion of Stylomyleodon lacus, a new fossil amiid from the Late Paleocene Paskapoo Formation of Alberta, and referred other specimens from the Edmonton Formation of Alberta to the same species. His descrip- tion also included a dentary and palatal teeth modified tor crushing. His relegation of the genus to the Amiidae was based on a correct inteipretation of the "splenial" (= coronoid) tooth plates (Estes, 1964). He suggested a relationship of Stylonujle- odon to Platacodon nanus ( at that time erroneously considered an amiid; see Estes, 1964) with the essential difference being hemispherical rather than Hattened tooth crowns. Jordan later ( 1928 ) noted the similarity of the two genera Kindleia and Stylomyle- odon and asserted the prior claim of his name Kindleia. Although he made no com- ment on Russell's attributing Stylomyleodon to the Amiidae, he rejected Russell's com- parison of that genus with Platacodon on the basis of Marsh's earlier conviction that the latter was mammalian. In reply to Jordan, Russell ( 1928b ) defended the valid- ity of his genus on the supposition that its dentary was distinct from that of Kindleia, although he did agree on the similarity of teeth and jaw fragments of the two genera. Russell (1929) further attempted to vali- date Stylomijleodon as a genus by com- paring his type with new specimens collected by Princeton University. This new material confirmed his association of the maxilla-dentary and palatine-coronoid den- titions, and also substantiated his interpreta- tion of Stylomyleodon as an amiid in which the coronoid teeth were specialized for crushing. He also admitted that there was insufficient Platacodon material to deter- mine any conclusive similariti(\s with Stylo- myleodon, but, r(>f(Mring to Hatcher's (1900, 1901 ) work, did insist that Platacodon was a fish. Simpson (1937) reported finding additional specimens of Stylomyleodon Rus- sell in the Fort Union Formation at Crazy Mountain Field sites of Montana. Estes ( 1964 ) , from his studies of amiid material from the Lance Formation of Wyo- ming, observed that whereas the type dentary referred by Russell to Stylomyleo- 40 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 don was the pcsterior portion of an amiid dentary, Jordon's type was the anterior portion. From this fact he confirmed the synonymy of Stylomtjleodon with the genus Kindleia, at that time beUeving that it was generically separated from Amia. Janot ( 1967 ) agreed with Estes on the synonymy of Stylomijleodon with Kindleia, but did not find sufficient cause to distinguish Kindleia generically from A7?j/fl. Russell (1967) con- tinued to leave the nomenclatural problem of Stylomyleodon-Kindleia unsettled. Estes and Berberian ( 1969) studied material from the Late Cretaceous Hell Creek Formation of Montana and confirmed Janot's proposi- tion that Kindleia is a synonym of Amia. They also suggested the possibility of synonymy of A. frantral ridges found in Amia calva. The size of the centrum indicated to Leidy that A. gracilis was a smaller species than A. calva (Leidy, 1873b). The vertebra 46 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 (ANSP 5360) corresponds approximately to the twelfth trunk vertebra in Ainia, since the aortal facets are oblong and unridged (Fig. 11). Although A. gracilis is small, it falls well within the size range of A. fra^osa and A. calva, and is considered as a nomen duhium. Estes and Berberian (1969: 10) suggested the possibility of synonymy of Paramiatus gurleyi (Plate 1) and Atnia frar- cnlnni height/width ratio. Thns A. russclli conforms with many of the most distinct characters of A. kehreri and A. valencien- nesi, and should be considered a synonym of the latter. Estes (1964) re-evaluated Dechaseaux's ( 1937) redescription of the Early Oligocene Amia munieri from France and noted simi- larities with A. fraiS,osa which included ( 1 ) styliform vomerine teeth, (2) branchiostegal rays rounded distally, ( 3 ) larger infraorbital 4 than infraorbital 5, and (4) similar parietal /frontal proportions. The principal difference between the forms is the small excavation for orbits in A. munieri. Since Dechaseaux's and Estes' studies, the speci- men (MNHN R4632, skull and associated cranial and postcranial elements) is being further prepared to display the cranial roof and palate more extensively. The frontal lacks a prominent excavation for the orbits as Estes (1964: 40) has noted, and in this feature A. munieri resembles A. sctitata and A. calva. A. mimieri is a very important form because it represents the only com- plete amiid specimen known from the Early Oligocene, and, as noted, it displays inter- mediate morphology of the cranial features among the species of Amia. A. munieri occurs very late in time in relation to the last known occurrence of A. jragosa in North America, and because there are no complete specimens known from this age, it represents a stage of evolution among the amiids that is not found in North America. Lehman ( 1951 ) described Fseudamia lieintzi (Troms0 Museum Naturhistorisk collections, Troms0, Norway) from a fairly complete articulated sptx-imen and two skulls from probable Eocene deposits in Spitzbergen. He differentiated this form from Amia on the basis of ( 1 ) Sinamia-\\ke metapterygoid and (2) presence of a con- cave notch on the dorsoposterior border of the operculum. Estes ( 1964 ) noted that Lehman was incorrect in his interpretation of tlie nature of the metapterygoid and operculum, and therefore suggested that Fseudamia might be placed in the genus Amia. From the examination of Lehman's plates, it appears that this form resembles A. fra^osa in its deep-bodied shape and low parietal /frontal ratio (approximately 0.410), and that it may be synonymous with A. valenciennesi and A. kehreri. I'urther preparation would possibly be helpful in uncovering palatal teeth, whose moiphology would aid in a more definitive description. Although the exact age of the Eocene deposit in which the specimen occurred is uncertain, this Spitzbergcni locality, if Early Eocene, lies on the possible migration route of amiids (and other vertebrates) between North America and Europe. Amia uintaensis (Leidy, 1873) Protamia tiintacnsis Leidy, 1873a: 98. Protamia media Leidy, 1873a: 98. Pappichthy.s plicatus Cope, 1873: 635. Pappichthtjs sclerops Cope, 1873: 635. Pappichthy.s laevis Cope, 1873: 636. Pappichthijs symphysis Cope, 1873: 636. Pappichthys corsonii Cope, 1873: 636. Pappichthys meditis Cope, 1884: pi. 4. Amia ivhiteavcsiana Cope, 1891: 2. Amia macrospondyla Cope, 1891: 2. Holotype. ANSP 5558, anterior tiimk vertebra. Paratypes. ANSP 8044, first anterior trunk vertebra; ANSP 3151, three posterior trimk vertebrae; ANSP 5622, basioccipital. Type locality and horizon. Henrv's Fork. North half of section 5, T 12 N, R 111 W, Sweetwater County, Wyoming; Bridger Formation. Age rouge. Torrejonian (Middle Paleo- cene) to Chadronian (Early Oligocene). Hypodi^m. Paleocene. Fort Union For- mation, Wyoming and Montana: PU 17117, maxillary; PU 17068, vertebrae and denta- ries; PU 162.36, disarticulated skull and trunk vertebrae; CM 25364, dentary; PU 17064, trunk vertebrae. Tongue River For- mation, Montana: PU 20578, basioccipital and vertebrae. Paskapoo Formation, Al- berta: ROM 4653, vertebrae. Eocene. Will wood Formation, Wyo- ming: PU 21173, basioccipitals; PU 17227, basioccipital and trunk vertebrae; PU 17649, 1 48 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 portion of cranium; PU 18760, skull frag- ments, dentary, and vertebrae. Wasatch Formation, Wyoming: AMNH 4635, dentaiy and maxilla. Golden Valley Formation, North Dakota: PU 18568, basioccipital. Green River Formation, Wyoming: USNM 18147, skull fragments and vertebrae; AMNH 785, complete caudal region; PU 13865, nearly complete specimen; MCZ 12916, disarticulated skull and associated vertebrae. Wind River Formation, Wyo- ming: AMNH 2437, dentary and skull frag- ments. Bridger Formation, Wyoming: CM 25362, portion of cranium and vertebral column; AMNH 4631, portion of cranium with dentaries, gular, and basioccipital; USNM 170976, maxilla; YPM 6238-6240, 6242, 6244, 6250-6253, 6257-6258, vertebrae and basioccipitals; USNM 170973, 5450, 3962, 3963, 3966, PU 20523, 10101, ANSP 2337-2339, vertebrae; USNM 2181, ANSP 5632, trunk vertebrae; USNM 3959, trunk and caudal vertebrae; ANSP 5580, mid- trunk vertebra; AMNH 2539, anterior por- tion of a left dentary, two premaxillae, right quadrate, left epihyal, anterior portion of an ectopterygoid, three trunk vertebrae, and numerous fragments of angular; USNM 3965, left dentary; USNM 3968, anterior dentary fragment; AMNH 2570, pre-maxil- lary fragment, fragments of angular, left quadrate fragment, trunk vertebra frag- ment, and a caudal vertebra; USNM 3960, PU 10099, 10110, vertebrae and a ural cen- trum; USNM 5476, basioccipital; USNM 3961, left dentary fragment. Washakie For- mation, Wyoming: FMNH 27465, 4509, ver- tebrae. Uinta Formation, Utah: CM 2382, maxillary fragment. Oligocene. Cypress Hills Formation, Saskatchewan: NMC 6197, trunk vertebra; NMC 6198, caudal vertebra. Known distribution. Montana, Wyo- ming, Utah, North Dakota, Alberta, and Saskatchewan. Revised diagnosis. Vertebral column with approximately 20 more vertebral seg- ments in total number (85) than A. fragosa, and five fewer trunk centra (31) and five more diplospondylous caudal vertebrae (21) than in the other long-bodied forms, A. scutata and A. calvo. Mid-trunk verte- brae subtriangular rather than ovoid. Pa- latal teeth sharp, greatly curved inwardly. Between 40-45 vomerine teeth as compared with 15-17 in A. fragosa, A. scutata, and A. calva. Hyomandibular more deeply notched between opercular process and extensor (dorsal) surface than in other species; opercular process relatively larger. Angle between alveolar ridge and exterior surface of the dentary forms a more acute angle than in the other species. Mandibular ramus less curved than in other species, so that angle between symphyseal ends of dentaries is relatively narrow. Greater mandible/head ratio (0.693) and head/ standard-length ratio (0.322) than any of the other forms: A. uintaensis has a head relatively longer and a mouth gape rela- tively wider than do other species. Most specimens are significantly larger than the other species, with a relatively greater de- gree of ossification of all bones. Greatest known standard-length 800 mm. Introduction Leidy (1873a) reported numerous dis- articulated vertebrae of a fossil fish related to Amia from the Bridger Formation of Wyoming. He distinguished a new genus Protamia from Amia by its "two oval fossae" ( aortal facets ) on the ventral surface of the centrum, and by large vertebrae character- istically with a much greater width to height proportion. Hijpamia, another new genus from the same locality which Leidy also related to Amia, was characterized by also being larger than A. calva, and by vertebrae whose sides converged into a "medium prominence excavated into a pair of oval fossae" deeper than those of Pro- tamia. Later ( 1873b ) , Leidy published a more complete and illustrated account of the various species of the new genera Protamia and Hypamia. In the same year Cope ( 1873 ) described a new amiid genus, also from the Bridger Formation, which he named Pappichthys. He distinguished this new genus from Amia by the "presence of Fossil Amiids • Boreske 49 only one series of teeth, instead of several, on the bones about the mouth." Osborn et al. ( 1878 ) reported other finds of Pci})- picJitJiys from the Bridger Formation which seemed to fit Cope's description. Cope (1884) further discussed his new genus, and rejected Leidy's prior nomenclatiue and description. New ton ( 1899 ) discussed this nomencla- tural controversy and asserted the validity of Leidy's genius Protamia, since Cope's later diagnosis \\'as no more effective in characterizing the new genus than Leidy's prior one. Newton bc^lieved that Cope's description of PappicJitlitjs as having only a single row of marginal teeth was taxo- nomically undiagnostic, since this condition would also include A. calva. Romer and Fryxell ( 1928 ) accepted Leidy's earlier description and genus as diagnostic, and referred PappiclitJujs to Protamia. They also mentioned Hypamia but found little to distinguish it from Amia. Hussakof (1932) continued to use Cope's name, however, and reported large speci- mens of Pappichthys from the Eocene of Mongolia. He also noted Cope's error in diagnosing the tooth characteristics of the genus, since Pappichthys {Protamia) has several rows of small teeth on the "splenial bone." In comparison with Amia he noted "points of difference in nearly every bone available for comparison," and concluded that Pappichthys was a valid genus, "not merely a group of large-sized extinct species of Amiatus." Estes (1964), like Romer and Fryxell (1928), referred Pappichthys to Protamia, and reported several vertebrae and a maxil- lary fragment from the Cretaceous Lance Formation of Wyoming. He inteipreted the increase in breadth over thickness of the vertebrae as a po.ssible "function of in- creased size," a condition that would also allow for tlie comparatively more massive nature of the maxillary fragment. He also considered the retention of this genus as arbitrary until enough materials were avail- able. Janot (1967) did not consider this single distinguishing characteristic of the vertebrae as sufficient foundation for the erecti(m of a new genus, and therefore sug- gested relerring Protamia to Amia. Estes et al. (1969) concurred with Janot in synonymizing Protamia with Amia. The present study confirms tliis synonymy; Leidy's species (1873a) has priority and the valid name of this fish is thus the oldest specific name, Amia uintacnsis. Revision of all forms referred now or in the past to Protamia is much needed, for these large amiids were diagnosed on char- acters of isohited vertebrae and skull frag- ments. This study gives more useful diagnostic characters that provide a basis on which the taxonomy of this group can be established. Fossil Record The major deposits carrying remains of Amia uintacnsis (Table 18) range in age from Middle Paleocene to Early Oligocene. Middle Paleocene specimens occur in the Fort Union, Tongue River, and Paskapoo formations and consist mostly of isolated and broken centra, and dentary and maxil- lary fragments. A nearly complete skull (PU 162.36) with associated trunk and caudal centra from the Bear Creek local fauna of Montana (Fort Union Formation) is the only articulated specimen from the Late Paleocene. The Eocene material in- cludes one complete articulated specimen (PU 13865), one complete caudal region (AMNH 785), and a disarticulated skull (MCZ 12916) from the Creen River Forma- tion. PU 13865 (Plate 3) has the axial skeleton intact in matrix, with a dislocated fifth centrum that is the only one available for three-dimensional measurements. This is also the only specimen in which a com- plete vertebral count can be taken. AMNH 785 provides excellent meristic information for the caudal region (Fig. 8C). CM 25362, from the Bridger Formation, consists of a left palatal and opercular series and an almost complete, disarticulated vertebral column that permitted the taking of a series of centrum measurements. Other skull frag- ments and vertebrae occur in many deposits 50 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 throughout the Eocene (Table 18). The latest occurrence of A. uintaensis is repre- sented by two isolated centra from the Cypress Hills Formation (Oligocene, Chad- ronian ) . Description Neurocranhan. Posterior to the spinal arterial foramina the basioccipital includes two fused vertebrae. As the basioccipitals display great variation in the morphology of the articular surface, it is difficult to char- acterize this form on the basis of this feature. However, the articular surface is generally kidney-shaped, with dorsal in- dentations bet\veen the neural facets, and ventrally there is an indentation distal to the aortal facets. In A. froii^osa and A. calva the basioccipital has ovoid articular surfaces with no dorsal indentations be- tween the neural facets (Estes, 1964: 29, fig. 15). In lateral view the distal articular surface of the A. uintaensis basioccipital is not perpendicular to the parasphenoid flanges; the dorsal half of this surface is more anteriorly directed than the ventral half. The parasphenoid is longer relative to its width than it is in either A. calva or A. fragosa, primarily in the region anterior to the ascending processes. At the point nearest the ascending processes, it lacks the pronounced convexity and die accom- panying anterior lateral notches found in A. calva and A. fragosa. The ascending processes are slightly less anteriorly oriented in ventral view than in A. calva, but more so than in A. fragosa ( Fig. 17 ) . The region posterior to the ascending processes is rela- tively shorter than in A. fragosa or A. calva; it is also more massive and more ventrally convex than in the other two forms. The posterior parasphenoid flanges resemble tliose of A. calva more tlian A. fragosa in outline as well as juxtaposition; those of A. fragosa are more laterally splayed than in A. uintaensis or A. calva. The tooth-bear- ing surface differs considerably from that of A. fragosa and somewhat from A. calva in outline and extent. As in A. calva, this surface extends anteriorly to the vomers, but its width is much greater and more constant than in A. calva, which is narrowly tapered anteriorly. Posteriorly, this surface extends further than in A. calva, but not as far as in A. fragosa. Approximately two- thirds of the tooth-bearing surface lies anterior to the ascending processes, while in A. calva this area is anteroposteriorly cen- tered, and in A. fragosa it is nearly all posterior. The tooth-bearing surface covers a greater portion of the ventral surface of the parasphenoid than in A. fragosa or A. calva; its basic outline is diamond-shaped, with the anterior apex widened and ex- tended to the vomers, while that of A. fragosa is subrectangular and that of A. calva is tear-drop shaped with the apex sharply protracted anteriorly. In A. uintaensis the distal edge of the suprascapular is convex as in A. calva, while in A. fragosa this edge is almost a straight line. The posterior border is more rounded distally than in A. calva and is convex rather than concave. In having the extrascapular rounded at the distal border, A. uintaensis is the same as A. calva and A. scutata, but differs from both of them in that th(^ posterior border is not concave, and from A. calva alone in lacking the distal posterior process. The anterior border is relatively straight, unlike the condition in A. calva and A. scutata, in which the lateral distal ends of the anterior borders are directly posteriad. As in A. fragosa the midline is shorter than in A. scutata and A. calva. As in A. scutata and in A. fragosa the pterotic is narrower at the anterior than posterior border, while in A. calva and, to an extent, in A. scutata the ends are sub- equal. As in A. fragosa they extend farther anteriorly and adjoin the frontal s postero- laterally. The dermosphenotic-pterotic su- ture is anterolaterally directed, as in A. scutata, but not as pronoimced as in A. calva. The anterolateral edge of the pterotic is indented and forms, witli the dermo- sphcnotic, an additional concavity in the outline of the cranial roof. Aside from this Fossil Amiids • Boreske 51 anterior indentation, the lateral borders are relatively straight, as eompared with the smoothly coneave exterior sides of the pteroties in A. scututa, A. calva, and A. fruf!,osa. The posterior border forms a smooth line, as in A. fragosa, and laeks the small lappet that A. scutata and A. calva display. The dermosphenotie is similar to that ot A. calva in relative size and outline, al- though it does not jut as deeply into the frontals. Its anterior border is rounded, as in A. calva, rather than sharply angular, as in A. fra^osa. The posterior half of the outer lateral border is indented to form a coneavity with the anterior tip of the pteroties. The parietal in A. uintaensis is elongated anteriorly, as in A. calva and A. scutata, while that of A. jra^osa is relatively square. The orbital excavation in the lateral sides of the frontal is shallow as in A. calva and A. scutata, while that of A. fragosa is characteristically deep (Fig. 28). The sen- sory canal cannot be determined. The frontals are more elongated relative to parietal length tlian in A. calva and A. scutata; the parietal /frontal ratio is only slightly smaller than that of A. fragosa (Table 7). The distal lateral border tapers anteromedially, and the anterior ends are relatively pointed anteriorly, forming a deep notch on the midline suture. There is a slight bifurcation of the anterior border of the nasal as in A. fragosa. The nasal bones are relatively narrower than in A. fragosa or A. calva, but are otherwise similar in shape and relative size. They are fairly well separated from the frontals, as in A. calva and A. scutata, rather than abutting them as in A. fragosa. The lacrimal in A. uintaensis resembles that of A. fragosa in general morphology, although it lacks the posterior notch for the anterior end of infraorbital 2 which is present in the other species of Amia. The lacrimal, like that in A. fragosa, is relatively longer and more tapered posteriorly than in A. scutata and A. calva. It is more dorsally convex than in the other forms, but only slightly more so than in A. fragosa. The infraorbital 5 in A. uintai'nsis is similar to that in A. fragosa and A. scutata, being less robust posteriorly than in A. calva. As in the other forms, it is narrower anteriorly than posteriorly. The ventral border is relatively straight, while that of the other forms is posteriorly convex. Infra- orbitals 2, 3, and 4 have not been identified. The vomerine tooth patch in A. uintaen- sis, as in A. fragosa, extends inore posteri- orly than in A. calva (Fig. 19). The \^omerine teeth are sharp and greatly curved posteriorly; they exceed those of A. fragosa and A. calva in number, each vomer bear- ing between 40-50 teeth, as compared to half that number in A. fragosa and A. calva. The rostral and antorbital are identical to that of the other species. Brancliiocranium. The suture between the anterior and posterior dermopalatine cannot be discerned. In A. uintaensis the dermopalatine has about twice the number of teeth as in A. calva, and the tooth patch extends more distad. The teeth are sharply pointed, as are the vomerine teeth. The hyomandibular is more deeply ex- cavated between the opercular process and the extensor ( dorsal ) surface, and the oper- cular process is more massive and extends further ventrad, forming a larger articula- tion surface, as compared with the other species of Amia. The articular surface of the quadrate is more robust than in other species of Amia and displays three cristae ventrally rather tlian dorsally as in A. calva and A. rolmsta (Janot, 1967: 144). The ceratohyal resembles that of A. calva and A. fragosa with the exception of its being thicker at the neck of the proximal end. The metapterygoid in A. uintaensis con- forms very closely to that of A. calva in outline and in the position of the anterior Fig. 19. Comparison of vomers of A, Amia calva; B, A. uintaensis; and C, A. fragosa. 52 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 basal process and the posterolateral otic process. The maxilla in A. uintaensis is more robust and relatively longer, and its pos- terior border is dorsoventrally wider than in the other forms, particularly A. fragosa. As in A. calva the small supramaxillary notch occurs more anteriorly than in A. fragoso. The dorsoposterior border is rounded, as in A. calva and A. scutata, rather than sharply angular, as in A. fragosa. Anteriorly the maxilla is deeper and more thickly ossified than in the other forms, but this may be a function of greater size. The supramaxilla resembles that of A. calva in general morphology, being elongated and narrowly tapered anteriorly, with a smooth- ly rounded posterior end conforming to the curve of the maxilla. The maxillo-supra- maxillary suture is straight as in A. calva. The premaxilla is identical to that of the other species. The dentary of A. uintaensis is similar to that of A. calva and A. scutata in lacking the dorsal shelf of the anterior lingual bor- der of the alveolar ridge which occurs in A. fragosa. The coronoids articulate more or less vertically on the alveolar ridge, as in A. scutata and A. calva. The anterodorsal region of the dentary slightly overlaps the ventral half, but not to the extent that it does in A. fragosa; A. uintaensis seems to be intermediate between A. fragosa and A. calva in this feature, the latter having no such ventral overlapping at the symphyseal edge. The coronoid articulation surface of the A. uintaensis dentary is thicker than in A. fragosa and A. calva, but only slightly more so than in A. scutata. At the termina- tion of this surface, this thickened area of bone forms the dorsal wall of the Meckelian groove, as in A. calva. The ventral wall of this groove is less well defined than in A. calva, witli A. scutata being intermediate. The anterior half of the dentary length in A. uintaensis is evenly tapered to the symphyseal edge; it is elongated and lacks the sharp curve present in A. fragosa at the midpoint of the alveolar ridge (Fig. 18). There is only a trace of such a curve in the dentaries of A. calva and A. scutata which are also more elongated and evenly tapered than in A. fragosa, although not to the ex- tent that they are in A. uintaensis. Anteri- orly, the bone is also relatively thicker than in A. fragosa and A. calva; A. scutata also displays this greater ossification at the anterior end of the dentary. Posteriorly, the dentary is very similar to that of A. calva. The coronoid teeth are sharp and conelike, extending to the midpoint of the lingual surface, as in A. calva. As Janot (1967) shows for A. robusta, the alveolar ridge is more horizontal in A. uintaensis and forms a more acute angle with the exterior surface of the dentary than it does in A. fragosa or A. calva; A. scutata is interme- diate between A. uintaensis and A. calva in this feature (Fig. 18). In A. uintaensis the first coronoid (symphyseal) overlies only the dorsal half of the anterior articular sur- face of the dentary, as in A. calva and A. scutata. The teeth are more sharply pointed than in any of the other forms (Fig. 18). The second coronoid is fragmentary, but appears to resemble that of A. calva with the exception of its having more sharply pointed teeth. The prearticular specimens available are fragmentary, but the lingual surface possesses blunt-conical teeth similar to those in A. calva and A. fragosa. Bor- sally, however, these teeth are as sharply pointed as the coronoid teeth. The angular is slightly longer and higher than that of A. calva. The posterior border is more ver- tical, with the articular notch less pro- nounced. It is more heavily ossified than in A. calva, but this may be a function of size. The surangular in A. uintaensis is basically similar to that of A. calva, al- though it is situated more dorsally and is more rounded at the dorsal edge. The gular is longer than that of A. calva and A. fragosa (Fig. 20). It is also slightly narrower at the posterior end than the anterior end, while the reverse is generally true in A. calva. Otherwise, the gular strongly resembles that of A. calva. Despite a few minor dissimilarities, the preopercu- lum resembles that of A. calva. There is a Fossil Amiids • Boreske 53 Fig. 20. Comparison of gulars of A, Amia calva; B, A. uintaensis; and C, A. fragosa. slightly more pronounced concavity in the ventroposterior border than is exhibited in A. calva; this concavity is altogether lacking in A. fragosa. The line of curvature is about the same as in A. calva; in A. fragosa the preoperculum is more deeply curved. The dorsal half is not quite as wide as the ventral half, while in A. calva both ends are fairly equal. In A. fragosa, however, the dorsal half is much narrower and more tapered than the ventral half, which is rela- tively wider and bulbous. The operculum in A. uintaensis is similar to that of A. calva and A. scutata in operculum-depth/ operculum-length (Table 7). The suboper- culum conforms in general morphology with that of A. calva, although it is slightly more robust, particularly in the posterior region. The corners tend to be angular, as in A. scutata and A. calva, rather than rounded, as in A. fragosa. The interoperculum is similar to that of A. calva, although more robust. The anterodorsal border is more convex than in A. calva, and is more deeply impressed into the preoperculum. The anteroventral border is narrowly tapered as in A. calva, rather than smoothly rounded as in A. fragosa. The first branchiostegal ray conforms to that of the other species. Al- though the lack of articulated material makes any count of the rays difficult, in MCZ 12916 there are 12 disarticulated branchiostegal rays on the right side of the cranial roof. As in A. fragosa the distal ends of the rays are consistently rounded, rather than squared as in A. calva. Post-cranial skeleton. The supracleithrum in A. uintaensis resembles that of A. calva and A. fragosa, excepting the dorsal articu- lation surface, which is rectilinear rather than pointed as in A. calva. The distal lateral border in the Paleocene specimens lacks the notch that occurs in A. calva, but this notch is present in the Eocene speci- mens. The metacleithrum in A. uintaensis is more elongated than in A. calva and A. fragosa. The dorsal end is narrower than in A. calva, and the ventral end is sciuared off. The cleithrum in A. uintaensis is largely similar to that of the other Ainia species, but is more massive at the proximal end than in A. calva, and the dermal sculpture covers a greater area than in A. calva, ex- tending to the distal border as in A. fragosa and A. scutata (Fig. 21). The mid-distal border is smoothly convex and lacks the notch ventral to the metacleithrum which is present in A. calva. The preceding study of the vertebral skeleton of A. calva revealed changes in height/ width proportions, position of chordal foramen, configuration of neural and aortal facets, and in the basapophyscal angles and length of basapophyses which may be used here to discern similar trends in A. uintaensis centra, for the fossil verte- brae display the same features characteris- tic of the Recent species even in disarticu- lated state. CM 25362 from the Bridger Formation is the only specimen that has a relatively complete, disarticulated, undistorted verte- B Fig. 21. Comparison of cleithra of A, Amia calva; B, A. scufata; C, A. uintaensis; and D, A. fragosa. 54 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 bral column; as the centra are separable this specimen is useful in comparisons with iso- lated vertebrae. There are 59 centra pres- ent: 25 trunk centra and 34 caudal centra, including two fused urals. Many of the preserved caudal centra are only fragments. Since the articulated specimen (PU 13865) has 85 vertebrae (see Table 9 for regional numbers) it may be assumed that about 25 vertebrae are missing from CM 25362. When comparing vertebrae from different regions of the column in the two specimens, it appears that CM 25362 lacks approxi- mately six trunk and approximately twenty caudal centra. The first anterior trunk cen- trum present in the CM 25362 series pos- sesses aortal facet configurations similar to those of the seventh vertebra of the articu- lated specimen ( PU 13865 ) . An articulated but separable series of six uncrushed an- terior trunk vertebrae (PU 10101), also from the Bridger Formation, aids in the reconstruction of the anterior region of the A. uintaensis vertebral column (Figs. 22- 25). The basapophyseal angles of these six PU 10101 vertebrae do not vary from 180 degrees. The first six anterior trunk verte- brae from a partly disarticulated vertebral column from the Paleocene specimen (PU 16236) resemble the six PU 10101 centra in length and shape of aortal facets, even though PU 16236 is a smaller individual. The height/ width ratio of these latter centra is difficult to determine, however, since the specimen underwent postdepositional crush- ing. The nearly complete vertebral series of the CM 25362 specimen has been used for the construction of the remaining trunk and caudal region in the model of the A. uintaensis vertebral column. The trunk centra of CM 25362 have been arranged according to basapophyseal angles that de- crease from 180 to 46 degrees, as in A. calva. Decreasing size was used to arrange the caudal vertebrae. Although A. uintaensis occurs much ear- Table 14. Angle of basapophyses, length, height, and width of vertebrae of Amia uintaensis compared with type specimens of synonymized taxa as illustrated IN figure 22 Relative Vertebral Number Specimen Angle of Basapophyses ( Degrees ) Length (mm) Height (mm) Width (mm) 6 A. uintaensis PU 10101 A. uintaensis PU 16236 P. uintaensis ANSP 8044 P.'sp. USNM 170973 A. uintaensis PU 10101 A. uintaensis PU 16236 A. whiteavesiarm NMC 6197 P. sp. FMNH P27465 A. uintaensis PU 10101 A. uintaensis PU 16236 P. sp. USNM 3966 P. medius USNM 3959 A. uintaensis PU 10101 A. uintaensis PU 16236 A. uintaensis PU 10101 A. uintaensis PU 16236 A. uintaensis PU 13865 A. uintaensis PU 10101 A. uintaensis PU 16236 P. uintaensis ANSP 5558 ISO" 180° 180° 180° 180° 8.0 32.0 45.0 6.0 31.0 44.0 8.0 32.0 46.0 5.5 19.0 29.0 8.0 32.0 44.5 8.5 28.5 39.0 8.5 29.0 40.0 9.0 28.0 36.0 10.0 33.0 43.0 8.0 30.0 41.5 8.5 21.5 29.0 8.5 22.0 30.0 10.0 33.0 44.0 7.5 33.0 40.8 11.0 33.5 44.0 9.5 31.5 39.0 4.5 16.5 21.5 11.0 34.0 42.5 9.0 33.5 34.0 10.5 32.5 40.0 Fossil Amiids • Boreske 55 licr in time than A. calva and A. scutata, it liodicd form tlian its contemporary, A. has approximately the same total number of fra^osa, which has a mean of 65 centra, centra (85), and like them is a longer- Tlu> vertebral column of A. umfaenm does, B 4. uintaensis PU lOIOI A. uintaensis PU 16236 P. uintaensis ANSP 8044 A . whiiteavesiana NMC 6197 R sp. USNM 3966 A. uintaensis PU 13865 P. sp. USNM 170973 P sp. FMNH P27465 R uintaensis ANSP 5558 P medius USNM 3959 Fig. 22. First anterior trunk vertebrae (A,B) of Amia uinfaens'is compared with type specimens of synonymized taxa (refer to Table 14 for data). 56 Bulletin Museum of Comparative Zoologij, Vol. 146, No. 1 11 R sp. P. sp. USNM 170973 USNM 3962 P. Sp. USNM 170973 "-^,/ P. plicotus AMNH2539 P sp. PU 20523 P Sp. USNM 170973 P sp. USNM 170973 12C 12d i Amia. sp. ANSP2337 P. plicafus USNM 170974 P. medius USNM 3959 12a p. medius P. sp. YPM 6238 FMNH P27465 P. medius USNM 3959 12b Amia sp. ANSP 2339 Fig. 23. Seventh through fourteenth mid-trunk vertebrae of Amia uinfaensis compared with type specimens of synonymized taxa (refer to Table 15 for data). Fossil Amiids • Borcske 57 however, differ meristieally from that of 36 (mean) in A. sctitata. The number of A. calva and A. scutata in number of verte- diplospondylous vertebrae is 20-21, as eom- brae in the various regions. There are 31 pared with 14-17 in A. calva and 15 in trunk eentra in A. uintacnsis (PU 13865), A. scutata. This variation from A. ra/i;« and as opposed to 37 (mean) in A. calva and A. scutata in the organization of the verte- 15 /? plicotus P. medius USNM 3958 USNM 3959 R medius YPM 6239 R sp. USNM 3966 P symphysis PU 10099 19a 19b P medius USNM 3959 R sp. USNM 3966 R sp USNM 3962 P sp. P sp. USNM 3966 FMNH P27465 P medius YPM 6240 P sp. USNM 3966 19C R sp. R sp- FMNH P27465 USNM 3963 P. sp. P medius USNM 3966 USNM 3959 R medius USNM 3959 R loevis USNM 3968 P uintaensis ANSP 3151 Fig. 24. Fifteenth through twenty-second posterior trunk vertebrae of Amio uiniaemis compared with type speci- mens of synonymized taxa (refer to Table 16 for data). 5(S Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 26 27 28 29 30 31 Psp. YPM 6242 /? uintaensis ANSP3I5I P. loevis PU 10109 R laevis PU 10109 P. media ANSP5632 P Sp. USNM 3963 Amia sp. ANSP2338 P symphysis PUIOIIO A. mocrospondyla NMC 6198 34 37 39 43 44 45 50 51 59 m P. laevis USNM 3968 P sp. FMNH PF4509 P sp. USNM 3966 P. medius USNM 3959 P. medius USNM 3959 P sp. USNM 5450 P. medius USNM 3959 P. corsoni USNM 3961 Fig. 25. Posterior trunk and caudal vertebrae of Am/o umfoensis compared with type specimens of synonymized taxa (refer to Table 17 for data). Fossil Amiids • Boreske 59 Table 15. Angle of hasapophyses, length, height, and width ok vehtehuae of Amia iiintacn.sis compared with type specimens of synonymized taxa as illustrated IN figuhe 23 Relative Vertebral Number Specimen Angle of Basapophyscs ( Def^rees ) Length (mm ) Height (mm) Width ( mm ) 7 8 9 11 12a 12b 12c 12d 14 P. sp. USNM 170973 P. sp. USNM 3962 P. sp. USNM 170973 P. plicatus AMNH 2539 P. sp. USNM 170973 P. .s7>. PU 20523 P. sp. USNM 170973 P. 77ic(lius YPM 6238 P. sp. FMNH P27465 P. mcdius USNM 3959 Aiiiiu sp. ANSP 2339 A7nia sp. ANSP 2337 P. ])licatus USNM 170974 P. mcdius USNM 3959 179° 8.5 19.5 25.5 7.0 20.0 28.0 178° 8.5 19.5" 26.0 177° 6.0" 18.0^ 24.0 8.0 21.0 25.5 174° 7.5 25.0 30.0 7.5 22.0' 25.0 6.0 19.0 24.0 171° 11.0 30.0 35.0 9.0 22.0 28.0 167° 10.0 29.0" 40.0 166° 10.0 29.0 38.5 163° 8.0 24.0 30.0 160° 7.5 19.0 23.0 Est. Taulk 16. Angle of hasapophyses, ijiingth, hkic;ht, and width of verteijuae of Amu/ uintactisis compared with type specimens of synonymized taxa as illustrated IN figure 24 Relative Vertebral Number 15 17 18 19 19a 19b 19c 20 21 22 Sjiccimen P. plicatus USNM 3958 P. mcdius IfSNM 3959 P. mcdius YPM 6239 P. sp. USNM 3966 P. sijmphysis PU 10099 P. mcdius USNM 3959 P. sp. USNM 3962 P. sp. USNM 3966 P. ,v/;. USNM 3966 P. sp. FMNII P27465 P. i7icdius YPM 6240 P. sp. USNM 3966 P. sp. FMNH P27465 P. sp. USNM 3963 P. sp. USNM 3966 P. Tticdius USNM 3959 P. mtY/tt/.v USNM 3959 P. /at't;i.v USNM 3968 P. uiiitacrisis ANSP 3151 Anglo of Basapophyses ( Degrees ) Length (mm) Height (mm) Width ( mm ) 156° 8.5 8.5 8.5 8.5 5.5 22.5 23.0 22.0 21.0 15.5 25.0 27.0 24.5 25.5 20.0 153° 9.0 9.0 24.0 23.0 29.0 28.0 149° 9.0 21.0 25.5 143° 8.5 11.0 23.0 28.0 27.8 33.0 139° 7.0 19.0 23.0 138° — — 136° 11.0 7.0 25.0 16.5 35.0 20.0 132° 9.0 9.0 21.0 22.0 27.0 26.0 122° 9.5 22.0 27.0 117° 10.0 12.0 26.0 28.0 30.0 29.0 60 Bulletin Museum of Coniparative Zoology, Vol. 146, No. 1 Table 17. Angle of basapophyses, length, height, and width of uintaensis compared with type specimens of synonymized taxa IN figure 25 vertebrae of Amia as illustrated Relative Vertebral Number Specimen Angle of Basapophyses ( Degrees ) Length ( mm ) Height ( mm ) Width ( mm ) 24 P. sp. YPM 6242 102° 9.0 17.0 22.0 26 P. tiintaensis ANSP 3151 97° 10.0 23.0 28.0 27 P. laevis PU 10109 90° 13.0 29.0 33.0 28 P. laevis PU 10109 83° 14.0 28.0 30.0 29 P. media ANSP 5632 80° 8.0 16.0 18.0 30 P. sp. USNM 3963 P. symphijsis PU 10110 62° 11.0 6.0 14.0 13.5 31 Amia sp. ANSP 2338 A. macrospondyla NMC 6198 46° 13.0 12.0 26.0 25.0 23.0 22.0 34 P. laevis USNM 3968 6.5 22.0 18.0 39 P. sp. FMNH PF 4509 7.0 19.0 18.0 43 P. s/;. USNM 3966 7.0 19.0 18.0 44 P. medins USNM 3959 6.0 16.0 18.0 45 P. 7ne(/ii« USNM 3959 5.5 17.5 18.0 50 P. s/;. USNM 5450 7.0 15.0 13.5 51 P. »!«//»« USNM 3959 5.0 17.0 11.0 59 P. corsonii USNM 3961 4.0 11.0 10.0 bral column into region.s and types of verte- brae appears to be a useful taxonomic character of A. nintaensi.^. The neural, aortal, and haemal facets do not appear to vary much from those of A. calva. The first six ventral aortal facets show basically the same pattern for both species (Figs. 11, 22). The angle of basa- pophyses in A. uintaensis differs from that of A. calva in two ways. The first six verte- brae all have basapophyseal angles of 180 degrees, and it is not until the seventh vertebra that these angles gradually begin to decrease. Because of this more posterior beginning in the decrease of the angles and because there are fewer tinmk vertebrae, the rate of decrease of the basapophyseal angle is greater. These angles range from ISO degrees anteriorly to approximately 45 de- grees posteriorly, about the same as the range for A. calva. The intracolumnar variation in centrum shape seen in the vertebral column of Re- cent A. calva also occurs in A. uintaensis ( Fig. 14). In some respects the latter shares certain characteristics with A. calva. The first centrum is broad and thin, and usually lacks basapophyses (Fig. 22). However, centra between the fourth and twentieth vertebrae begin to acquire an almost sub- triangular outline, as opposed to the sub- elliptical form of the A. calva trunk centra (Fig. 12). The subtriangular shape may be a function of greater size of the centra. The chordal foramen is open in all known vertebrae of A. uintaensis from the Paleo- cene. Eocene, and Oligocene, but is often filled with detritus during fossilization. Estes (1964: 42) observed that Cretaceous specimens as well as the Late Faleocene specimen (PU 16236) had the chordal foramen smoothly closed with bone. A re- Fossil Amiids • Boreske 61 examination of PU 16236 reveals that the chordal foramen is actually filled with fine sediment rather than bone, so that the character of the closed foramen can only be applied to the Cretaceous specimens. Chordal foramen position in all specimens shows slight intracolumnar variation along the tiimk as in A. calva, although occurring more dorsally. In the caudal region there is virtually no difference between the two forms. Leidy characterized "Protamia" uintaensis on the basis of five centra and one basioc- cipital. His height/ width proportions were described in relation to those in an un- diagnostic intracolumnar standardization of the centra of the A. calva vertebral column. My measurements of the anterior trunk centra reveal that the holotypc ANSP 5558 has a width 1.3 times the height, and para- type ANSP 8044 has a width 1.6 times the height. Other paratype centra are posterior trunk centi-a with width/ height ratios of approximately 1:1. Romer and Fryxell (1928: 521) described a displaced posterior trunk centrum as having a height of 10 mm and a width of 12.5 mm, about the same as in ANSP 5558. Estes (1964: 43), in his discussion of the height/ width proportions of A. uintaensis centra, misinterpreted Leidy 's (1873a, 1873b) diagnosis of "Proto- mia' uintaensis and Romer and Fryxell's (1928) diagnosis of "Paramiatus ^wleyij' indicating that vertebrae of tlie former were three times as wide as deep, those of the latter two times. Estes was correct, how- ever, in his assumption that there is intra- columnar variation in height/ width ratios. The general pattern of intracolumnar variation in the A. uintaensis vertebral column is quite similar to that of A. calva; there is the same trend from horizontally elliptical centra to circular or vertically elliptical centra ( Fig. 14 ) . Thus the earlier diagnoses of A. uintaensis using height/ width ratios that attributed the proportions of the anteriormost trunk vertebrae to the entire column are undiagnostic. On the basis of isolated centra and skull material, the most commonly used character in differentiating A. uintaensis from A. calva has been the former's greater size. How- ever, the articulated specimen (PU 13865), which is the smallest known A. uintaensis, is only 146 mm longer than A. fra^osa (FMNH 2201) and 16 mm longer than the largest A. calva known to me (UMMZ 197683). Estes (1964) suggested that the widening of the A. uintaensis vertebrae might be a function of its greater size; Gould's (1966) statement that internal elements generally increase at allometric rates to provide sufficient surface area to maintain the external surface area offers a partial explanation as to why the large A. uintaensis vertebrae have greater width in proportion to height than they do in smaller amiid vertebrae. Discussion Two species of Protamia, one species of Ilypamia, six species of Pappichthys, and three species of Amia have been described on vertebral characters from isolated centra and disarticulated cranial elements (Table 19). With the exception of Amia ichiteaves- iana, A. selwyniana, and A. macrosponclyla from the Oligoeene Cypress Hills Forma- tion of Alberta, all these taxa are based on material from the Bridger Basin, Bridger Formation, of Wyoming. Each of these 12 taxa will be re-evaluated in the following discussion. Of the twelve species and four genera, 'Trotamia" uintaensis (Leidy, 1873a) is the oldest name. Leidy's type specimens are all trunk vertebrae. The holotypc ANSP 5558 (Fig. 22) is approxi- mately the sixth anterior vertebra and displays the characteristic subtriangular outline of other specimens. The paratypes include trunk vertebrae ( ANSP 8044, 3151 ), and a large basioccipital (ANSP 5622). The holotypc vertebrae and the basioccipital are considered diagnostic for Amia uintaen- sis, on the basis of their possessing the characteristic subtriangular vertebral out- line, and a kidney-shaped articular surface of the basioccipital. Leidy (1873a) described Protamia media from two large trunk centra from the 62 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 Bridger Formation of Wyoming. His main criterion for distinguishing this form from A. calva and from the other species of "Protarnia" was that the vertebrae were twice the size of A. calva vertebrae and "somewhat smaller than Protamia uintaen- sis" (Leidy, 1873b). The holotype USNM 2181 appears to be from the anterior trunk region ( approximately the seventh or eighth centrum, as suggested by its proportions and configurations of aortal facets). The basapophyseal angle is approximately 178- 180 degrees. The paratype ANSP 5632 is from the posterior trunk region, with an 80- degree basapophyseal angle, which is ap- proximately equivalent to the twenty-ninth centrum in A. uintaensis (Table 17; Fig. 25). Cope ( 1884, plate 4, figs. 7-20) figured "PappiclitJiys mcdius" on the basis of 14 disarticulated centra from the same locality (USNM 3959). Eight of these are from the trunk region and correspond to centra within the anterior to mid-trunk region of A. uintaensis (Tables 15-16; Figs. 23-24). The remaining six centra correspond to cen- tra in the caudal region (Tables 16-17; Figs. 24-25). Cope gave no description, but in figuring these specimens he allocated to them his own genus, emending Leidy's (1873a) prior nomenclature. Both Leidy and Cope had apparently assumed that the characteristics of one or a few vertebrae represented those of the entire coliunn. Both species fall well within the size range of A. uintaensis (Tables 14-17), and are here considered synonyms of the latter. Leidy (1873a) describc-d Uypamia ele- gans from one small trunk vertebra. He characterized this form as possessing a cen- trum that was characteristically "short in proportion with its breadth, and it presents sutural impressions for a contiguous pair of neural arches" (Leidy, 1873b). ANSP 5580 appears to be from the mid-trunk region, comparable to approximately the nineteenth centrum as suggested by its proportions and configuration of aortal facets. The basapo- physeal angle is 138-139 degrees. These character-states and the small size are not unique, occurring as the do in all the other species of Amia; Hypamia elegans is there- fore a nomen duhium. Cope (1873) described Pappiclithys plicatus from the anterior portion of a large left dentary (AMNH 2539). Other type material included two premaxillae, a right quadrate, a left epihyal, an anterior portion of an ectopterygoid, three trunk vertebrae, and numerous fragments of angulars. He characterized this form primarily on the basis of dermal sculpture of the "cranial fragments being roughly grooved." The angular in A. uintaensis is generally marked by more pronounced dermal sculp- tiu-e than the other mandible elements. His diagnosis of the vertebrae (USNM 3958) is based on proportions and mor- phology of neural and aortal facets, both of which correspond to various trunk ver- tebrae in A. uintaensis (Tables 15-16; Figs. 23-24). The description of the re- maining elements conforms with other elements of A. uintaensis. Pappichthys plicatus is tlierefore a synonym of the latter. Cope (1873) described Pappichthys sclerops from a large left dentary. He characterized this form as possessing a dentary "more compressed and deeper" than that in A. calva and other species of "Pap- pichthys." The dentary (USNM 3965) in all respects greatly resembles all dentaries that have been referred to A. uintaensis, and I regard Pappichthys sclerops as a synonym of the latter. ('ope (1873) described Pappichthys laevis from a large anterior dentary frag- ment (USNM 3968). Other type materials include a premaxillary fragment, fragments of angulars (AMNH 2570), a left quadrate fragment, a trunk vertebra fragment, and a caudal vertebra. Although Cope distin- guished this taxon from other species of Pappichthys on vertebral proportions, vari- ances in dermal sculpture, dentary alveolar count, and obliqueness of alveolar face, these character-states occur in A. uintaensis. PappicJitJujs laevis is therefore a synonym of the latter. » Fossil Amiids • Boreske 63 Cope (1873) described Pappichthys sym- pJu/sis from two large fragments of trunk- vertebrae and a iiral (USNM 3960). His diagnosis rests primarily on eonfiguration of neural faeets and basapophyseal length. Osborn et cil ( 1878: 104) later reported two eaudal vertebrae as cotypes (PU 10099, 10110). Cope (1873) described Pappich- thys corsonii from 12 centra (USNM 5475- 5476), a basioccipital (USNM 5476), and a left dentary fragment (USNM 3961). He distinguished this form from Pappiclithys sympliysis on different neural facet mor- phology, basapophyseal length, and height/ width proportions. Merrill (1907: 14) cites ^'PappicJitJujs sympliysis = Pappichthys cor- sonii' without further discussion. The cen- tra of both forms conform to centra in the vertebral column of A. uintaensis (Table 17; Fig. 25) and the characters assigned to the dentary and basioccipital of Pappich- thys corsonii are also found in A. uintaensis; thus both P. sympliysis and P. corsonii arc synonyms of A. uintaensis. From the Early Oligocene Cypress Hills Formation, Saskatchewan, Cope (1891) described Amia wliiteavesiana from an an- terior vertebra (NMC 6197), and Amia macrospondyJa from a caudal vertebra (NMC 6198). Both these forms were founded on variations of vertebral charac- ters (height/ width proportions, lack of basapophyses, and chordal foramen posi- tion) that are also represented in the verte- bral column of A. uintaensis. The type centrum of A. tiJiiteavesiana corresponds approximately to the second anterior verte- bra in A. uintaensis (Table 14; Fig. 22), that of the type centrum of A. macrospon- (Jyla with the thirty-first centrimi in A. uintaensis (Table 17; Fig. 25). Prior to the appearance of Cope's ( 1891 ) publica- tion. Ami (1891), in his review of the Cypress Hill fauna, mistakenly listed A. whiteavesiana under the name A. selwyni- ana. A. macrospondyhi and A. ivhiteavesi- ana are here considered synonyms of A. uintaensis; A. sehcyniana is a iiornen nudum. Comments on European and Asian Forms Janot ( 1967 ) described a large amiid, Amia rohusta, from the Late Paleocene of France, on the basis of disarticulated material. She distinguished this form from A. calva and A. russeUi on the angle of the ventral border of the dentary face, and on morphology of the parasphenoid tooth- bearing surface in addition to other minor morphological differences. Many of the di- agnostic elements or associations on which A. uintaensis is based, such as coronoid and vomerine teeth, regional vertebral counts and dorsal cranial elements, are missing in her material. The elements that she does figiue, however, closely resemble the com- parative bones in A. uintaensis. Simi- larities include rounded distal ends of branchiostegal rays (also in A. fragosa), subtriangular morphology of trunk verte- brae, extensive surface of parasphenoid tooth-patch, and shallow orbital notch in frontal (also in A. scutata and A. calva). These marked similarities suggest that A. rohusta is a synonym of A. uintaensis. Current work on the relationship of the North American and European continents in the Early Cenozoic ( McKenna, 1972) indicates that they were connected until tlie Early Eocene and that there is great sim- ilarity between the Paleocene and Early Eocene mammalian taxa at that time. There is thus no zoogeographic problem in- herent in synonymizing these two species. Hussakof (1932) described Pappichthys mongoliensis from disarticulated elements from the Late Eocene Ulan Shireh beds of the Shara Murun region. Inner Mongolia (collected by the American Museum Cen- tral Asiatic Expeditions.) At the time of Hussakof's description, this collection (AMNH 6372) represented the most exten- sive material of ''Pappichthys." The collec- tion includes numerous dentaries, maxillae, three gulars, three opercula, three cleithra, an hyomandibular, a supracleithrum, a vomer, and trunk and caudal vertebrae. 64 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 Hussakof distinguished this form from A. calva by the length of the dentaries and the morphology of the operculum, and from species of "Pappichthys" and "Pro- tamia" on the basis of comparison of verte- bral size. A comparison of the Mongolian material with A. uintaensis shows some dis- similarities, but there is still a closer affinity between this form and A. uintaensis than with the other species of Amia. The vomer bears numerous sharp vomerine teeth; the hyomandibular is deeply arched, and the lingual face of the dentaries conforms to that of A. uintaensis. The dentary, however, is quite elongated anteriorly, the supra- cleithrum is narrower, and the dorsal border of the operculum is short and ascends at a 30-degree angle rather than being horizontal as in A. uintaensis (and in other Amia species). The extrascapular is narrow and tapered to a point rather than flattened medially. Thus, although Pappich- thys mongoliensis is similar to A. uintaensis in many features and is clearly related to it, it also differs in some respects. It un- doubtedly belongs to the genus Amia, and retention of all the Mongolian specimens in Amia mongoliensis seems the most practical alternative at this time. The Mongolian higher vertebrate taxa indicate that the Turgai Straits at least partially isolated Mongolia from Europe during at least part of the Cretaceous, Paleocene, and Eocene, and that probably little exchange took place until the Late Eocene (Szalay and Mc- Kenna, 1971: 280-281). It may be possible that A. mongoliensis evolved from A. uintaensis during this migration. Amia cf. uintaensis Hypodigm. Cretaceous. Lance Forma- tion, Wyoming: CM 256, YPM 6311, trunk vertebrae; UCMP 56276, two fragments of a single vertebra; UCMP 56277, one com- plete vertebra, one vertebral fragment, one left maxillary fragment. Hell Creek Forma- tion, Montana: AMNH 6385, trunk vertebra; MCZ 9334, dentary tooth tips. Aguja For- mation, Texas: UMM collections, maxillary fragment. Ojo Alamo Formation, New Mexico: USNM collections, trunk vertebra. Discussion Cretaceous specimens of large amiids occur in both Lance and Hell Creek for- mations and consist mostly of isolated and broken centra, and teeth that have been identified primarily on the basis of size. The characteristic subtriangular out- line of the trunk vertebrae is even more pro- nounced in these Cretaceous specimens, wherein the lateral centrum walls between the basapophyses and the aortal facets are concave ( Fig. 26 ) . The chordal foramen is, as Estes ( 1964: 42) noted, closed with bone, as are one-third of the vertebrae referred to A. fragosa from the Lance Formation. How- ever, Estes observed lateral concavities be- tween the neural facets and basapophyses in a large vertebral centrum (AMNH 6385) from the Hell Creek Formation (mistakenly cited by him as AMNH 6835 from the Oldman Formation of Alberta). Estes apparently confused neural with aortal facets and thus figured the vertebra upside down. Correct orientation of the centrum (Fig. 26) shows concavities be- tween the basapophyses and the aortal facets. Thus, Estes was incorrect in con- cluding that A. fragosa, A. calva, and the Eocene specimens of A. uintaensis "also seem to lack the concavity between the 'basapophysis' and neural arch present in the large Cretaceous specimens." Two other specimens from the Lance Formation (YPM 6311, CM 256; Fig. 26) also show the prominent concavities between the basapophyses and aortal, rather than neu- ral, facets. In addition to the vertebrae, Estes described a maxillary fragment as being larger and more robust than that of A. fragosa, although "characteristically amiid in tooth implantation and general shape." A more complete maxillary frag- ment (UMM collections) from the Aguja Formation ( Big Bend National Park, Brewster County, Texas) conforms with Estes' (1964) description. Fossil Amiids • Borcnke 65 B Hlii H Fig. 26. Comparison of different Cretaceous vertebrae. Am'ia cf. uinfaensh: A, anterior trunk vertebra, CM 256, Lance Formation, Wyoming; B, posterior trunk vertebra, AMNH 6385, Hell Creek Formation, Montana; C, mid-trunk vertebra, YPM 6311, Lance Formation, Wyoming. Chondrichthyes: D, E, G (thin section), trunk vertebrae, FHKSCM 13024-9, Black Creek Formation, North Carolina; F, trunk vertebra, MCZ 12879, Peedee Formation, North Carolina. Cetacean: H, caudal vertebra, FHKSCM 13025, Calvert Formation?, North Carolina. 1 rz dorsal, 2 ^ articular surface, 3 ::= ventral 66 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 Only three new centra and a maxillary fragment have been identified since Estes' (1964) study. The vertebrae, as noted above, differ in certain minor respects from the Paleocene and Eocene specimens. Whether or not this material actually repre- sents A. uintaensis or an earlier stage of evolution can only be determined when more complete Cretaceous material is avail- able. Amia scutata Cope, 1875 Amia dictycephala Cope, 1875: 3. Amia exilis Lambe, 1908: 12. Holotijpe. USNM 5374, incomplete spec- imen lacking the head and body anterior to the middle of the dorsal fin; anal and part of dorsal and caudal fins well preserved. Type locality arul horizon. Florissant, Colorado. East half of section 2, T 13 S, R 71 W, Teller County, Colorado; Flor- risant Formation. Age Range. Chadronian (Early Oligo- cene) to Orellan (Middle Oligocene). Hypodigm. Oligocene. Cypress Hills Formation, Saskatchewan: NMC 6200, 6205, vertebrae; NMC 6201, basioccipital. Chad- ron Formation, South Dakota: PU 17172, left dentary with posterior coronoid bearing teeth, and a trunk vertebra. Lower Brule Formation, South Dakota and Nebraska: FMNH PF4508, PF4509, CM 3814, verte- brae; FMNH PF4506, right vomer bearing teeth. Florissant Formation, Colorado: PU 10172, nearly complete specimen (counter- part = YPM 6243, anterior half; USNM 4087, caudal half); YPM 6241, complete caudal region (with counterpart); UMMP V-57431, nearly complete specimen; USNM 3992, partial specimen, lacking skull and tail; AMNH 2802, nearly complete skiill; AMNH 2670, partial specimen, lacking skull and caudal region; AMNH 2671, caudal region. Known distribution. South Dakota, Ne- braska, Colorado, and Saskatchewan. Revised diagnosis. Vertebral meristics similar to those of A. calva, but head/ standard-length proportion is intermediate between that of A. uintaensis and A. calva. Extrascapular thicker at distal end than in A. calva, with concave posterior border. Pterotic more similar to that of A. uintaen- sis than of A. calva; anterior portion narrow and extended laterally to the frontal. Or- bital excavations more marked than in A. calva, but not as deep as in A. uintaensis or A. fragosa. Preoperculum resembles that of A. uintaensis more than that of A. calva, being narrower dorsally than ventrally. Symphyseal incurving of the dentary less than in A. calva, but greater than in A. uintaensis. Ventroposterior process of cleithrum heavily sculptured as in A. fragosa and A. uintaensis. Infraorbital 4 larger than infraorbital 5 as in A. fragosa and A. uintaensis. Ossification of cranial bones ex- tensive as in other fossil species, greater than in A. calva. Greatest known standard- length 390 mm. Introduction Cope's (1875: 3) description of Amia scutata is based on a specimen lacking the head and body anterior to the middle of the dorsal fin, from the Middle Oligocene Florissant Fomiation near Florissant, Colorado. He distinguished this form from Amia dictyocephala (found in the same deposit; Cope, 1875) and Amia calva by its larger scales "of which only seven and a half longitudinal rows are visible above the vertebral column." Cope de- scribed A. dictyocephala from two partially complete specimens lacking skidls and caudal fins (USNM 3992, AMNH 2670), two complete caudal regions (AMNH 2671, USNM 4087), and a nearly complete skull (AMNH 2802); Osborn et al. (1878) later described another specimen of A. scutata from the same deposit. Tliis specimen was more complete, consisting of an axial skeleton and a crushed skull. They believed A. scutata to be a valid form, differing from A. calva in having a proportionately larger head. Comparison of known specimens of A. scutata revealed that the counterparts to the specimen described by Osborn et al. ( PU 10172 ) were separated and sold to two Fossil Amiids • Borcskr 67 Fig. 27. A, Amia scutata UMMP V-57431; B, A. scufafa PU 10172; C, A. "dictyocephala" USNM 3392; D, A. "dicfyocepbala" AMNH 2670. different miiseiims. The caudal portion of and is one of the paratypes used by Cope the counterpart was found in the National (1875) in his description of A. dictijo- Museum of Natural History (USNM 4087) cephala. The anterior region was found 68 Bulletin Museum of Comparative Zoologij, Vol. 146, No. 1 unlabeled at the Yale Peabody Museum (YPM 6243; Plate 4).i In 1967 another nearly eompk^te speci- men was discovered from the same deposit (Fig. 27A) and Cavender (1970: 42) re- ported the specimen A. dictyocepliala as differing from A. calva in having a larger infraorbital 4, in the sculptin-e of cleithrum, and "by its proportionately larger head and orbit, and somewhat shorter body." Fossil Record Other than the Florissant Formation, the only deposits from which elements of A. scutata can be identified are the Cypress Hills Formation of Saskatchewan, Chadron Formation of South Dakota, and the Lower Brule Formation of South Dakota and Nebraska. Becker (1961: 38) reported amiid scales (UMMP collections) from the Late Oligocene Passamari Formation and Middle Oligocene Grant Horse Prairie Shale of Montana (Becker, 1962). Since no specific characters for scales of Amia have yet been determined, it is best to allocate this material to Amia sp. Skinner et al. (1968: 415) has reported Amia .sp. vertebrae (F:AM 42947) from the Early Miocene Turtle Butte Formation of South Dakota. Only two specimens were found; since the vertebrae of A. scutata and A. calva are morphologically and meristieally similar. Skinner et al.'s identification is the only possible one at this time. The strati- graphic range of A. scutata is therefore lim- ited to the Early and Middle Oligocene. Description Neurocranium.. The basioccipital (PU 10172, NMC 6201 ) is similar to that of A. calva. The only available parasphenoid (PU 10172) is poorly preserved, but closely resembles that of A. calva in length and position of ascending processes. The extrascapular in A. scutata differs slightly from that of A. calva in that the distal end is relatively thicker and the 1 The counterparts ( USNM 4087, YPM 6243) to PU 10172 have been .subse(iuently accjuired by the Museum of Natural History, Princeton University. L< orbital length D- orbital depth ^a^dermosphenotlc angle A. fragosa D/L=O.I76mn. A. uintaensis D/L=O.I55mn. 18° Z^"! 40 = I34<' A scutata D/L=O.I32mn. I5« A calva D/L"O.IOOmn. 15° IB" 40=145° Z4»r 4.0=135° A. cf. scutata O/L'0.121 [■0=137° Fig. 28. Orbital dimensions of ^m\a spp. posterior lappet is less pronounced; also, the posterior border is more convex (Fig. 15). As in A. calva, however, the proximal anterior corner is squared off, and the medial suture is relatively long. The pterotic in Ax. scutata resembles that of A. uintaensis more than that of A. calva in general morphology, since the anterior half is narrower than the posterior half; in the Recent species the ends are nearly sym- metrical. The anterior border extends fur- ther laterally than in A. calva, and, as in A. uintaensis, adjoins the distal lateral side of the frontal, rather than the posterior border as in A. calva. The dermosphenotic, parietal, frontal, and nasal of A. .scutata conform to these bones in A. calva. The parietal/ frontal ratio is marginally within the lower limit of the range of A. calva (Table 7). The orbital excavation in the Imxssil Amuus • liorcske 69 frontal (Fit:;. 2cS) is greater than in A. caha bnt less than in A. jruii^osa or A. uintaensis. Snprascapulars, antorbitals, and rostrals are not preserved. The laerinial is similar to that of A. calva, b(>arint!; a posterior noteh for the reeeption of infraorbital 2, but in A. scutata the laeri- nial is more robust. Infraorbital 2 and infra- orbital 3 are similar to these bones in A. calva. Infraorbital 4 is more massive pos- teriorly than in A. calva; it exeeeds infra- orbital 5 in dorsoventral length, and the posterodorsal corner, which in A. calva is markedly acute is, in A. .scutata, more squared off. This bone more closely re- sembles that of A. jraii^osa: it is not avail- ablc> for comparison in A. uintacnsis. Infra- orbital 5 is less massive posteriorly than in A. calva; in this feature it resembles that of A. fra'^osa. It is also, as in A. fra^osa and A. uintacnsis, dec>per anteriorly than in A. calva. Branchiocranium. The supramaxilla in A. scutata is elongated and tapered to a point anteriorly, with a relatively straight ventral border as in A. calva. It is slightly longer and more robust posteriorly, the posterodorsal border being higher and less obliquely curved than in A. calva, A. uin- tacnsis, and A. jra<^osa. The preniaxilla resembles that of A. calva. The maxilla is wider posteriorly and more ossified anteri- orly than that of A. calva, but otherwise agrees with the bone in the Recent .species. Dermopalatine, autopalatine, entoptery- goid, ectopterygoid, metapterygoid, and vomer are not pr(\served. Ho\v(^ver, conical vomerine teeth are displayed on PU 10172, and resemble those of A. calva rather than those of A. frarmal struc- ture is limited to the cent(>r and dorsal re- gion of this part of the cleithrum. Th(> vertebral column of A. scutata re- sembles that of A. calva both in number of ccMitra (Table 9) and in general morphol- ogy of the centra. The head /standard- length proportion (0.312) is greater than in A. calva (0.271 ), but less than in A. uintaen- 70 Bulletin Museuui of Comparative Zoology, Vol. 146, No. 1 sis (0.322). The insertion of peetoral fin/ standard-length and insertion of anal fin/ standard-length ratios nrv both within the ranges of A. calva, althongh the latter pro- portion for A. scutata is somewhat greater than the mean for A. calva ( F'ig. 31 ). Discussion In the same paper as his deseription of Aryiia scutata. Cope (1875: 3) described Amia clictyocepliala, also from the Florissant Formation. A. dictyocephala was distin- guished from A. scutata by having 10 to 12 supravertebral scale rows, and 35 vertebrae between the anterior dorsal fin pterygio- phore and the posterior anal fin ptervgio- phore (USNM 3992 AMNH 2670). "lie further characterized this form from a skull (AMNH 2S02) that "possesses twelve branchiostegal rays, and a relatively smaller orbit than in Amia calva." A re-examination of these specimens in the previous section on meristics showed that Cope's supra- vertebral scale row count was in error, and there is no perceptible difference in this feature between Recent and fossil Amia species (Table 8). In A. calva, the range for the number of centra between the in- sertion of the dorsal fin and the terminus of the base of the anal fin is 33-37. In the type specimen of A. dictyocephala (USNM 3992) the number of centra is 35, and the mean number in specimens of A. scutata is 36; there is clearly no way that this fcnitiue can be used to distinguish A. dictyocephala from A. scutata and A. calva. Cope, on the basis of AMNH 2802, thought that an orbit in A. dictyocepJiala was smaller than one in A. calva, but the small size was due largely to the constriction of the orbit that resulted from crushing of the dcrmosphenotic and upward displacement of infraorbital 5. The characters that Cope used to differentiate A. dictyocephaki from A. scutata are un- diagnostie, and my studies of the specimens show no morphological or meristie differ- ence; A. dictyocepJiala is here considered to be a synonym of A. scutata. Lambe (1908: 12-13) described Amia exilis from a single basioccipital (NMC 6201 ) and two mid-trunk \ c>rtebrae ( NMC 6200, 6205) from the Farly Oligocene Cypress Hills Formation of Saskatchewan. The temporal occurrence of these elements is equivalent to that of A. scutata. Lambe's description of the basioccipital conforms to that of A. scutata in being more extensively o.ssified than in A. calva. His diagnosis of the two centra is founded on height/ width proportions, ehordal foramen position, basa- pophyseal angle, and configuration of nc>ural facets. Because A. scutata resembles A. calva in vertebral morphology, the charac- ters that Lambe uses to distinguish A. exilis are undiagnostic; I therefore consider A. exilis iis a synonym of A. scutata. Amia cf. scutata Hypodi^m. Miocene. Pawnee Creek Formation, Colorado: UCMP 38222, nearly complete cranial roof, infraorbitals 4 and 5, nearly complete anterior portion of palate, two branchiostegal rays, maxillae, and right dentary. Description The general morphology of the cranial roof resembles both A. scutata and A. calva in parietal/ frontal ratio (Table 7), rectan- gular parietals, and shape of dermosphen- otic and nasal (Fig. 29). The extrascapular more closely resembles that of A. scutata in its greater width and less pronoimced distal posterior lappets. The pterotic also resem- bles that in A. scutata in its being narrower anteriorly than posteriorly, and in bordering die frontal laterally rather than posteriorly. The size and depth of the orbital excavation is intennediate between that of A. scutata and A. calva (Fig. 28). The maxilla is similar to that of A. calva, being less robust posteriorly than that of A. scutata. The branchiostegal rays are squared off distally, as are tliose of both A. calva and A. scutata. Infraorbital 4, although posteroventrally in- complete, is clearly closer to that of A. scu- tata than A. calva in being relatively larger than infraorbital 5, and in the posterodorsal corner being squared off rather than acute as in A. calva. Infraorbital 5 resembles that I Fossil Amiids • Boreske 71 of A. scutata in size relative to infraorbital 4, the anterior end bcnng narrower than in A. scutata; this featnre eontributes to lessen- ing the relative width of the orbit. The dentary resembles that of A. scutata in being wider anteriorly than in A. calva; the dorsal lingual surfaee only slightly overlaps the ventral lingual siuface as in A. scutata (Fig. 18); Meckel's groove is thus similar to that of A. scutata. There is no available palate in A. scutata for comparison. The number of vomerine teeth is 18 and 21, which is bracketed by the range for A. calva (Estes and Berberian, 1969: 5). As Estes ( 1964 ) noted for this specimen, these teetli are sharper and more incurved ex- ternally than internally; this disparity is more distinct in this form than in the extant species. The hyomandibular, entopterygoid, ectopterygoid, dermopalatine, and pre- maxilla are poorly preserved, but appear to resemble these bones in A. calva. Discussion Estes (1964: 36) and Estes and Tihen (1964: 454) referred to this specimen as Amia sp. (and in error gave the source as White River Formation). The .specimen resembles A. scutata in some elements, A. calva in others, and is intermediate in several character-states, notably bone thick- ness and size of orbits. It does, however, appear to show a stronger resemblance to A. scutata than to A. calva, particularly in Fig. 29. Amia cf. scutata DC 38222, Late Miocene, Pawnee Creek Formation, Colorado. 72 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 the morphology of the extrascapiilar, ptcro- tic, dentary, and infraorbitals 4 and 5, and I have thus compared it with the fossil species. Since this is a form that is both morphologically and temporally interme- diate between A. scutata and A. calva, it is difficult to determine whether or not this specimen actually represents A. scutata or a later stage of evolution leading to A. calva, but it is at least of interest in documenting the slow phyletic development toward A. calva in mid-Cenozoic time. Amia cf. calva Hijpodi^m. Pliocene. Lower Valentine Formation, Nebraska: UCMP 65851, an- terior portion of left dentary and a trunk vertebra; UMMP 521S7, right nasal, ectop- terygoid fragment, unidentified cranial fragments; UMMP 421S5, right dentary fragment. Ogallala Formation, Kansas: UMMP 55574-55578, three right and two left dentary fragments; UMMP 55579, in- complete right cleithrum; UMMP 55583, a right extrascapular; UMMP 55580, a right maxilla; UMMP 55585, a left premaxilla; UMMP 55586, several scales. Discussion Smith (1962), and Estes and Tihen (1964) described as Amia sp. a nasal and dentary, and cranial fragments from the Lower Valentine Formation, Nebraska. Wilson (1968) described as Amia calva denatry fragments, a premaxilla, a maxilla, an extrascapular, an incomplete cleithrum, and several scales from the Ogallala Forma- tion, Kansas. This Early Pliocene material resembles A. calva more closely than does the Miocene A. cf. scutata specimen noted above; the elements are very lightly ossified as in the Recent species. The cleithrum is distinctly A. calva-\ike in its lack of distal marginal dermal sculpture. The dentary fragments are also thinly ossified as in A. calva, but are slightly wider relative to the dentary in the Recent species, as in the Miocene form. Temporally, this Pliocene material is later than the Miocene form and earlier than A. calva; morphologically, how- ever, the available elements conform with A. calva. Amiidae incertae sedis Hypodi^m. Cretaceous. Paluxy Forma- tion, Texas: SMUSMP 62270, dentary frag- ments, premaxillary fragment, vertebrae, maxillary fragments, and an unidentified palatal bone bearing teeth; FMNH 7050, basioecipital; FMNH 7051, mid-trunk ver- tebra; FMNH 7052, anterior trunk vertebra fragment; FMNH 7053-7054, anterior trunk vertebrae; FMNH 7055, caudal vertebra; FMNH 7056, small vertebrae; FMNH 7049, unidentified palatal bone bearing teeth. Description The dentaries are fragmentary (Fig. 30); the only diagnostic features available for comparison with other amiid forms are related to the anterior region of the dentary. The surfaces pits on the exterior side of the dentary are relatively larger and deeper than in any species of Amia. The dentaries lack the dorsal shelf adjacent to the lingual side of the alveolar ridge seen in A. fragosa. The coronoid articulation surface descends directly from the alveolar ridge, as in a Uro- cles dentary from the Late Jurassic (Pur- beck) of England (BMNH 48236). The lingual surface above the Meckelian groove is relatively short, even more so than in Amia uintaensis, and the groove itself is quite wide, more so than in BMNH 48236. The anterior portions of the dentaries are relatively straight, rather than incurved as in Amia fragosa, and are evenly tapered to the symphyseal edge. The dentary and pre- maxilla teeth are broken, but in dorsal view the interior surfaces of the broken teeth are very even, lacking the serrated outline seen in other species of Amia. Only the anterior portion of the premaxilla is present; it bears nine alveoli, conforming in this respect with all Am,ia species. The premaxilla, although incomplete, displays the anterior (ventral) edge of the large foramen that is character- istic of Amia. Only part of the anterior maxilla is present in the specimens avail- able, and since the more diagnostic aspects Fossil A muds • Boreske 73 occur posteriorly, it is difficult to determine smaller fragment (SMUSMP 62270) bears any affinities witli particular specic\s; the pillar-shaped teeth with nipple-like tips, as anterior portions that are available gener- in the tooth-bearing palatal bones in species ally conform with those of Amia. The of Amia. Posterior to the spinal arterial specific bones to which the palatal frag- foramina the basioccipital includes one ments belong cannot be identified. The fused vertebra. As in Amia fra^osa and Fig. 30. Amiidae incerfae sedis, Early Cretaceous, Poluxy Formation, Texas: A'-A-, anterior portion of left den- tary; B, premaxlllary fragment; C, anterior portion of rigfit maxilla; D, unidentified palatal fragment. XO-15 74 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 Amia caJva, the basioccipital has an ovoid articular surface with no dorsal indentations between the neural facets. The large verte- brae are thickly ossified, as in the Creta- ceous specimens of Amia cf. uintaensis. The chordal foramina are closed and the only available large mid-trunk centrum dis- plays the pronounced triangular outline characteristic of Amia uintaensis. None of the large vertebrae display the character- istic Amia aortal facets; they do, however, possess neural facets, and the mid-trunk centra bear basapophyses. The small verte- brae are also thickly ossified and the chordal foramina of the trunk vertebrae are closed. As Traquair (1911: 39) noted for Amiopsis cloUoi, the lateral sides of the vertebrae are marked by a number of vari- able excavations, or "oval fossae" (Fig. 7). These smaller mid-trunk vertebrae, unlike the large ones, display both aortal and neural facets, as well as basapophyses and lateral oval fossae. Discussion Thurmond ( 1969 : 88 ) reported "various fragments of an undetermined amiid" from the Paluxy Formation of Texas, which is the earliest known occurrence of amiids in North America. He further noted that amiid material occurred both in freshwater and marine zones and that a further descrip- tion of this material would be the subject of a later study. He was uncertain as to whether the amiids occurring in the marine zones were actually marine or were freshwater forms secondarily deposited in the marine areas. None of the material can be referred to Atnia since it displays charac- teristics of Amia uintaensis, Amia fragosa, Urocles, and Amiopsis, as noted in the above description. The vertebrae suggest the possibility of more than one form: the large vertebrae are subtriangular and re- semble Am'a uintaensis in morphology, with the exception of the lack of aortal facets on the trunk vertel:)rae. The small vertebrae are Amia frag,osa-\ike in morphol- ogy; they possess aortal facets, but also display lateral oval fossae characteristic of Amiopsis. In reviewing the European Juras- sic and Cretaceous Urocles, Lange (1968) found little morphological justification to warrant continued generic distinction be- tween Urocles and species described by Woodward ( 1916 ) as belonging to Amiop- sis from the Purbeck Beds near Wevmouth, Dorset. Lack of knowledge of the skull of Amiopsis makes it impossible to compare cranial elements with those of other amiids; the singular postcranial feature charac- terizing Amiopsis is the lateral o\'al fossae of the v^ertebrae. Although Lange suggests that both Amiopsis and Ainia evolved inde- pendently from different Urocles species- groups, it is premature to attempt to do more than indicate morphological similar- ities or dissimilarities since the phylogenetic relationship of Amiopsis with Urocles or Amia cannot be clearly defined until a much-needed review of the taxon has been completed, and until more Amiopsis mate- rial is made available for study. The Paluxy material shows resemblances to two early Aryiia species, Amia uintaensis and Amia fragosa, as well as to the Late Mesozoic European amiids, Urocles and Amiopsis. Whether the Paluxy material represents one or more forms intermediate between Atnia and Urocles (or Amiopsis) or whether it belongs to some other group of amiids that became extinct before the end of the Cretaceous cannot be deter- mined, since taxonomic evaluation of this material is limited by the lack of articulated specimens. SPECIMENS REMOVED FROM THE AMIIDAE Miller (1968: 468-470, pi. 1, figs. 1, 3, 7-9) questionably identified as Protamia sp. one large (FHKSCM 13025) and three small centra (FHKSCM 13024-9) recovered from a channel sandstone cut into the Up- per Cretaceous Black Creek Formation, Phoebus Landing, North Carolina. Since all known Ainia are freshwater forms and since these centra were associated with vari- ous marine vertebrates. Miller (1968: 467) concluded that the channel sandstone con- Fossil Amiids • Boreske 75 tained a mixed fauna, "the ehannel sand- stone formed in an estuarine or tidal en- vironment." My studies indicate that these specimens are not amiid. The smaller vertebrae are horizontally ovoid. A. uintaensis trunk cen- tra have concavities between the basa- pophyses and aortal facets ( Fig. 26, A-C ) . A thin section (Fig. 26, G) through the articular surface of one of the Nortii Caro- lina specimens (FHKSCM 13924-9) has a radial structure resembling that of S(juatimi and other sharks (Hasse, 1882, tables 17- 18). All layers are laminated parallel to the exterior surface and are crossed by various perpendicular vascular foramina. Their articular surfaces are slightly concave, while those of Ainia are markedly so. Each of the small vertebrae bear horizontal basapoph- yses as in Recent Sqiialus, and are best referred to the elasmobranchs. The large vertebra is a cetacean caudal (Fig. 26, H), possibly belonging to the Cetotheriidae (Clayton Ray, 1971, personal communication). The centrum is ovoid, with very slightly concave articular surfaces, and lacks a chordal foramen, as well as ventral facets. The dorsal facets for the accommodation of metapophyses are well defined. Since this centrum is from a ma- rine mammal, it is more probably from the Miocene (Calvert Formation?) than from the Cretaceous Black Creek Formation. Eastman (1899) described Amiopsis dar- toni from a partial opercular series, pectoral fin, and associated cycloid scales from the Late Jurassic marine Sundance Formation, South Dakota. Eastman felt that the many "stout ribs" associated with the pectoral fin suggested a well-ossified Aniia-hkc verte- bral column and the semicircular operculum conformed with that of A. ctilvu. Since the scalers are covered supt^rficially with ganoine and appear elliptical, Eastman placed this form among the Amiidae. He allocated the generic name, Amiopsis, on a temporal basis. According to Bobb Schaeffer, (1971, personal communication) the holotype (USNM 4792) and the paratypes (MCZ 9696, USNM 4793) are to be tentatively referred to the Leptolepidae on the basis of morphology of opercular series and pec- toral fin lepidotrichia. Schaeffer is currently studying the Late Jurassic North American fishes and is including a more extensive discussion of this material in his review. SUMMARY AND CONCLUSIONS This survey of the osteology, morpho- metries, and meristics of the North Amer- ican fossil amiids indicates that the extant and fossil forms fall into foiu- groups worthy of specific status: ( 1) Amia fragosa, (2) A. uintaensis, (3) A. scutata, and (4) A. colvci. All these forms, excepting A. fragosa, have somewhat elongated bodies (approximately 85 centra) and shai-p, conical coronoid and palatal teeth. Al- though the coronoid and palatal teeth of A. uintaensis are more sharply curved in- wardly, the teeth are closer in morphology to those of A. scutata and A. calva than to the styliform teeth of A. fragosa. A. uintaensis, A. fragosa, and A. scutata all have a larger infraorbital 4 than infraorbital 5, greater degree of ossification of cranial elements, deeper orbital notch in the frontal, greater head/ standard-length, and generally larger parietal /frontal ratio. These charac- ter-states clearly set the fossil species of Amia apart from the Recent A. calva. Articulated specimens have yielded more information on the osteology of A. fragosa. A. fragosa is a short-bodied form (approxi- mately 65 centra) with a smaller number of caudal lepidotiichia than in the other species of Amia, styliform palatal and coro- noid teeth, deeper orbital excavation in the frontals, square parietals, and a short box- like skull having relatively short mandibles diat occupy about half the head-length. The styliform crushing palatal teetlr of A. fragosa suggest a durophagous habit, rather than the more predaceous habit indicated by the sharp palatal teeth of A. uintaensis, A. scutata, and A. calva. Although it is known that A. calva includes molluscs and crusta- ceans in its diet, perhaps A. fragosa was more exclusively adapted for shell crushing than the Recent species. 76 Bulletiti Museuin of Comparative Zoologtj, Vol. 146, No. 1 Fig. 31. Skull and body structure of A, Amia calva; B, A. scufafa; C, A. uinfaensis; and D, A. fragosa. Fossil Amiids • Boreske 77 PLEISTOCENE PLIOCENE POST-BLANCAN BLANCAN- HEMPHILLIAN CLARENOONIAN FOSSIL LAKE BEOS (lOAHO FM.) WAKEENEY It. (OGALLALA FM.) LOWER VALENTINE FM. BARSTOVIAN MIOCENE HEMINGFORDIAN ARIKAREEAN EUBANKS l.f. (PAWNEE CREEK FM.) TURTLE BUTTE FM. WHITNEYAN OLIGOCENE ORELLAN CHAORONIAN RUBY PAPER SHALE (PASSAMARl FM.) GRANT HORSE PRAIRIE SHALE FLORISSANT FM. ORELLA MEMBER (BRULE FM.) CHADRON FM. CYPRESS HILLS FM. OUCHESNEAN EOCENE UINTAN BRIDGERIAN WASATCHIAN CLARNO FM. HORSEFLY RIVER BEDS UINTA FM. WASHAKIE FM. BRIDGER FM. WIND RIVER FM. FOSSIL LAKE BEOS (GREEN RIVER FM.) 6OLOEN VALLEY FM. WASATCH FM. GRAYBULL BEDS (WILLWOOO FM.) CLARKFORKIAN TIFFANIAN PALEOCENE TORREJONIAN PUERCAN MAASTRICHTIAN CRETACEOUS CAMPANIAN ALBIAN BEAR CREEK l.f, (FORT UNION FM.) SILVER COULEE l.f. (FORT UNION FM.) MELVILLE FM. SAUNDERS CREEK l.f. (PASKAPOO FM.) CEDAR POINT QUARRY l.f. (FORT UNION FM ) MEDICINE ROCKS l.f. (TONGUE RIVER FM.) ROCK BENCH l.f. (FORT UNION FM4 TULLOCK FM. MANTUA If. (FORT UNION FM.) HELL CREEK FM. LANCE FM. OJO ALAMO FM. AGUJA FM. EDMONTON FM. JUDITH RIVER FM. "MESAVERDE" FM. OLDMAN FM. BUTLER FARM l.f. (PALUXY FM) 7X Table 18. Major deposits containing remains of Amia in the WESTERN interior OF THE UnITEU StATES AND CaNAUA 78 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 Seven genera and twenty-three amiid species (Table 19) have been described in the literature. Estes (1964) synonymized Stylomyleoclon lacus with Kindleia fra^osa, and Estes and Berberian ( 1969 ) referred the genus Kindleia to Amia, thereby con- firming the suggestion of Janot (1969). Paramiatus p,iirleyi (Romer and Fryxell, 1928) is unquestionably a synonym of A. fragosa. Regardless of possible synonymy with European taxa, the sti'atigraphic range of A. fragosa is remarkably long, extending as it does from the Late Creta- ceous through the Middle Eocene. Al- though A. fragosa is better known than the other fossil species, and was extensively described by Estes (1964), O'Brien (1969), and Estes and Berberian (1969), its phylo- genetic relationship to them and to A. calva could not be understood without compara- tive information on both the other fossil forms and A. calva (Fig. 32). A. newherrianus and A. depressiis (Marsh, 1871), and A. gracilis (Leidy, 1873a), described from undiagnostic ver- tebral characters, are considered here as nomina duhia. A. iiintaensis is a form having a relatively greater body-length than the other species of Amia. It has approximately the same total number of vertebrae as A. calva and A. scutata, but the arrangement of the coliunn varies meristically from them. Its head is more elongated tlian that of the other forms, with the jaws occupying over two-thirds of the head-length. The vouier- ine teeth are sharp (as are the palatal and coronoid teeth), as they are in A. scutata and A. calva, but are more than twice as numerous as in these later forms. The pres- ent study confirms the opinions of Romer and Fryxell ( 1928), Estes ( 1964), and Estes and Berberian (1969) that the differences between Amia and Protamia, Hypamia, and Pappichthys are insufficient for the recogni- tion of any of the latter as genera distinct from Amia. Hypamia elegans (Leidy, 1873a) is considered a m)men duhium, be- ing based on vertebral characters that can- not be distinguished from those of the other species. Protamia media (Leidy, 1873a), Pappichthys symphysis, P. corsonii, P. medius, P. plicatus, P. sclerops, P. laevis (all described by Cope, 1873), as well as Atnia macrospondyla and A. whiteavesiana (Cope, 1891), are all considered here as synonyms of A. uintaensis; they were based on undiagnostic vertebral characters and morphology of the skull elements. Material of large amiids from the Late Cretaceous Lance and Hell Creek formations is referred to A. cf. uintaensis, since the material differs only in minor respects from the Paleocene and Eocene specimens. It cannot be deter- mined whether this material represents ac- tual populations of A. uintaensis or an earlier stage of its evolution. The strati- graphic range of A. uintaensis extends from the Paleocene to the Early Oligocene. A. scutata, an Early to Middle Oligocene long-bodied form, shares cranial characters with both A. uintaensis and A. calva. Al- though it has closer morphometric and meristic affinities to the Recent form, it is structually and temporally intermediate be- tween A. uintaensis and A. calva; it resem- bles the more primitive A. uintaensis in the moi-phology of Meckel's groove and coro- noid articulation surface of the dentary, greater ossification, and in having an elon- gated skull with a greater head/ standard- length than in A. calva. A. dictyocephala (Cope, 1875) is considered a synonym of A. scutata; it was based on undiagnostic meristic characters. In the evolutionarv con- tinuum, A. scutata appears to be an inter- mediate stage between A. uintaensis and A. calva (Fig. 32). A more direct line of evolution exists between A. scutata and A. calva; this is supported by Miocene and Pliocene amiid material that displays cra- nial elements closely transitional between the two species. Thus the Recent species of A. calva had begun at least by the begin- ning of the Pliocene, and A. calva was ap- parently distinct from A. scutata by that time. It appears that A. fragosa represents an amiid population that survived until the Middle or Late Eocene and had no phylo- genetic affinities with the modern form be- yond this time. I Fossil Amiids • Boreske 79 /Im/'o colva RECENT PLEISTOCENE POST-BLANCAN Rl ANT AM HEMPHILLIAN PLIOCENE CLARENDONIAN Ami a cf. calva BARSTOVIAN A mi a cf . scut at a MIOCENE HEMINGFORDIAN ARIKAREEAN WHITNEYAN OLIGOCENE ORELLAN CHADRONIAN Ami a scuta ta - i DUCHESNEAN , J EOCENE UINTAN BRIDGERIAN \ y WASATCHIAN ^ragosa Amia uintaensis CLARKFORKIAN 'V" / PALEOCENE TIFFANIAN TORREJONIAN PUERCAN \ \ / - MAASTRICHTIAN \ Amia ct uintaensis CRETACEOUS CAMPANIAN \ ; ALBIAN Al niidae mcertae sedi \ s Fig. 32. Suggested phylogenetic relationships within the genus Am\a. 80 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 In the North American fossil record, fossil remains unquestionably those of the family Amiidae first occur in tlic Lower Cretaceous (Albian) sediments of Texas. However, none of the material can be referred to any known species of Araia; it displays charac- ter-states resembling those of Amia uintaen- sis and Amia frcifi^osa, as well as the Euro- pean Urocles. Some of the vertebrae re- semble those of Amiopsis. The Paluxy mate- rial may represent either one or more forms transitional between Amia and the Late Mesozoic European Urocles (or Amiopsis) , or an as yet undescribed line. The body- length of Amia fra^osa appears to be a primitive feature derived from the earlier amiids Urocles, Siruimia, Ikechaoam,ia, and Amiopsis. Despite thc>ir different vertebral columns, Amia frafi^osa and A. uintaensis show similar morphology of the cranial elements, but the nature of the probable common origin of these forms is still uncer- tain in the absence of a more complete fossil record. Remains of amiids referable to or close to Amia fragosa and A. uintaensis have been described from the Paleocene, Eocene, and Oligocene of Europe, and the Eocene of Asia. Additional but still not definitive evi- dence supports Estes' ( 1964 ) and Estes and Berberian's (1969) suggested synonymy of A. russelU (Late Paleocene, France), A. kehreri (Middle Eocene, Germany), and A. munieri (Early Oligocene, France) with A. fragosa. Pseudamia Jieintzi (Eocene, Spitzbergen ) and A. valenciennesi ( Eocene, France) are also possible synonyms of A. fragosa. A. valenciennesi is the oldest name and would take precedence over A. fragosa. Cranial similarities confirm the synonymy of A. rohusta (Late Paleocene, France) with A. uintaensis. European and North American fossil Amia occurred in freshwater deposits and apparently occupied a habitat much like that of the Recent species. According to Westoll (1965: 19-20) the distribution of freshwater vertebrates is a useful indica- tion of "direct continental communication," Table 19. Amiid genera and species of various AUTHORS discussed IN TEXT IN RELATION TO THE REVISED lAXONOMY Atnia calva Kindlcia fragosa Stijlomijleodon lactis Amia fragosa Paramiattts f^tirlcyi Amia scutata A7nia dictyocepliala Aitiia cxilis Protamia uintaensis Protamia media PappicJi th ys m edius PappicJithys pJicatus Pappichthys sclerops Pappichthys lacvis PappicJithys symphysis PappicJithys corsonii Atnia loJiiteavcsiana Amia macrospondyla Amia depressus Amia newJwrriamis Amia gracilis Hypamia elcf^ans Arnia sehvyniana . . Amia calva Amia fragosa .Amia sctitata Amia uintaensis nomina dubia .nomen nudum since ". . . descendents of a common stock on different modern continents must have used essentially a terrestial route." The present study further amplifies similarities in the Paleocene and Early Eocene amiid fossil record of North America and Europe. This distribution of amiids adds to the similarity of assemblages of Paleocene and Early Eocene lower vertebrates (Estes et al., 1967) and mammals (McKenna, 1972) on the two continents. The occurrence of Pseudamia Iwintzi in the Eocene deposits of Spitzbergen may be additional evidence for the existence of the De Geer migration route (bridging Europe, Spitzbergen, and North America during the Paleocene and until the close of Sparnacian time), espe- cially if suggested relationship to A. fragosa could be demonstrated. The Asian form A. mongoliensis resembles A. uintaensis in minor respects but is sufficiently distinct in itself to be maintained as a separate species. Fossil Amiids • Boreske 8i LITERATURE CITED Agassiz, L. 1843. Recherches sur les poissons fossiles. Neuchatel, tomes 1-5, atlases 1-5. Allis, E. 1889. The anatomy and development of the lateral line system of Amia calva. J. Morphol., 2: 1-540. . 1897. The cranial muscles and cranial and first spinal nerves in Amia calva. J. Mor- phol., 12(3): 1-814. Ami, H. 1891. On some extinct Vertebrata from the Miocene rocks of the north-west Territories of Canada recently described by Professor Cope. Science, 18: 53. Andreae, a. 1892. Vorliiufige Mitteilung iiber die Ganoiden ( Lepidostetis und Amia) des Mainzer Beckens. Verb. Nat. med. Ver. Hei- delberg (N.F.), 5: 7-15. . 1895. Beitriige zur Kenntniss der fos- silen Fische des Mainzer Beckens. Abh. Senck. naturf. Ges., 18: 351-365. Becker, H. 1961. Oligocene plants from the Upper Ruby River Basin, southwestern Mon- tana. Mem. Geol. Soc. Amer., 82: 1-127. . 1962. Two new species of Mahonia from the Grant-Horse Prairie Basin in south- western Montana. Bull. Torrey Bot. Club, 89: 114-117. Blair, W., A. Blair, P. Brodkorb, F. Cagle, AND G. Moore. 1968. Vertebrates of the United States. New York: McGraw-Hill Press, 450 pp. BoLK, L., E. Goppert, E. Kallius, and W. Lu- BOSCH. 1936. Handbuch der vergleichen- den Anatomie der Wirbeltiere 4. Berlin: Ur- ban und Schwarzenberg, 1116 pp. Boreske, J. 1972. Ta.xonomy and taphonomy of the North American fossil amiid fishes. (Abstr.) Bull. Geol. Soc. Amer., 4(1): 3-4. Bridge, T. 1877. The cranial osteology of Amia calva. J. Anat. Phys., 11: 605-622. Cartier, D., and E. Magnin. 1967. La crois- sance en longueur et en poids des Atnia calva L. de la region de Montreal. Canad. J. Zool., 45: 797-804. Cavender, T. 1968. Freshwater fish remains from the Clarno Formation Ochoco Moun- tains of north-central Oregon. Ore Bin, 30 (7): 125-141. . 1970. A new find of Amia dictyoce- phala Cope from the Middle Tertiary Floris- sant Lake Beds of western North America. Abst. paper given at 50th Annual Meeting, Amer. Soc. Ichth. Herp.: 42. Cope, E. 1873. On the extinct Vertebrata of the Eocene of Wyoming, observed by the ex- pedition of 1872, with notes on the geology. Ann. Rept. U.S. Geol. Surv. Terr. (1st series), No. 6: 545-649. . 1875. On the fishes of the Tertiary shales of South Park. Bull. U.S. Geol. Surv. Terr. (2nd series). No. 1: 3-5. . 1884. The Vertebrata of the Tertiary Formations of the West. Rept. U.S. Geol. Surv. Terr. (Hayden), 3: 745-746. . 1891. On Vertebrata from the Tertiary and Cretaceous rocks of the northwest Terri- tory. Book 1. The species from the Oligocene or Lower Miocene of Cypress Hills. Contrib. Geol. Surv. Canad. Paleo., 1: 2-4. Dean, B. 1898. On the dogfi.sh (Amia calva), its habits and breeding. Rept. Comm. Fish- eries, Game, and Forests New York, No. 4: 1-10. Dechaseaux, C. 1937. Le genre Amia, son his- toire paleontologique. Ann. Paleont., 26: .3- 16. Eastman, C. 1899. Jurassic fishes from the Black Hills of South Dakota. Bull. Geol. Soc. Amer., 10: 397-407. Estes, R. 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyo- ming. Univ. Calif. Publ. Geol. Sci., 49: 1- 180. , AND P. Berberian. 1969. Amia ( = Kimlleia) fragosa (Jordan), a Cretaceous amiid fish, with notes on related European forms. Breviora, Mus. Comp. Zool., No. 329: 1-14. , , AND C. Meszoely. 1969. Lower vertebrates from the Late Cretaceous Hell Creek Formation, McCone County, Montana. Breviora, Mus. Comp. Zool., No. 337: 1-33. , M. HeCHT, AND R. HOFFSTETTER. 1967. Paleocene amphibians from Cernay, France. Amer. Mus. Novitates, No. 2295: 1-25. -, AND J. TiHEN. 1964. Lower vertebrates from the Valentine Formation of Nebraska. Amer. Midi. Nat., 72(2): 453-472. Goodrich, E. 1930. Studies on the Structure and Development of Vertebrates. London: Macmillan Co. Ltd., xxx + 837 pp. Gould, S. 1966. Allometry and size in ontog- eny and phylogeny. Biol. Reviews, 41: 587- 640. Ha.mmett, F., AND R. Hammett. 1939. Pro- portional length growth of gar ( Lepisosteus platyrhincits DeKay). Growth, 3(2): 197- 209. Hasse, C. 1882. Das natiirliche System der Elasmobranchier. Jena: Verlag von Gu.stav Fischer, pp. 77-284. Hatcher, J. 1900. The Carnegie Mu.seum pale- ontological expeditions of 1900. Science, 12 (.306): 718-720. . 1901. Some new and little known fos- sil vertebrates. Ann. Carnegie Mus. Pitts- burgh, No. 1: 128-129. Hay, O. 1895. On the structure and develop- 82 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 ment of the vertebral column of Amia. Publ. Field Columb. Mus. Zool., 1(1): 1-54. . 1911. Tlie Pleistocene of Indiana. Ann. Kept. Ceol. Nat. Res. Indiana, .36: 552. . 1917. Vertebrata mostly from stratum No. 3, at Vero, Florida; together with de- scriptions of new species. Rept. Geol. Surv. Florida, 9: 43-68. . 1923. The Pleistocene of North Amer- ica and its vertebrated animals from the states east of the Mississippi River and from the Canadian Provinces east of longitude 95 de- grees. Bull. Carnegie Inst., .322: 336-382. HoFFSTETTER, R., AND J. Gasc. 1969. Verte- brae and ribs of modem reptiles. In Biology of the Reptilia. New York: Academic Press, xvi -|- 468 pp. HuBBS, C, AND K. Lagler 1967. Fishes of the Great Lakes Region. Ann Arbor: University of Michigan Press, viii -|- 289 pp. HussAKOF, L. 1932. The fossil fishes collected by the Central Asiatic Expeditions. Amer. Mus. Novitates, No. 553: 1-16. Imbrie, J. 1956. Biometrical methods in the study of invertebrate fossils. Bull. Amer. Mus. Nat. Hist., 108: 217-252. Janot, C. 1966. Aiuia riisselli nov. sp., nouvel Amiide ( Poisson holosteen ) du Thanetien de Berru, pres de Reims. C. R. Soc. Geol. France, 1966(3): 142. . 1967. A propos des amiides actuels et fossiles. Colloq. Intern. C. N. R. S., 163: 139-153. Jordan, D. 1906. The fishes of Samoa. Bull. Bureau Fisheries, 25: 2.37. . 1919. The genera of fishes. Publ. Le- land Stanford Junior University, 1919: 259. . 1925. Opinion 89. In Opinions ren- dered by the International Commission on Zoological Nomenclature. Smithsonian Misc. Coll., 73(3): 27-29. . 1927. Kindleia, a new genus of cichlid fishes from the Upper Cretaceous of Alberta. Canad. Field Nat., 41: 145-147. . 1928. Note on Kindleia and Stylomy- leodon ( fossil fish from Cretaceous deposits of Alberta). Canad. Field Nat., 42: 47. , and B. Evermann. 1896. The fishes of North and Middle America. Bull. U.S. Nat. Mus., 47: 112-113. Lagler, K., J. Bardach, and R. Miller. 1962. Ichthyology. New York: John Wiley and Sons, Inc., vii + 523 pp. Lambe, L. 1904. Progress in vertebrate pale- ontology in Canada. Trans. Roy. Soc. Canada, No. 2: 27-43. . 1908. The Vertebrata of the Oligocene of the Cypress Hills, Saskatchewan. Contrib. Paleo. Canada, 3(4): 12-13. Lange, S. 1968. Zur Morphologic imd Tax- onomie der Fischgattung Urocles aus Jura nnd Kreide Europas. Palaeontographica, Abt. A, 131: 1-78. Lehman, J. 1951. Un nouvel Amiide de I'Eocene du Spitzberg, Pseudamia lieintzi. Troms0 Mus. Arshefter, 70: 1-11. Leidy, J. 1873a. Notice of remains of fishes in the Bridger Tertiary Formation of Wyoming. Proc. Acad. Nat. Sci. Philadelphia, 1873: 97-99. . 1873b. Contributions to the extinct vertebrate fauna of the western territories. Rept. U.S. Geol. Surv. Terr. (Hayden), 1: 185-189. Liu, T-S., H-T Liu, and T-T Su. 1963. The discovery of Sinamia zdanskyi from the Ordos Region and its stratigraphical significance. Vert. Palasiatica, 7(1): 1-30. Lund, R. 1967. An analysis of the propulsive mechanisms of fishes, with reference to some fossil actinopterygians. Ann. Carnegie Mus., 39(15): 195-218. Marsh, O. 1871. Communication on some new reptiles and fishes from the Cretaceous and Tertiary. Proc. Acad. Nat. Sci. Philadelphia, 1871: 103-105. McKenna, M. 1972. Was Europe connected directly to North America prior to the Mid- dle Eocene? In Evolutionary Biology, Vol. 6. New York: Appleton-Century-Crofts, pp. 179— 189. Merrill, G. 1907. Catalogue of the figured specimens of fossils, minerals, and ores in the Department of Geology, LTnited States Na- tional Museum. Bull. U.S. Nat. Mus., 53: 6-15. Miller, H. 1968. Additions to the Upper Cre- taceous vertebrate fauna of Phoebus Land- ing, North Carolina. J. Elisha Mitchell Sci. Soc, 84: 467^71. Newton, E. 1899. On the remains of Amia, from the Oligocene strata in the Isle of Wight. Quart. J. Geol. Soc. London, 4: 1- 10. Nybelin, O. 1963. Zur Morphologic und Ter- minologie des Schwanzskelettes der Actinop- teiygier. Arkiv. Zool, 15(35): 485-516. O'Brien, D. 1969. Osteology of Kindleia frag- osa Jordan (Holostei: Amiidae) from the Edmonton Formation ( Maestrichtian ) of Alberta. M.A. Thesis, University of Alberta: vi -1- 118 pp. Osborn, H., W. Scott, and F. Speir. 1878. Paleontological report of the Princeton scien- tific expedition of 1877. Contrib. Mus. Geol. Arch. Princeton, No. 1: 102-104. Piton, L. 1940. Paleontologie du gisement Eocene de Menat ( Puy-de-Dome, flore et faune). Mem. Soc. Sci. Nancy, 56: 1-303 pp. Reighard, J. 1903. The natural history of Amia Fossil A muds • Bote she 83 ralva Linnaeus. Mark Anni\cisaiy Vohinie (art. t): 57-109. RoMKH, A., AND F. FiiYXKLi,. 1 92S. Purdiuiatiis ^urlcyi, a ni-w (Iccp-hocliccl aiiiiid fish from tlie Eocene of Wyoniinj,'. Aiiiei. |. Sei., 16 (90): 519-527. |{us.st:LL, L. 1928a. A new fossil fish from the Paskapoo Beds of Alberta. Aiiier. |. Sei., 15 (86): 103-107. . 19281). The jfenera Kiiullcut and Sty- loviylcodon. Anier. J. Sei., 15(87): 264. . 1929. Tiie validity of the .ueniis Sttj- IniiujUoiUm. Auier. J. Sei.. 17( 100): 369-;571. . 1967. Paleontology of the Swan Hills area, north-central Alheila. Roy. Ontario Mus. C.'ontril)., No. 71: 1-31. Sc:hakffeh, B. 1967. Osteichthyan vertebrae. |. Linn. Soe. (Zool.), No. 47: 185-195. Shufixut, R. 1885. The osteology of Ainiu calva. Washington: Government Printing Of- fice, 93 pp. SiMi'.soN, (;. 1937. The Fort Ihiion of the Crazy Mountain Field, Montana, and its inannnalian faunas. Bull. U.S. Nat. Mus., 169: \ + 287 pp. , A. RoK, AND R. Lewontin. 1960. Quantitative Zoology. New York: Harcourt, Brace, and Co., iv + 440 pp. Skinneh, M., S. Skinneh, and R. Gooius. 1968. C-enozoic rocks and faunas of Turtle Butte, south-central South Dakota. Bull. Anier. Mus. Nat. Hist., 138: 381-436. Smi'iii, C. 1962. Some Pliocene fishes from Kansas, Oklahoma, and Nebraska. Copeia, 1962(3): 505-520. Stensio, E. 1935. Smainia zdanskyi, a new amiid from the Lower C^retaceoiis of Shan- tung, China. Pahoiit. Sinica, .ser. C, No. 3: 1-48. Swift, C. 1968. Fossil bony fishes from Flor- ida. I'laster Jacket, No. 7: 2-11. SzALAY, F., AND M. Mc:Kenna. 1971. Begin- ning of the age of uiaiumals in Asia: tin- Late Paleocene Cashato fauna, Mongolia. Bvill. Amer. Mus. Nat. Hist., 144: 270-317. Thomson, K., and K. IIaiin. 1968. (Jrowth ;nid form in fossil rhipidistian fishes ( Cro.ssopter- ygii). J. Zool., 156: 199-223. Thuhmond, J. 1969. Lower vertebrates and paleontology of the Trinity Croup. Ph.D. Thesis, Southern Methodist Unixersity ( Libr. Congr. No. 70-19259). 1.36 pp. Univ. micro. Ann Arbor, Mich, (di.ss. ab.str. 31: 2156). TnAQUAut, R. 1911. Les poissons wealdiens de Bernissart. Mem. Mus. Roy. Hist. Nat. Belg., 6: 1-65. Westoll, T. 1965. ecological evidence bear- ing upon continental drift. Symposium on Continental Drift. Royal Soc. London, 1965: 12-25. WiUTEHousE, R. 1910. The caudal fin of the Teleostomi. Proc. Zool. Soc. London, 1910: 590-627. Whitley, C. 1954. More new fish names and records. Aust. Zool., 12: 57. Wilson, R. 1968. Systematics and faunal anal- ysis of a Lower Pliocene vertebrate assem- blage from Trego County, Kansas. Contril). Mus. Paleo. Univ. Mich., 22(7): 75-126. Woodward, A. 1916. The fossil fi.shes of the English Wealden and Purbeck formations. Palaeontogr. Soc. London, Pt. 1, 1915: 1-48. 84 Bulletin Museum of Comparative Zoology, Vol. 146, No. 1 5 cm afab. B .J; m .: '-^ -■ 1 y •^fe%i ' • ■•■-Tf^ ^ •'i. - . i Plate 4. Amio scu/ofa. Middle Oligocene, Florissant Formation, Colorado: A, counterpart YPM 6243; B, counter- part USNM 4087; C, PU 10172. I i us ISSN 0O27.4100 Bulletin OF THE Museum of Comparative Zoology An Analysis of Variation in the Hispaniolan Giant Anole, Anolis ricordi Dumeril and Bibron ALBERT SCHWARTZ HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS, U.S.A. VOLUME 146, NUMBER 2 19 APRIL 1974 PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Bulletin 1863- Breviora 1952- Memoirs 1864-1938 JoHNSONLV, Department of MoUusks, 1941- OccAsioNAL Papers on Mollusks, 1945- Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- malian Hibernation. Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae (MoUusca: Bivalvia). Whittington, H. B., and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolu- tion of Crustacea. Proceedings of the New England Zoological Club 1899-1948. (Complete sets only.) Publications of the Boston Society of Natural History. Authors preparing manuscripts for the Bulletin of the Museum of Comparative Zoology or Breviora should send for the current Information and Instruction Sheet, available from Editor, Publications OflBce, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. © The President and Fellows of Harvard College 1974 AN ANALYSIS OF VARIATION IN THE HISPANIOLAN GIANT ANOLE, ANOLIS RICORDI DUMERIL AND BIBRON ALBERT SCHWARTZ' Abstract. The nominal Hispaniolan species of giant anole, Anolls ricordi, is considered to be in actuality composed of three distinct allopatric spe- cies: A. ricordi, A. barahonae, and A. baleatus. Subspecies of all three species are described, but only A. baleatus is well represented in collections. A theoretical history of this species complex upon Hispaniola is presented. The Hispaniolan giant anole, Anolis ri- cordi Dumeril and Bibron, 1837, has been known to science for more than a century; yet only in the hist 35 years has it become evident that this species is not homoge- neous in its characteristics throughout Haiti and tlie Republica Dominicana. The spe- cies was first named (as Anolis ricordii) from Santo Domingo, as the entire island was known at that historical period, but specimens seem to have been rare in col- lections thereafter. Schmidt (1921: 10) re- ported four A. ricordi from two Dominican localities. Cochran (1941: 133) Hsted 24 specimens (all but one of which were in the National Museum of Natural History) from 11 localities. Mertens (1939: 68-70) studied 17 specimens in European collec- tions and was the first to recognize that there were two readily distinguishable pop- ulations that he considered subspecies: A. r. ricordi in Haiti, and A. r. baleatus Cope in the Republica Dominicana. Williams ( 1962 ) reviewed the species in more detail and examined 90 specimens. For this suite of anoles, he described A. r. barahonae \ 1 Miami-Dade Community College, Miami, Florida 33167. Bull. Mus. Co from tlie Sierra de Baoruco in the south- western Republica Dominicana. Still later, Williams (1965) studied an additional 80 specimens and named A. r. leberi from Camp Perrin on the extreme distal portion of the Haitian Tiburon Peninsula. Thus, with increasing quantities of material from more diverse localities, our knowledge of the distribution and variation in this species has increased accordingly. A great many problems remain, however, when one deals in detail with the variation in A. ricordi. Williams (1962, 1965) pointed out that records of the species were of such a scattered nature (especially on the Tiburon Peninsula but also elsewhere on the island) that intergrades between several of the subspecies remained unknown and also that there were no specimens available from large areas between named populations. Williams and Rand (1969), in their excellent summary of the geographic differentiation in all species of Hispaniolan anoles, pointed out (p. 15) that Anolis ri- cordi was composed of "several described subspecies, some of which are sharply enough distinct to raise the question of pos- sible species status." This is most especially true of the taxa ricordi, baleatus, and bara- honae, all of which are extremely well characterized by both pigmental and struc- tural details, but all of which occupy areas (extensive in the cases of ricordi and ba- leatus) without known intergradation be- tween them or without close geographic approximation. Thus, the closest ap- mp. Zool., 146(2): 89-146, April, 1974 89 90 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2 o CM I 00 i IZ k. ■s. u o 0) lA 3 1 .a o D5 o 01 0 w» C u 01 •~ C c o -c 0 i. 0 ■< 'u 01 a Q. 0 0 II (A o -J 3 CO 0 w o> k. M •V lo (V 0) "o 3 3 -o c c o ^ < _D E ., .^ > ~z 1 o V* u J) O u k! ^ E 0 d) 'C ->t "5 IB vt 0 35 II -Q tA 1 1 ^ 0 1 > < 5 c 01 vt 01 ^ Lk oe II II a o II VI 01 o II < 3 ^- O a V) 0 f* c 0 0 c o -c 0 w ■♦- "a E E VI $ 0 "o _o < J3 0) "o 0 ■a c "a a w 0 >. < Q. '■*- 0) > ^ =: O M 0 1 1 u D II (U 0 c ■D c tA '> CQ l^ 01 a ^ 01 0 c 0 c ^ -o o c 0 "o c o 05 E c _0 D O) O E ^ tA a 0 VI T3 'c 0 Q. 0 -a c 01 c 01 c 1 1 0) c D D _3 c o 1 k. 0 3 3 VI -C -C 01 0 *- *• "3 o -o 0 0) (J '5 T3 3 0 ■a c o o kj 0 O w 01 w» < u> ^ 3 J) _Q c ^ 1 >- ^ c 1 "a 0 tn *L» (11 < < <1) 1. o o tA 01 c 0) 0 c j: lA 0 c c 0) c 0) c ^wi 0 0 u J ^ _o «/) 0 II — o _c II o -^ ^ tn ^ u 0£ t in E 0 "o 0) 01 o 0 1 X > C 1 'E 0 > 0 c 01 a 0 O "5 E •A X -SI D C "o k- -0 1 c o 0) «r **- o 1 VI 3 o Q. lA 2 O "o c VI -a 0 -Q ,^ k-' 0 u tA k- _o c iA Ji 0 < 3 1 01 D O) 0 II 0 3 1 3 01 c ristically harahoiuie in squamation." The specimen (AMNH 51230) from half- way between Enriquillo and Oviedo is a young male with a snout-vent length of 121. Since this lizard presumably came from the lowlands of the Peninsula de Barahona, it might logically be expected to be alhocellatiis. However, the lizard is now 118 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2 drab patternless brown, and there are no indications that it was ever spotted. Pre- sumably albocellatus and harahonae inter- grade between Enriqiiillo (which Hes at the extreme southeastern corner of the Si- erra de Baoruco) and Oviedo (which Hes well down on the Peninsula de Barahona). Several facts have prompted my naming this lonely specimen. First, I have exam- ined the Enriquillo specimen noted by Wil- liams, and, although it shows some indica- tion of vertical crossbars, they are not any more conspicuous than those in some more recently taken A. /;. harahoruie from the Baoruco highlands (Williams examined only 17 harahoime at the time of its original description; I have studied almost twice this number). Secondly, the xeric to semi- arid region south of the Sierra de Baoruco has come to be known as an area of local differentiation at the subspecific level for a variety of reptiles; this alone is no reason for naming albocellatus, of course. Thirdly, although since 1964 when the holotype was collected both I and others have spent con- siderable time on the Peninsula de Bara- hona and in the vicinity of Oviedo, we have never seen or secured another A. harahonae in this region. In September 1966, the very severe hurricane Inez passed directly across the Peninsula. What had once been high- canopied semi-arid forest (as at Oviedo) has been either totally destroyed or been reduced (by 1969) to a landscape of bare snags with some leafy growth just now be- ginning to appear but at a much lower canopy-level than previously. The changes between the Oviedo area in 1964 and 1969 are so massive that, upon my first visit there after Inez, I was unable to orient myself in reference to our older collecting localities! Certainly this entire region has suffered greatly, and, with the destruction of trees, it seems reasonable to assume that A. hara- honae has suffered equally. The population may never have been high, since such semi- arid woods are not at all optimal habitat for any of the Hispaniolan giant anoles, and the destruction of the habitat must surely have affected A. h. albocellatus adversely. Since persistent visits to this area have yielded no new material, and since the liz- ard may presently be very rare, I have de- cided upon the present course rather than wait in hope for someone to secure a sec- ond ( or more ) lizard. Remarks. The Peninsula de Barahona has been shown to have distinctive subspe- cies (or even species) of a variety of rep- tiles. Species that have described endemic subspecies south of the Sierra de Baoruco include: Sphaerodacttjhis difficilis Barbour, Leiocephalus Imrahonensis Schmidt, Am- eiva chrysolaema Cope, Ameiva lineolata Dumeril and Bibron, Arnphisbaena gona- vensis Cans and Alexander, and Dromicus parvifrons Jan. Endemic Peninsula de Barahona species are: Anolis longitibialis Noble, Typhlops sijntherus Thomas, Lep- totyphlops pyrites Thomas, and Uromacer ivetmorei Cochran. Only one amphibian, Eleutherodactylus alcoae Schwartz, is re- stricted to the Peninsula. To the former list can now be added Anolis harahonae. The eastern half of the Peninsula, although xeric, was originally clothed in dry forest, much of it upon a series of limestone ter- races, the highest point of which is the Loma Gran Sabana, having an elevation of 1082 meters in the north and descending to Cerro Caballo, and Loma de Chendo, hav- ing elevations of 322 and 233 meters, re- spectively, to the south. West of this ridge, the land descends abruptly to Acacia-cac- tus desert to the east of Cabo Rojo, and this habitat continues to the Dominico- Haitian border at Pedernales. Presumably, A. b. albocellatus occurs throughout the eastern half of the Peninsula in the for- merly high-canopied forests of the lime- stone terraces. The holotype was secured by Richard Thomas during the day in a viny tangle in semi-xeric woods near Oviedo; the lizard was in an edge situation, since beyond the dense vine tangle the woods thinned to more scrubby and cleared areas. The name albocellatus is from the Latin HisPANioLAN Giant Anole • Schivartz 119 "albus" for "white" and "ocellus" for "eye," in allusion to the white spots that are typi- cal of the holotype. Anolis baleatus Cope Eti])ristis baleatus Cope, 1864, Pvoc. Acad. Nat. Sci. Philadelphia, p. 168. Type locality. Santo Domingo; holo- type, British Museum (Natural History) 1946.8.29.22. Definition. A giant species of Hispanio- lan Anolis characterized by the combina- tion of large size (males to 1/Iaria Trinidad Sanchez Province, Repub- lica Dominicana, one of a series collected by native collectors on 28 November 1971. Original number ASFS V34486. Paratypes. CM 54119-26, MCZ 125628- 33, ASFS V34502-13, same data as holo- type; AMNH 6017, Villa Riva, Duarte Province, Republica Dominicana, C. R. Halter, May-July 1915. Associated specimens. REPCBLICA DOMINICANA: Duarte Province, Los Bracitos (AMNH 41465-66); ca. 4 km NE Ponton (Rio Cuaba) (ASFS V2987); San- chez Ramirez Province, 1 km SE La Mata (ASFS V33650-51); La Vega Province, 12.8 km NW Bonao, 1200 feet (366 meters) (ASFS V4317); 71 km NW Santo Domingo (= near La Cumbre) (MCZ 128369); San Cristohal Province, 5.0 mi. (8.0 km) NE Gonzalo, 1000 feet (305 meters) (ASFS V29420-21). Definition. A subspecies of A. haleatus characterized by the combination of mod- HisPANioLAN Giant Anole • Schwartz 127 ally 4 scales between second canthal scales, 8 vertical rows of loreal scales, 3 scales be- tween the supraorbital semicircles, 5/5 scales between the interparietal and the supraorbital semicircles, moderate number of vertical dorsal scales (14-22; mean 17.1), moderate number of ventral scales (15-32; mean 22.4), nuchal crest scales very high to high (usually) to moderate or even low (rarely) in both sexes, body crest scales extremely variable, modally moderate in both sexes, but with some oc- cinrences of high and many occurrences of low body crest scales, subocular scales almost always separated from supralabial scales by one row of scales, both sexes some shade of green (usually dark) with foiu- pale green crossbars and with bright sky-bhie blotches along the junction of the green dorsal color and the paler venter (less prominent in females than in males), dewlap in males pale yellow to orange, in females pale yellow to orange but with much dark brown to grayish streaking or smudging, throat in males deep yellow-or- ange and immaculate or with very faint greenish dots, in females yellow-green to bright yellow, always with some darker green dots, rarely marbled with dark green, but never streaked with that color. Distribution. Northeastern Republica Dominicana, from Duarte, Sanchez Rami- rez, La Vega, and northern and eastern San Cristobal provinces, to the base of the Peninsula de Samana (Caiio Abajo); in- tergrades with the subspecies to the south and east in the region of El Seibo Province. Description of holotijpe. An adult male with a snout-vent length of 137 and a tail length of 250; snout scales between second canthals 4, 7 vertical rows of loreal scales, 3 scales between the supraorbital semicir- cles, 6/6 scales between the interparietal and supraorbital semicircles, vertical dor- sals 16, horizontal dorsals 23, ventrals 26, one row of scales between the suboculars and supralabials, fourth toe lamellae on phalanges II and III 30, nuchal crest scales very high, body crest scales moderate; in life, dorsum dark green with four pale green crossbars, the dark green color blending fjuickly at the junction of the dor- sal and ventral color into a series of diag- onally directed sky-blue areas that give a ragged appearance to the jmiction of the dorsal and ventral colors; dorsal crossbands continue onto the tail; cascjue gray-green, eyeskin pale pea-green; dewlap pale yel- low-orange, chin slightly deeper yellow-or- ange, throat yellow-orange, immaculate except for some vague pale greenish smudges posterolaterally. Variation. The series of A. b. caeruleo- lattis consists of 20 males and 17 females. The largest male (ASFS V34505) has a snout-vent length of 148, the largest fe- male (AMNh' 6017) 145. The male is from the type locality, the female from Villa Riva. Snout scales at the level of the second canthals range between 2 and 5; the mode is 4 ( 14 specimens ) . The verti- cal loreal rows vary between 6 and 10; the mode is 8 ( 15 specimens ) . There are 2 or 3 scales between the supraorbital semicir- cles (mode 3). There are modally 5 scales between the interparietal and the supraor- bital semicircles; 5 scales are involved in 52 percent of the combinations; actual counts are 4/4 (3), 4/5 (6), 5/5 (10), 5/6 (7), 6/6 (4), 6/7 (1), 4/6 (1), and 5/7 ( 1 ) . Vertical dorsals range between 14 and 22 (mean 17.1), horizontal dorsals be- tween 15 and 25 ( 19.9 ) , and ventrals be- tween 15 and 32 ( 22.4 ) . Of 16 adult males, four have the nuchal crest scales very high, 11 have them high, and one has them mod- erate. Of 17 adult females, four have the nuchal crest scales very high, ten have them high, and three have them moderate. In the adult males, the body crest scales are high in six males, moderate in eight, and low in two, whereas in the adult fe- males, these scales are high in five, mod- erate in six, and low in six. All but two lizards (6 percent) have the suboculars separated by one row of scales from the supralabials. In a series of 12 adult male topotypes, the dorsal ground color was recorded as some shade of green (usually dark green) 128 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2 with four pale pea-green crossbands. The patterned hke adults except that the sky- dorsal green color blends quickly ventro- blue lower edges to the dorsal color were laterally into a series of irregular sky-blue absent and the dewlap was streaked brown patches or blotches that mark the border and gray basally. The chin and throat between the dorsal green and the pale yel- were immaculate pale green. There are no low to cream venter. These sky-blue color data on the other juveniles, and none patches are often prominently extended of them presently shows any pattern, onto the lateral margins of the venter as a Comparisons. A. h. caeruleolatus dif- series of diagonal, posteriorly directed fers from all previously described subspe- areas, which, upon preservation, are still cies in».having the sky-blug patching along prominent features of the lower sides. The the lower sides. In having four dorsal pale upper surface of the head was gray-green green body bands, caeruleolatus differs to brown, the eyeskin pale pea-green. The strikingly from multistruppus with its mul- dorsal banded pattern of dark and light tiple banding; in addition, the dewlap of green continues onto the tail. The dewlap multistruppus is pale and often grayish, in is pale yellow-orange, yellow, or orange, contrast to the generally brighter dewlaps and the chin is slightly deeper yellow-or- of caeruleolatus. From nominate baleatus, ange, concolor with the throat, which is caeruleolatus differs in having the throat either immaculate (usually) or with very yellow to yellow-green rather than bright faint greenish dots or smudges. Eleven fe- yellow to orange, and female caeruleolatus male topotypes were colored and patterned have the throat with dark green markings, dorsally like the males, with the pattern From high upland sublitnis, caeruleolatus extending onto the tail, but there is only a differs in having the sky-blue blotches ven- vague indication of the ventrolateral sky- trolaterally and in lacking ventral mark- blue pigmentation. The necks of females ings, and whereas caeruleolatus has com- were often streaked with dark and pale parably pigmented dewlaps^ those in greens. The chin and throat were yellow sublimis are generally paler and often suf- to yellow-green, regularly with some fus^d at least basally with gray. The dor- darker green dots, blotches, or occasionally sal patterns of both sublimis and caeruleo- marbled with dark green. The female dew- latus are comparable, since both are lap was yellow to pale orange, streaked crossbanded. with dark brown or grayish. As far as meristic counts are concerned, Two females from the haitises region caeruleolatus differs from the named sub- near Gonzalo were deep to emerald green species in the following ways. Compared in life with yellow dewlaps having varying with baleatus, caeruleolatus has modally 8 amounts of brown streaking or smudging; (rather than 7) vertical loreal rows, and a the limbs were contrastingly banded dark lower mean number of ventral scales (22.4 and pale green. The throats were bright versus 23.8). There is also a strong ten- yellow to bright green, with scattered dency for adult caeruleolatus to have mod- deeper green spots in each case. In a pair erate to low body crest scales, whereas in from La Mata, the dorsa were bright baleatus the tendency is toward high to green, somewhat marbled with yellow and moderate body crest scales. Compared yellow-green, the upper surfaces of the with multistruppus, caeruleolatus has mod- heads were pale fawn, the eyeskin pale ally 4 (rather than 2) snout scales at the grayish green, and the dewlaps orange in level of the second canthals, 8 rather than both sexes. 7 vertical rows of loreals, 5/5 rather than The series of A. b. caeruleolatus includes 4/4 scales between the interparietal and four juveniles and subadults with snout- the supraorbital semicircles, and a lower vent lengths from 60 to 91; the largest of mean of vertical dorsal scales (17.1 versus these is a topotype that was colored and 18.6). With regard to body crest scales, HisPANioLAN Giant Angle • Schwartz 129 these two subspecies show the same situa- tion as caeruleolatus and baleatus. Com- pared with .suJ)limis, caeruleolatus has 4 (rather than 2) snout scales at the level of the second canthals, 8 (rather than 7) ver- tical rows of loreals, 5/5 (rather than 4/4) scales between the interparietal and the supraorbital semicircles, and lower means of both vertical dorsals (17.1 versus 19.2) and ventrals (22.4 versus 25.1). A. h. suh- limis has not been recorded as having the dorsal body crest scales other than high, in contrast to the strong tendency in cae- ruleolatus of having these scales moderate to low. Discussion. A. h. caeruleolatus centers in the extremely mesic eastern portion of the Valle de Cibao in that area that has the most rainfall in the Republica Domin- icana. I have already commented on the specimens from Los Bracitos, Duarte Prov- ince; these specimens are old and pattern- less and are from a locality in the Cordillera Septentrional which is, farther west, occu- pied by A. h. baleatus; I include them with caeruleolatus provisionally. The specimen from Ponton, Duarte Province, is a juve- nile (ASFS V2987; snout-vent 60) and is presently patternless; no color data are available. It too I only provisionally re- gard as caeruleolatus. The two specimens from La Vega Province (ASFS V4317, MCZ 128379) are also without color data in life, and the former is a patternless ju- venile (snout-vent 69). Specimens from these last two localities also require verifi- cation as to subspecfic status. A. h. caeruleolatus presumably inter- grades with four subspecies: baleatus, mul- tistruppus, the subspecies on the Peninsula de Samana, and subspecies to the south- east. Only in the last case are specimens that I interpret as intergradient known, and they will be discussed under the de- scription of the southeastern subspecies. No intergrades are known between the Samana subspecies, baleatus, or multistrup- pus. Distance between caeruleolatus and the nearest localities for these subspecies are: Samana subspecies — 13 kilometers (Caiio Abajo and 5 km NW Sanchez); baleatus — 50 kilometers (Los Bracitos and Pena); multistruppus — 12 kilometers (12.8 km NW Bonao and Guaigiii). Of these presumed areas of contact, that between caeruleolatus and the Samana subspecies is not unexpected; the area between the two known localities is very open and relatively barren and devoid of trees and appears al- ways to have been so. There are fine high swamp-forests in the western part of this intervening region, and it is possible that intergrades between these two distinctive subspecies will be encountered in these forests. Most puzzling is the absence of intergradation between caeruleolatus and multistruppus. The specimen from north- west of Bonao is a juvenile, but it does not show the characteristic multiple crossbands of both young and adult multistruppus. It may be that multistruppus occupies only the foothills of the Cordillera Central and that the zone of intergradation between multistruppus and caeruleolatus is very abrupt. Remarks. A. b. caeruleolatus is known from sea level to an elevation of 1000 feet (305 meters) in the haitises region near Gonzalo and 1200 feet ( 366 meters ) north- west of Bonao. Specimens were secured primarily from native collectors; the long series of topotypes is due to the industry of the inhabitants of Cafio Abajo. The Cafio Abajo area is one of cafetales and cacaotales with high canopied shade-trees, and the lizards apparently are extremely abundant in this optimal habitat. The pair of lizards from La Mata were secured by me while they were copulating on the side of a large shade-tree in a cafetal about 4 feet (1.2 meters) above the ground at 1225 hours. The two females from Gon- zalo were taken during the day on large trees adjacent to a small spring in the haitises; the surrounding area was under heavy cultivation, but the doline slopes were covered locally with undisturbed for- est. The name caeruleolatus is from the Latin "caeruleus" for "blue" and "latus" 130 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2 for "side," in allusion to the sky-blue lower sides of this subspecies. Anolis baleatus samanae new subspecies Holotype. CM 54105, an adult male, from 7.6 mi. (12.2 km) NE Sanchez, 1000 feet (305 meters), Samana Province, Re- publica Dominicana, one of a series col- lected by native collectors on 28 November 1971. Original number ASFS V34474 . Paratypes. ASFS V34475-79, same data as holotype; USNM 193990-92, same local- ity as holotype, native collectors, 27 No- vember 1971; MCZ 125634, 5.0 mi. (8.0 km) NW Sanchez Province, Republica Do- minicana, J. Aria, 27 November 1971; ASFS V34495-96, 5.0 mi. (8.0 km) NW Sanchez, Samana Province, Republica Dominicana, J. Aria, 28 November 1971; CM 54127-30, 5.0 mi. (8.0 km) NW Sanchez, Samana Province, Republica Dominicana, J. Aria, 30 November 1971; MCZ 12563.5-39, USNM 193993-4001, 5.0 mi. (8.0 km) NW Sanchez, Samana Province, Republica Do- minicana, J. Aria, 1 December 1971; ASFS V34514, ASFS V34836-38, Las Terrenas, Samana Province, Republica Dominicana, native collector, 28 November 1971; ASFS V1904, 6 km E Sanchez, Samana Province, Republica Dominicana, R. Thomas, 30 Oc- tober 1963; AMNH 28651, Samana, Sa- mana Province, Republica Dominicana, J. King, August 1924; AMNH 39817-23, AMNH 42285, Laguna, Samana Province, Republica Dominicana, W. G. Hassler, Oc- tober-December 1929; USNM 61928, Cayo Hondo, Samana Province, Republica Do- minicana, W. L. Abbott, February 1919. Definition. A subspecies of A. baleatus characterized by the combination of mod- ally 2 snout scales at level of second can- thai scales, 7 vertical rows of loreal scales, 3 scales between the interorbital semicir- cles, 4/4 scales between the inteiparietal and the supraorbital semicircles, moderate number of vertical dorsal scales ( 13-20; mean 16.6), moderate number of ventral scales (16-29; mean 22.1), nuchal crest scales very high to high (usually) to mod- erate or low (rarely) in both sexes, body crest scales high to moderate but often low in both sexes, subocular scales almost al- ways separated from supralabial scales by one (rarely 2) row of scales; dorsum in both sexes in life blotched dark green, greenish, dull gray-green, brown, or black- ish, dewlaps in males dull yellow to pale yellowish orange, in females very pale yel- low to pale yellowish orange, streaked with blackish or brown basally, and chin and throat in males cream to yellowish or yel- low-orange, mottled with black or gray, in females pale green to greenish yellow with dark green to brown streaking or even re- ticulate. Distribution. The Peninsula de Samana in the northeastern Republica Dominicana, and apparently islets in the Bahia de Sa- mana. Description of holotype. An adult male with a snout-vent length of 145 and a tail length of 222 (regenerated); snout scales between second canthals 3, 6 vertical rows of loreal scales, 3 scales between the supra- orbital semicircles, 4/4 scales between the interparietal and the supraorbital semicir- cles, vertical dorsals 18, horizontal dorsals 19, ventrals 21, one row of scales between the suboculars and supralabials, fourth toe lamellae on phalanges II and III 30, nuchal crest scales very high, body crest scales high; in life, dorsum mottled dull greens and gray-brown with whitish (almost cream but suffused with pale gray); upper surface of head mixed dark brown and gray, venter dull greenish, dewlap orange, chin and throat creamy to yellowish, not marked with green. Variation. The series of 54 A. b. sama- nae consists of 32 males and 22 females. The largest male (AMNH 39807) has a snout-vent length of 157; the largest fe- males (CM 54130, USNM 193994) have snout-vent lengths of 145. The male is from Laguna, the females from 5.0 mi. NW Sanchez. Snout scales at the level of the second canthals range between 2 and 5; the mode is 2 (24 specimens). The verti- cal loreal rows vary between 5 and 9; the HisPANioLAN Giant Angle • Scliwartz 131 mode is 7 (25 specimens). There are 2 or 3 scales between the supraorbital semicir- cles (mode 3). There are modally 4 scales between the interparietal and the supraor- bital semicircles; 4 scales are involved in 43 percent of the combinations: actual counts are 3/3 (2), 3/4 (3), 4/4 ( 17), 4/5 (8), 5/5 (13), 5/6 (5), 6/6 (1), 6/7 (2), 4/6 (1), and 3/5 (1). Vertical dorsals range between 13 and 20 (mean 16.6), horizontal dorsals between 13 and 27 (19.3), and ventrals 16-29 (22.1). Of 30 adult males, 14 have the nuchal crest scales very high, 15 have them high, and one has them moderate; in 20 adult females, nine have the nuchal crest scales very high, nine have them high, one has them mod- erate, and one has them low. Body crest scales in males are high in three lizards, moderate in 16, and low in ten; in females, 11 have these scales moderate and ten have them low. The suboculars are separated from the supralabials by one row of scales in all but four specimens (7 percent), which have them in contact, and one spec- imen (2 percent), which has 2 rows of scales in this position. A. ]). samanae is basically a blotched liz- ard, and no adults show any indication of crossbanding. The body is irregularly blotched with blackish, dark green, dull green, gray-brown, and occasionally there are sky-blue areas along the ventrolateral margin of the dorsal coloration in males, but these areas are not so prominent as in caeruleolatus. Regardless of the dorsal shades, the upper surface of the head is mixed dark brown and shades of gray in both sexes. The hindlimbs are finely barred with pale and dark green. The venter is dull greenish in both sexes. The dewlap in males varies from dull yellow or pale yellowish orange to orange, and the chin and throat are yellowish, cream, or yellow-orange, mottled with black or gray. In females, the dewlaps are very pale yel- low, pale yellow-orange, or grayish orange, at times streaked with blackish or brown basally, and the chin and throat ground color is pale green, marbled, streaked, or even reticulate with dark green to (rarely) brown. There arc one juvenile (AMNH 28651; snout-vent length 40) and two subadult (snout-vent lengths 92 and 97) A. 1). sa- manae. The subadults are old and discolored but their patterns seem not to differ from those of full adults. The juvenile on the other hand, has fom- bold pale crossbars on the dorsum, the pattern continuing onto the tail. This young individual has the umbilicus still present and is presumably near hatchling size. Comparisons. Since samanae and cae- ruleolatus are adjacent geographically, the most pertinent comparisons are between them. Examples of these two populations, as noted in the introduction to the present paper, were available to me simultaneously and I was struck with their differences in life. A. 1). samanae is a blotched lizard whereas caeruleolatus is a crossbanded one; the latter subspecies also typically has sky-blue ventrolateral blotches, a feature absent (or occasionally poorly expressed) in male samanae. Male dewlap colors are similar in both subspecies, although fe- male dewlap colors in samanae seem some- what paler than those of caeruleolatus. The chin and throat markings of the two subspecies are quite distinct; in male cae- ruleolatus, the throat is deep yellow to yellow-orange, at best with very faint gray- ish dots or smudges, whereas in male samanae the throat is yellowish or cream to yellow-orange, mottled with black or gray. In female caeruleolatus, the throat is yellow to yellow-green, always with some dark green dots, blotching, or mar- bling, whereas in samanae females, the throat is pale green, greenish yellow, or yellow-green, with dark green to brown streaking or reticulum. The only subspecies thus far described which is blotched like samanae is the Cor- dillera Central suhlitnis, although caerul- eolatus may show a marbled dorsum in some areas. No pigmental or pattern dif- ferences separate samanae and sublimis, since in both dorsal coloration and color of 132 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2 the dewlap the major color involved is green. However, the throat in male sub- limis is pale green, whereas in samanae it is cream to yellow-orange. Certainly rnulti- struppus and samanae are easily distin- guished in the field by their very different dorsal patterns, for example, and haleatus, with its very, very bright chin and thioat, both of which are immaculate, is quite dis- tinctive from samanae. In meristic data, samanae differs from caeruleolatus in having 2 (rather than 4) snout scales, 7 (rather than 8) vertical rows of loreals, and 4/4 (rather than 5/5) scales between the inteiparietal and the supraorbital semicircles. From multistrup- pus, samanae differs in having a lower mean of vertical dorsal scales ( 16.6 versus 1S.6), and the same difference occurs be- tween samanae and su])Umis ( 16.6 versus 19.2) and in ventrals (22.1 versus 25.1). From haleatus, samaiuie differs in having 2 (rather than 4) snout scales, 4/4 (rather than 5/5) scales between the inteiparietal and the supraorbital semicircles, and lower means in both vertical dorsals ( 16.6 versus 17.5) and ventrals (22.1 versus 23.8). The nuchal crest scales in samanae are more consistently very high to high than they are in any of the other subspecies of A. haleatus. Discussion. As pointed out in the dis- cussion of A. h. caeruleolatus, there are no intergrades known between that subspe- cies and samanae. The isthmus of the Pe- ninsula de Samana is much cleared and locally even barren, but there are large western swampy areas that support mag- nificent hardwood forests toward the land- ward side. These forests may well support intermediates between samanae and cae- ruleolatus, or, because of their proximity to the mainland, they may be inhabited by caeruleolatus. Specimens from 5.0 mi. NW Sanchez, that locality for samanae which is closest to a known locality for caeruleo- latus ( 18 kilometers ) , show no tendencies toward the crossbanded condition of cae- ruleolatus. A. h. samanae is the only Hispaniolan giant anole known by specimens from any off-shore island or islet. The specimen from Cayo Hondo, taken by W. L. Abbott, constitutes this record, although I am un- able to locate this islet. I assume it is one of the archipelago within the Bahia de Sa- mana. Remarks. All but one A. h. samanae se- cured by myself and parties were native- collected. The exception is a lizard taken by Richard Thomas, one of two seen on a small tree and in a vine tangle in a steep limestone ravine east of Sanchez. The area of the type locality is in the uplands of the Sierra de Samana on the road between Sanchez and Las Terrenas. Thus newly constructed road passes through superb mesic high-canopied forest, and much of the area is not yet seriously disturbed. Ob- viously from the number of lizards secured by natives in this region, A. /;. samanae is common. The range is not high, with a maximum elevation of 1673 feet (510 me- ters) in Monte Las Caiiitas; this mountain lies between Sanchez and Las Terrenas. Specimens from Las Terrenas itself were secured by natives from near-coastal mesic cafetales and cacaotales, and lizards from northwest of Sanchez were in similar situ- ations. Only three other reptiles (Diplo^lossus sternurus alloeides Schwartz, Leiocephalus personatus pyrrholaemus Schwartz, and Dromicus parvifrons niger Dunn) are known to have differentiated at the subspecific level on the Peninsula de Samana. Sphaero- clactylus clenchi Shreve and Sphaeroclac- tijlus samanensis Cochran both occur there and have as yet unnamed populations, one of which in each case is limited to the pen- insula. It is also of interest to note that in Anolis clisticJius Cope, the Samana popula- tion is identical to the population on the soutliern shores of the Bahia de Samana (ignigularis Mertens), but that the range of this subspecies is interrupted at the head of the Bahia de Samana by A. cl. domini- censis Reinhardt and Liitken (see Schwartz, 1968: 280-81, for details). HisPANioLAN Giant Angle • Sclucmiz 133 Anolis baleatus litorisilva new subspecies Holotype. USNM 193977, an adult male, from 1.2 km SSW Piinta Cana, La Altagracia Province, Rcpviblica Domini- cana, one of a series collected by Danny C. Fowler and Bruce R. Sheplan, on 24 November 1971. Original number ASFS V35095. Paratypes. ASFS \'35096-100, same data as holotvpe; CM 54113-14, MCZ 125616-17, 5.5 km SSW Punta Cana, La Altagracia Province, Republica Domini- cana, D. C. Fowler, 27 November 1971; ASFS V29090, Juanillo, La Altagracia Province, Republica Dominicana, native collector, 24 July 1971; ASFS V961-62, 0.5 mi. NW Boca de Yuma, La Altagracia Province, Republica Dominicana, R. F. Klinikowski, R. Thomas, 2 September 1963; ASFS VI 136, 2.5 km NW Boca de Yuma, La Altagracia Province, Republica Domin- icana, native collector, 4 September 1963; ASFS V17573, 4 km NW Boca de Yuma, La Altagracia Province, Republica Domi- nicana, A. Schwartz, 13 June 1969; ASFS V17616, 2 km NW Boca de Yuma, La Al- tagracia Province, Republica Dominicana, J. B. Strong, 15 June 1969. Definition. A subspecies of A. I)(ileatus characterized by the combination of 2 or 4 scales at level of the second canthal scales, 7 vertical rows of loreal scales, 3 scales be- tween the interorbital semicircles, 4/5 scales between the interparietal and the supraorbital semicircles, low number of vertical dorsals (13-19; mean 15.9), low number of ventral scales ( 18-26; mean 21.3), nuchal crest scales always very high to high in both sexes, body crest scales high (rarely) to moderate or low, suboc- ular scales usually separated from supra- labial scales by one row of scales; dorsum in life varying from light blue-brown to light greenish brown in males, dull brown to olive-brown in females, blotched with creamy to gray, dewlap in males bright orange, brownish in females, and chin and throat (including lips) bright orange in males, pale yellow-green in females. Distri])ution. Extreme eastern Repub- lica Dominicana in La Altagracia Province, from Punta Cana to the \'icinity of Boca de Yinna. Description of holotype. An adult male with a snout-vent length of 136 and a tail length of 183 (regenerated); snout scales between sc^cond canthals 2; 6 vertical rows of loreal scales, 3 scales between the supra- orbital semicircles, 5/5 scales between the interparietal and the supraorbital semicir- cles, vertical dorsals 15, horizontal dorsals 16, ventrals 25, subocular scales in contact witli the supralabial scales, fourth toe la- mellae on phalanges II and III 31, nuchal crest scales very high, dorsal body crest scales high; in life, dorsum blotched light blue-brown and light green-brown; venter pale gray-green; chin, lips, and dewlap bright orange. Variation. The series of 16 A. h. litori- silva is composed of six males and ten fe- males. The largest male (MCZ 125616) has a snout-vent length of 158, the largest female (ASFS V961) 131. The male is from 5.5 km SSW Punta Cana, the female from 0.5 mi. NW Boca de Yuma. Snout scales at the level of the second canthals range between 2 and 5; there are t\vo modes, 2 and 4, each with five individuals. The vertical loreal rows vary betwe(Mi 6 and 9; the mode is 7 (nine specimens). There are 2 to 4 scales between the supra- orbital semicircles (mode 3). There are modally 4/5 scales between the interpari- etal and the supraorbital semicircles; 5 scales are involved with 59 percent of the combinations; actual counts are 4/4 (4), 4/5 (6), 5/5 (5), and 5/6 (1). Vertical dorsals range between 13 and 19 (mean 15.9), horizcmtal dorsals between 14 and 22 (18.5), and ventrals between 18 and 26 (21.3). Of four adult males, three have the nuchal crest scales very high and one has them high; of five adult females, two have these scales very high and three have them high. In the males, the body crest scales are high in one and moderate in three, and in the females, these scales are 134 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2 moderate in two and low in three. The siiboculars are separated from the siipra- labials by one row of scales in all but one specimen (6 percent). A. h. litorisilva is essentially a blotched lizard whose colors do not include bright or even medium greens. The color notes on the holotype apply equally well to the other adult males — the dorsum is blotched with bluish browns and light greenish browns, without any clear greens, and the blotching is often more pronounced on the head than on the body. In females, the dorsum is dull brown to olive-brown with only occasional slight remnants of a lighter green pattern on the head; the blotching in the female involves creamy to gray pig- mentation. The venter is pale gray-green or whitish green in males, pale greenish gray in females. The dewlap in all adult males was recorded as bright orange, and brownish in females. In males, the chin (including the lips) is bright orange, and pale yellow-green in females. The upper surface of the head in males is blotched like the body and is dark chocolate in fe- males. In females, the upper surfaces of the hindlimbs were recorded as olive- brown, blotched with cream to gray like the dorsum. The series of A. h. litorisilva contains seven juveniles and subadults (snout-vent lengths 45 to 88). The smallest juvenile (ASFS V17573, female) was bright green in life with four pale buffy crossbands and dark green shadow-bars between the cross- bands; the tail was ringed cream and dark gray, and the venter was pale green. The dewlap was yellow-green and gray. A slightly larger female (ASFS V17616) with a snout-vent length of 57 was yellow-green dorsally and without bands, the head was brown; the eyeskin was green, and the venter yellow-green. The tail was banded black and yellow-green, and the dewlap was mainly brown with the scale rows yel- low-green. A still larger female (ASFS V1136) with a snout-vent length of 67 was green, faintly crossbarred with grayish green, and there were charcoal smudges on the neck. Two male subadults with snout- vent lengths of 71 and 83 (ASFS V35099- 100) from the type locality were recorded by Fowler as follows: "One with a strong vertical banding pattern alternating brown- green and white-gray, which extends from tip of tail to the head where it becomes slightly more diffuse; on the other, the dor- sal groimd color is dull brown with rem- nants of banding pattern only around head; the ventral ground color of the first is gray- green with brown mottling, the second is dull gray-brown; in both juveniles, the dewlap is orange-green and the chin and lips are green." The largest subadult (ASFS V29090) was patternless green above, and the dewlap was orange with charcoal stripes; the specimen is a female. Comparisons. Because of its blotched (rather than crossbanded) pattern, litori- silva requires comparison with samanae and suhlimis. The general effect of the dorsa of all three subspecies is quite sim- ilar, but samanae and suhlimis are much the brighter lizards, with greens predomi- nant in the dorsal pigmentation. On the other hand, litorisilva is a much more drab lizard, without clear greens in the adults, the tendency being toward more sombre hues, primarily shades of browns. From all other described subspecies, litorisilva differs in being blotched rather than cross- banded and also in having much less gaudy dorsal colors. In meristic counts, litorisilva differs from the remaining sub- species in the following ways. From cae- ruleolattis, litorisilva differs in having 7 (rather than 8) vertical loreal rows, and lower means of vertical dorsals (15.9 ver- sus 17.1) and ventrals (21.3 versus 22.4). From rnultistruppus, litorisilva differs in lower means of vertical dorsals ( 15.9 ver- sus 18.6) and ventrals (21.3 and 22.3). From stihlitiiis, litorisilva differs in having lower means of vertical dorsals ( 15.9 ver- sus 19.2) and ventrals (21.3 versus 25.1). From haleatus, litorisilva differs in having lower means of vertical dorsals ( 15.9 ver- sus 17.5) and ventrals (21.3 versus 23.8). Meaningful comparisons of litorisilva with HisPANiOLAN Giant Anole • ScJiwartz 135 other subspecies in counts of snout scales, and scales between the interparietal and the supraorbital semicircles, are impossible since litorisilva has a bimodal condition in the former (and the bimodes are 2 and 4, those counts which occur singly as the mode in the other subspecies) and has a mode of 4/5 in the latter (whereas all other species have either 4/4 or 5/5). Considering the fairly large series of litori- silva (16 specimens), these two "abnormal" conditions are puzzling. At least in the case of 4/5 counts, the absence of 3/3 or 3/4 counts in litorisilva suggests that this subspecies tends toward a 5/5 count. Discussion. A. /;. litorisilva appears to be the extreme eastern isolate of the more widespread A. haleatus stock. It occupies semi-arid forests on and near the coast (as at Juanillo and Punta Cana) and on the limestone ridge behind Boca de Yuma. Both situations are far more xeric than is customary for A. haleatus, and the faded nongreen coloration of the adults is doubt- less a response to the dry and open to dense forest conditions of this region. Nevertheless, individuals are quite con- spicuous at night as they sleep exposed. A. b. litorisilva presumably comes into con- tact with the subspecies to the north and west (named below) but intergrades are presently unknown; in the vicinity of Hig- iiey (the closest locality for the adjacent subspecies) the lizards are more brightly colored and crossbanded and quite unlike litorisilva. Remarks. All but one specimen of li- torisilva were collected by myself and par- ties. Individuals were found sleeping in primarily coastal forest (to which the name, from "litus" for "shore" and "silva" for "forest," refers in Latin) at elevations from 4 to 15 feet (1.2 to 4.6 meters) above the ground. Generally, juveniles sleep closer to the groimd and in more dense situations than adults. One juvenile was taken from a roadside Acacia, a most un- usual situation (since Acacia is a distinct xerophyte) for any giant anole. Several adults were taken in dense viny tangles, sleeping on the woody vines; the advan- tage of this situation was made (piite ob- vious wIkmi I attempted to catch a large adult at night by hand. The light from my flashlight wakened the lizard almost im- mediately, and although 1 was extremely careful not to jar any of the vines, this was a vain endeavor. At the first jostling, the lizard jumped to the ground and escaped in the dry leaf litter and understory. Anol'is haleatus scelestus new subspecies Holotype. CM 54106, an adult male, from 5.1 mi. (8.2 km) E Santo Domingo (from Rio Ozama), Distrito Nacional, Re- publica Dominicana, one of three collected by David C. Leber and Richard Thomas on 18 June 1964. Original number ASFS V2460. Paratopes. ASFS V2461-62, same data as holotype; MCZ 125618-27, 8.4 mi. (13.4 km) NE La Romana, 100 feet (31 meters). La Romana Province, Republica Domini- cana, B. R. Sheplan, 22 November 1971; CM 54115-18, USNM 193981-89, 8.4 mi. (13.4 km) NE La Romana, 100 feet (31 meters). La Romana Province, Republica Dominicana, D. C. Fowler, A. Schwartz, 17 July 1971; MCZ 16321, La Romana, La Romana Province, Republica Dominicana, E. Leider, 1922; ASFS V29284-300, 0.2 mi. (0.3 km) N Otra Banda, 350 feet (107 me- ters). La Altagracia Province, Republica Dominicana, D. C. Fowler, A. Schwartz, 26 July 1971; ASFS V21699-700, 1 km NE Higiiey, La Altagracia Province, Republica Dominicana, J. R. Dennis, R. Thomas, 16 August 1969; USNM 193979-80, 0.7 mi. (1.1 km) W Higiiey, La Altagracia Prov- ince, Republica Dominicana, R. Thomas, 29 August 1963; ASFS V1038, 1 mi. (1.6 km) W Higiiey, La Altagracia Province, Republica Dominicana, R. Thomas, 3 Sep- tember 1963; ASFS V28757, 15.5 mi. (24.8 km) E San Pedro de Macoris, Rio Cumay- asa. La Romana Province, D. C. Fowler, 12 July 1971; ASFS V28910-16, 15.5 mi. (24.8 km) E San Pedro de Macoris, Rio Cumayasa, San Pedro de Macoris Province, Republica Dominicana, D. C. Fowler, A. 136 Bulletin Musewyi of Comparative Zoology, Vol. 146, No. 2 Schwartz, 16 July 1971; ASFS V28847, 15.5 mi. (24.8 km) E San Pedro de Macoris, La Romana Province, Repiiblica Dominicana, A. Schwartz, 15 July 1971. Associated specimens. REPUBLICA DOMINICANA: La Altagracia Province, 1 km SE Las Lisas (ASFS V17434-35); San Cristobal Province, 8 km N Yamasa, 200 feet (61 meters) (ASFS V28656). Definition. A subspecies of A. haleatus characterized by the combination of mod- ally 2 scales at level of the second canthal scales, 7 vertical rows of loreal scales, 3 scales between the supraorbital semicircles, 5/5 scales between the interparietal and the supraorbital semicircles, low number of vertical dorsals ( 12-20; mean 15.4 ) , low number of ventral scales ( 17-28; mean 21.1), nuchal and body crest scales always very high to high in both sexes, subocular scales usually separated from supralabial scales by one (occasionally two) row of scales; dorsum in both sexes either green with three pastel green crossbands or dark green flecked with light green, cream with some greenish to brownish green smudges, dewlap in males deep yellow to deep or- ange, streaked or smudged with dark brown to charcoal, and throat in females dark green marbled with yellow and pale green (males unrecorded). Distribution. Southeastern Republica Dominicana, from the Sierra de Yamasa and the vicinity of Santo Domingo in the west, east to the region about Higiiey and Las Lisas in La Altagracia Province. Description of Jiolotype. An adult male with a snout-vent length of 152 and a tail length of 267; snout scales between second canthals 4; 8 vertical rows of loreal scales, 2 scales between the supraorbital semicir- cles, 4/5 scales between inteiparietal and supraorbital semicircles, vertical dorsals 16, horizontal dorsals 16, ventrals 22, sub- ocular scales separated from supralabial scales by one row of scales, fourth toe la- mellae on phalanges II and III 34, nuchal crest scales high, body crest scales moder- ate; in life, dorsum olive-green with six pastel green crossbands, tail and venter light green; dewlap dark yellow. Variation. The series of 61 A. b. sceles- ttis consists of 27 males and 34 females; a large number of the specimens are juve- niles and subadults. The largest male (ASFS V29284) has a snout-vent length of 180, the largest female (ASFS V29286) 147; both are from near Otra Banda. Snout scales at the level of the second canthals range between 2 and 4; the mode is 2 (32 specimens). The vertical loreal rows vary between 5 and 8, with a mode of 7 (25 specimens). There are 1 to 4 scales be- tween the supraorbital semicircles (mode 3). There are modally 5/5 scales between the interparietal and the supraorbital semi- circles; 5 scales are involved in 49 percent of the combinations; actual counts are 3/4 (2), 4/4 (14), 4/5 (14), 5/5 (17), 5/6 (11), 6/6 (1) and 4/6 (1). Vertical dor- sals range between 12 and 20 (mean 15.4), horizontal dorsals between 15 and 25 (18.8), and ventrals between 17 and 28 (21.1). Of 11 adult males, nine have the nuchal crest scales very high and two have them high. Of 16 adult females, nine have these scales very high and seven have them high. Body crest scales in males are high in two lizards, moderate in eight, and low in three; in females, the body crest scales are high in two, moderate in eight, and low in six. Fifty-three specimens have the sub- oculars separated from the supralabials by one row of scales, whereas in four lizards ( 7 percent ) these scales are in contact, and in two lizards (3 percent) they are sepa- rated by tv/o rows of scales. In general, both sexes of A. b. scelestiis show a pattern of about six or seven fine crossbands that are often obscured by dor- sal blotching. Colors are shades of greens, with brighter green the base color and the blotching tending toward darker shades. The crossbands are lighter pastel shades of green, and in some lizards the dorsal ground color is olivaceous. Another vari- ant, which is somewhat more prevalent in females, is an olive green to dark green HisPANioLAN Giant Angle • Schwaiiz 137 dorsum, flocked with pale green. Two fe- males from near Higiiey showed still an- other style of body pattern and color, with the dorsal ground color cream with some dark green to brownish green snuidges, and the neck with alternating pale blue and charcoal markings, the pale blue mark- ings persisting onto the cheeks. In males the upper surface of the head is brown, and in females it is mixed brown and green, with the snout and supraocular scales deep green in some lizards. In fe- males, the chin and throat are dark green, marbled with yellow and pale green. The dewlap is rather \'ariable; in males it has been recorded as dark yellow or deep yel- low to orange or dark orange, whereas in females the dewlap varies from yellow to dark orange with dark brown, olivaceous, or charcoal streaking, marbling, or smudg- ing. Although there are no color notes in life, in the preserved lizards the eyeskin is regularly pale gray, and I presume that in life the eyeskin is set off from the rest of the head color in some pigmental fashion. Many specimens of both sexes have the lower sides tigroid with "stripes" extend- ing conspicuously onto the lateral sides of the abdomen. There are 34 juvenile and subadult A. b. sceJestus, with snout-vent lengths between 46 (USNM 193989) and 94 (ASFS V21699-700). Three juveniles (snout-vent lengths 46-61) have umbilici still present. This entire suite of young lizards shows a remarkable diversity in dorsal pattern. Even small specimens may be either uni- color green (usually with a vertical nuchal white crescent and a white subocular spot ) , green with three or four yellow body bands, or there may be many more bands resulting from the inteiposition of pale body bands between the primary pale body bands. One specimen (ASFS \'29296; snout-vent, length 70, male ) has both pale body bands and interstitial pale blotching, whereas another lizard (MCZ 125621; snout-vent length 86, female) already shows the adult pattern of several fine pale crossbands on a green ground. The largest subadults, however, (ASFS V21699-700; snout-vent lengths 94, male and female) are both presently unicolor and show no indications of the adult body banding. That a single juvenile may demonstrate a pattern change is shown by the following notes on ASFS V28757, a female with a snout-vent length of 54: "Alive, emerald green with about foiu' pale yellow cross- bands on body; dead — seven narrow brown body bands which are hollowed, and the dorsal groimd color now pale yellow- green." The dewlap in young males is or- ange, in young females from dull yellow streaked with charcoal to charcoal. Comparisons. In color and pattern, A. h. scelestus differs from all other subspe- cies. No other named population has six or seven narrow dorsal crossbands; even multistnippus is much more conspicuously banded than scelestus and lacks any sort of dorsal blotching. A. h. scelestus is known to intergrade with more northern caeruleolatus and is presumed to meet li- torisilva. In each case, there is no difficulty distinguishing the adjacent forms chromat- ically. A. h. caeruleolatus typically has (in males) sky-blue blotches along the junc- tion of the dorsal and ventral colors, and is prominently crossbanded with three dor- sal crossbands. A. h. litorisilva is a blotched lizard, the dorsal colors much more drab than those of scelestus, tending toward browns and brownish greens. Perhaps scelestus most closely resembles multi- stnippus, but, although both are banded, the bands in multistruppus are much finer and much more numerous than the six or seven pale dorsal crossbars in scelestus. A. h. scelestus, with modally 2 snout scales, differs from caeruleolatus, which has 4 snout scales. In having 7 vertical lor- eal rows, scelestus differs from caeruleola- tus, which has 8 rows. In having 5/5 scales between the interparietal and supraorbital semicircles, scelestus differs from samanae, multistruppus, and .mhlimis, all of which have 4/4. A. h. scelestus has the lowest 138 Bulletin Museum of Comparative Zoology, Vol. 146, No. 2 mean of vertical dorsals ( 15.4 ) of all named subspecies, being most closely ap- proached by litorisilva (15.9). A. b. sceles- tus males are larger than those of any other subspecies ( 180 in scelestus, 158 in litori- silva, which is second largest) and in fact this subspecies exceeds all other Hispanio- lan giant anoles in size, being most closely approached by male A. r. ricordi, which reach a snout-vent length of 160. Discussion. I am uncertain that all specimens included in scelestus should be so associated. This is especially true of the specimen from near Yamasa (ASFS V28656); this is a juvenile male and its taxonomic status remains somewhat in doubt, since it is young. It is also possible that specimens from Santo Domingo like- wise are not identical with more eastern lizards, although the two samples agree fairly well. A. b. scelestus and A. b. caeruleolatus in- tergrade in the region of El Seibo Prov- ince; I have examined the following mate- rial from El Seibo which I consider intergradient: 3.5 mi. (5.6 km) S Sabana de la Mar (ASFS X7877); 2.1 mi. (3.4 km) N El Valle (ASFS X7861-62); 3 km N El Valle (ASFS V3157-58); 10.5 km N Hato Mayor (ASFS V35329-30). This series consists of three juveniles and four young adults (with snout-vent lengths between 112 and 127). The single adult male (ASFS X7877) was tannish gray in life with darker brown blotches, a pale green venter, and an orange dewlap. Two adult females (ASFS X7861-62) were pale pea- green with vertical gray bars, the upper surface of the head grayish tan, venter green, and dewlap grayish orange. The lower jaw and throat were green mottled with darker green. In general this series seems closer to caeruleolatus than to scel- estus, but the male lacks sky-blue ventro- lateral markings. On the other hand, the vertical gray bars, recorded for the female, resemble the pattern of scelestus rather than that of female caeruleolatus. It seems likely that caeruleolatus and scelestus in- tergrade in this region. Remarks. Almost all ASFS scelestus were secured while the lizards were asleep at night. Typical situations are lowland cacaotales and cafetales with their high canopied shade-trees, along lowland streams (as at Otra Banda and Yamasa), and in woods associated with limestone cliffs (east of Santo Domingo). The long series from the Rio Cumayasa is from the high riverine woods along that stream; re- markably, we secured only juveniles and subadults at this locality, despite three nocturnal visits. One juvenile from this lo- cality was taken on the exposed branch of an Acacia tree along an open road. Per- haps the most remarkable place whence A. b. scelestus has been taken is the locality northeast of La Romana. This place is a deep and well-wooded ravine through which flows a clear stream; however, the ravine is completely surrounded on all sides by cane fields, and the ravine woods are completely isolated at the ravine rim from other such ecologies, if they even still exist in this area. A. b. scelestus was ex- ceptionally abundant in this particular and very restricted strip of riverine gallery for- est. Elevations above ground recorded for sleeping scelestus range from 2 to 20 feet (0.6 to 6.1 meters), with juveniles usually sleeping much lower than adults. The alti- tudinal distribution of A. b. scelestus is in general low, with recorded elevations from sea level to 200 feet. It is likely that this subspecies also occurs in the uplands of the Cordillera Oriental, but as yet there are no specimens from areas within that rather low-lying but mesic and well-for- ested massif. The name scelestus is from the Latin for "unlucky, wretched," in allusion to the dif- ficulties involved with collecting this sub- species at the La Romana ravine noted above. The transition betu'cen scelestus and //- torisilva must be very abrupt; the two sub- species are known from localities separated by only 28 kilometers. The habitats of the two subspecies are quite different, with scelestus inhabiting very mesic situations HisPANiOLAN Giant Angle • Schwartz 139 and litorisilva xeiic coastal woods. Inter- estingly, this same eastern region of the Repiiblica Dominicana is also an area of abrupt changes in subspecies of Anolis di- stichus, where the subspecies ifi,ni^\ i j Epigonus wbustus 1S9 ti^ty Y^ars. Vaillant (1888: 25) remarked Epigonus lenimen 193 that E. telescopus was recognized in ancient Epigonus crassicaudus 197 times, and Risso (1810: 303) reported that Species Incertae Hedis 199 j-j-jj^ gpecies was prized for its firm, deli- Aek„S^!Sr°!!!' ::;:::::::::::::::::::::::::: loE! cious-tasti„g „,«,, although it was rardy Literature Cited 200 taken. The presence of common names tor Appendix 203 E. telescopus in vocabularies of western Mediterranean and North Atlantic fishing Abstr.\ct. a study of the deep-sea Apogonidae communities ( Doderlein, 1889 ) provides results in a revision of the genus Epigonus additional evidence of man's long-term Rafinesque. Twelve species are recognized in- awareness of the species. £. telescopus is eluding two new forms — E. oligolepis and £...,, • n i i • . i i i. „r pectirUfer. E. fragilis (Jordan and Jordan) is still occasionally sold in the markets of southwestern Europe. ' This paper is based on a portion of a thesis Two other species of EpigOnus are cap- presented to Harvard University in partial ful- tured by commercial fishermen. E. denti- fillment of the requirements for the Ph.D. in culcitUS is edible ( Dieuzeide et al., 1953: ^i?'"g>- ^ r -1^) and is taken in the Mediterranean. - Department of Marine Science, University of y-r .., .i ,i . ,■ ^^t■^^■ >., f^^ c u \ri J c^ u . I 171 • 1 oo-m 1 Uutil recently this form was mistaken tor South Florida, St. Petersburg, Florida 33/01 and ^■'^_, , _, , , Museum of Comparative Zoology, Harvard Uni- the yOUng of E. teUsCOpUS. E. crassicaudus \ersity, Cambridge, Massachusetts 02138. is caught by Chilean fishermen. Like E. Bull. Mus. Comp. Zool., 146(3): 147-203, September, 1974 147 148 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 telescopus, it is not taken in sufficient numbers to support a separate fishery but is captured by fishermen trawhng for more abundant deepwater organisms. Although African Epigonus are not pres- ently exploited, tropical eastern Atlantic stocks may represent future sources of pro- tein for mankind. Surveys sponsored by the Organisation of African Unity and the U.S. Agency for International Development revealed these fishes are "of possible po- tential importance ( not necessarily by pres- ent marketing standards) [Williams, 1968: 79]." The same may be true for Caribbean and Gulf of Mexico Epigonus; however, complete data have not been compiled for the latter areas. A major hindrance to the evaluation of deep-sea cardinalfish stocks has been taxo- nomic confusion. The systematic history of Epigonus began in 1810 with Risso's de- scription of Pomatomus telescopus and Rafinesque's account of its synonym Epi- gonus niacrophthahnus. During the fol- lowing seventy-one years, work on the genus was primarily limited to re-descrip- tions of E. telescopus and discussions of its biology (e.g., Cuvier, 1828; Valenciennes, 1830; Capello, 1868; Moreau, 1881). The surge in oceanographic exploration during the last quarter of the nineteenth and beginning of the twentieth centuries rapidly increased the number of nominal Epigonus AAke species. Among the forms described between 1881 and 1920 were Apogon pandionis Goode and Bean, 1881; E. occidentalis Goode and Bean, 1896; Hynnodus atherinoides Gilbert, 1905; Oxij- odon macrops Brauer, 1906; and Hynnodus me galops Smith and Radcliffe, 1912. In the following decade, three new species and two new genera appeared in the liter- ature. Much of the confusion associated with the taxonomy of Epigonus stems from ma- terial described prior to 1930. Early taxa were based on small samples. Because many nations participated in oceanographic research, specimens were deposited in scattered institutions and descriptions ap- peared in diverse publications. Conse- quently, it was difficult for workers to obtain either comparative material or a broad overview of the group's systematics. These shortcomings were aggravated by inaccurate, under-illustrated descriptions based on ill-considered characters. It was common, for example, to use dentition patterns to define generic boundaries, yet tooth arrangements are difficult to observe, easily damaged, and subject to ontogenetic and geographic variation. As a result, an inordinately large number of Epigonu.s-\ike forms was recognized by the end of the 1920's. Although generic taxonomy was stream- lined by Fowler and Bean in 1930 and Matsubara in 1936, species-level taxonomy became increasingly complex. New forms were described in 1935, 1950, 1954, and 1959. In addition, misidentifications of Epigonus were published in several widely circulated works on regional faunas (e.g.. Smith, 1949b and 1961; Gosline and Brock, 1960). The aim of the present study is to clarify the species-level systematics of the Epi- gonus-\ike fishes. Data from traditional characters are evaluated and augmented by information from characters not pre- viously examined for this group. A special effort is made to discuss features such as dentition patterns that caused taxonomic confusion in the past. The ecology, func- tional anatomy, zoogeography, and evolu- tion of Epigonus will be discussed in future works on the genus. METHODS Measurements were made to the nearest tenth of a millimeter by tlie use of Helios needlepoint dial calipers; characters larger than 190 mm were measured with a meter rule or GPM Anthropometer. Measure- ments routinely taken include: Standard length (SL) — from tip of snout to base of caudal fin. EriaoNus Systematics • Mayer 149 Head length (HL) — from tip of snout to tip of opercular spine. Body depth — between dorsal and v(Mitral surfaces of body at level of peKic fin base. Head height — from quadratomandibular joint vertically to bony rim above eye. Eye diameter — between anterior and pos- terior margins of orbit as defined by first and sixth suborbitals. Snout length — from tip of snout to an- terior margin of orbit. Interorbital width — shortest distance be- tween bony rims above eyes. Maxillary length — from tip of snout to posterior margin of maxilla. Lower jaw length — from tip of mandible to quadratomandibular joint. Caudal peduncle depth — shortest dis- tance loetween dorsal and ventral sur- faces of caudal peduncle. Caudal peduncle length — from posterior- most anal fin ray to caudal fin base. First spine length (first spine of first dorsal fin, D,I; first spine of second dorsal fin, DJ; second spine of anal fin, AH; pelvic fin spine, PJ) — from base to tip of spine along anterior edge. Counts were made under a dissecting microscope with the use of dissecting needles or insect pins. A Fibre-Lite High Intensity Illuminator proved invaluable for examinations of oral, branchial, and visceral structures. Gill raker and branchiostegal counts were made on the left side of speci- mens; remaining counts and measurements were made on the right side whenever possible. Counts made include: fin spines (indicated by Roman numerals), fin rays (indicated by Arabic numerals), branchi- ostegal rays, rakers on first gill arch, lateral line scales, pyloric caeca, vertebrae (pre- caudal + caudal, including hypural fan), pleural and epipleural ribs, and basal ptervgiophores between neural spines 9 and 10. Osteological data were obtained from radiographs taken at the Woods Hole Oceanographic Institution, the Museum of Comparative Zoology, and the Harvard University School of Public Health. Holo- types of Oxyodon iiiacrops and Scepterias Icninwn were radiographed at the Zoolo- gisches Museum der Humboldt Universitat and Australian Museum, respectively. More comprehensive osteological studies were based on cleared and stained specimens prepared by trypsin digestion (Taylor, 1967). Osteological terminology follows that presented by Gosline ( 1961 ) and Mead and Bradbury (1963). Suborbital bones are numbered from 1 to 8 beginning with the rostralmost element (lacrimal). Statistical data were analyzed with the use of the Harvard Computation Labo- ratory's IBM 360/65 digital computer. Standard techniques described by Mayr (1969: 189-193) and Simpson et al. (1960: 65-68, 83-88) were employed for analyzing meristic data. Morphometric characters were examined with the aid of regression techniques specified by Simpson et al. (1960: 215-233, 238) and Bailey (1959: 91-99). Before undertaking regression analyses, morphometric data were plotted against SL. Graphs were drawn according to a BMD 05D plotting routine (Dixon, 1967: 71 ) and served as visual tests for linearity of scatter. Only characters exhibiting linear scatters were analyzed by regression tech- niques. As a second precaution against nonlinearity, subadult specimens ( < 40 mm SL) were excluded from statistical samples. Data from several morphometric char- acters are presented both as ratios (i.e., percent of SL or HL) and as regression parameters. The former are intended only as identification aids. As Royce (1957: 17) points out, heterogenic growth makes the use of ratios in fish taxonomy inefficient and may lead to erroneous conclusions. Collection and institution names are ab- breviated as follows in this paper: ABE —Collection of Dr. T. Abe, Tokyo AM — Australian Museum, Sydney 150 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 BMNH — British Museum (Natural History), London BPBM — Bernice P. Bishop Museum, Honohihi CM — Carnegie Museum; collections presently housed in FMNH, Chicago DM — Dominion Museum, Welling- ton EBM — Estacion de Biologia Marina, Universidad de Chile, Viiia del Mar FMNH —Field Museum of Natural History, Chicago IRSN — Institut Royal des Sciences Naturelles de Belgique, Brus- sels ISH — Institut fiir Seefischerei, Ham- burg LACM — Los Angeles County Museum of Natural History, Los Ange- les MCZ — Museum of Camparative Zo- ology, Harvard University, Cambridge MNHN —Museum National d'Histoii-e Naturelle, Paris MZF — Museo Zoologico di Firenze, Florence RUSI — J.L.B. Smith Institute of Ich- thyology, Rhodes University, Grahamstown SAM — South African Museum, Cape Town SMF — Natur-Museum Senckenberg, Frankfurt am Main SU — Stanford University; collec- tions presently housed in the California Academy of Sci- ences, San Francisco TABL — Tropical Atlantic Biological Laboratory, Miami UMML — Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami USNM — National Museum of Natural History, Washington, D.C. UZM — Universitetets Zoologiske Mu- seum, Copenhagen ZMB — Zoologisches Museum der Humboldt Universitiit, Berlin Descriptions are based on material listed by Mayer (1972: Appendix II). Additional data were obtained from examinations of the seventeen specimens listed below. All seventeen fishes were radiographed. E. robustus: ISH 1132/66, 3 specimens, 121.1-142.5 mm, WALTHER HER- WIG Sta. 237/66, 36°00S, 52°58'W, 800 m. ISH 189/71, 9 specimens, 147.0-198.0 mm, WALTHER HER- WIG Sta. 121/71, 37°44S, 54°43'W, 800 m. ISH 269/71, 1 specimen, 147.5 mm, WALTHER HERWIG Sta. 340/ 71, 38°50'S, 54°25'W, 1000 m. ISH 430/71, 1 specimen, 124.1 mm, WALTHER HERWIG Sta. 348/71, 38°20'S, 54°33 W, 997-1040 m. E. fmgilis: LACM 32668-6, 1 specimen, 72.5 mm, 2 mi. off Haleiwa, Oahu, Hawaii, 65 fms. SU 32262, 2 speci- mens, 90.0-93.9 mm, Honolulu, Hawaii. Distributions were determined from material examined and from published accounts. Because of the confusion in Epigomis taxonomy, published data were used only if species identifications could be verified from included descriptions, illustrations, etc. Data from specimens of doubtful identity were not considered. A complete list of station data taken from the literature is provided by Mayer (1972: Appendix II ) . No attempt has been made to provide exhaustive synonymies for Epigonus spe- cies. References are cited only if they (1) are taxonomically or zoogeographically im- portant; (2) provide outstanding descrip- tions, illustrations, or synonymies; or (3) represent verifiable misidentifications. Non- taxonomic accounts have been omitted, as liave references to cruise summaries and faunal lists. SYSTEMATICS Statistical data are presented in tables accompanying species descriptions; meristic Epigonus Systematics • Mayer 151 characters witli low variability are reported ill the text as value, followed in parentheses by number of specimens exiiibiting that v^alue. Meristic and mensural data from holotypes of new species are presented in the Appendix. Genus Epigonus Rafinesque, 1810 Epifiomis Rafinesque, 1810: 64. (Type .species: Epigonus macrophthahnus Rafinesque, 1810 by in()n()t>p\'. A synonym of Pomatomus telescopus Risso, 1810.) Tt'h'scops Bleeker, 1876: 261. (Type .species: Poiuatonniti tclescopiuiu [sic!] Risso, 1810 by original designation. Pomatomus deemed in- applicable. ) Pomatomichthijs Ciglioli, 1880: 20. (Type species: Pomatomiclithys constanciae Giglioli, 1880 by monotypy. A synonym of Pomatomus teles- copus Risso, 1810.) Hynnodiis Gilbert, 1905: 217. (Type species: Hijnnodus athcrinoides Gilbert, 1905 by mono- typy. A synonym of Epigonus occidentaJis Goode and Bean, 1896.) Oxyodon Brauer, 1906: 287. (Type species: Oxyodou Diacwps Brauer, 1906 by monotypy.) Xystramia Jordan, 1917: 46. (Type species: Glossamia pandionis Goode and Bean, 1881 by original designation. Glossamia deemed inapplicable. ) Scepterias Jordan and Jordan, 1922: 44. (Type species: Scepterias fragilis Jordan and Jordan, 1922 by monotypy.) Paraliynnodus Barnard, 1927: 525. (Type species: Parahynnodus robustus Barnard, 1927 by mono- typy- ) Diapiosis. Epip^onus is distinguished from other lower perciform genera by a mosaic of characters including 8 suborbital bones, all lacking subocular shelves; large, thin-walled swimbladders with postero- dorsal ovals; VII or VIII first dorsal fin spines; 1,9 or 1,10 second dorsal fin ele- ments; 11,9 anal fin elements; 15-23 pectoral fin rays; and 17-35 gill rakers. No member of the genus exhibits fang-like conical teeth, such as are found in Cheilo- dipterus, or anteriorly projecting teeth, such as are found in Rosenblattia. Description. Body elongate, fusiform; dorsal and ventral profiles slightly convex, similar. Mouth oblicfue, terminal; upper jaw protrusile; maxilla excluded from gape. sheathed l)y lacrimal anteriorly, free pos- teriorly; supramaxilla absent. Eye large, round or oval. Nostrils paired, rounded or slit-like, two on each side of head. Premaxillae, mandibles, vomer, and pahi- tines edentulous or bearing conical teeth; tongue and endopterygoids rarely dc>ntiger- ous; ectopterygoids edentulous. Gill rakers moderate to long, 17-35; branchiostegal rays 7 (3 + 4); pseudobranchiae present. Opercular .spine either weak, flattened and poorly ossified, or pungent and bony; spine ventral to one or more horny or mem- branous spinelets. Preopercle with double edge; angle frequently produced. Dorsal fins VII-1,9, VII-1,10, or VIII- 1,10, separated by distinct interdorsal space; rudimentary subcutaneous eighth spine present in seven-spined forms. Anal fin 11,9; pectoral fins 15-23; peKic fins 1,5: caudal fin forked, 9 + S principal rays, upper- and lowermost rays unbranched. Scales large, deciduous, ctenoid. Lateral line complete, extending parallel to dorsal profile on dorsolateral surface of trunk, descending to midline on posterior portion of caudal peduncle, continuing on tail; lateral line scales 33-51; canal simple, broadening into deltoid or Y-shaped tube at rear edge of scale. Scale pockets cover- ing most of body including occiput, soft dorsal, anal, and caudal fins; scales absent from snout; no axillary scale at base of PJ spine. Suborbitals 8, all lacking subocular shelves. Vertebrae 25; basapophyses on vertebrae 3 or 4. Predorsals 3, first and second interdigitating between neural spines 2 and 3, third located behind neural spine 3. Caudal skeleton with 2 autogenous haemal .spines, 6 hypurals (hypural 1 = parhypural sensu Monod, 1968), 3 (>purals, 2 (rarely 1) pairs of uroneurals. Actinosts 4, 3VL' borne by scapula. Swimbladder large, thin-walled, lacking anterior or posterior projections to cranium and neural arches; diaphragm absent; oval posterodorsal; retia mirabilia well devel- oped. Stomach U- or Y-shaped; pyloric 152 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 caeca .5-34, may be modified into lu- minescent organs; intestines simple, folded into three segments. Specimens dioecious; no evidence of hermaphroditism or oral brooding. Habitat: Engybenthic; continental slope between approximately 200 and 1200 meters. Key to Species of Epigonvs la Opercular spine weak, poorly ossified, or absent (opercular spine refers to the ventralniost reinforced projection from the posterodorsal edge of the opercle) 2 b Opercular spine pungent, bony 7 2a Lateral line scales 46-51; tongue eden- tulous or bearing scattered tooth patches 3 b Lateral line scales 33-36; tongue cov- ered with tooth patches (Fig. lA) E. oligolepis 3a Gill rakers 23-34; premaxillary teeth short, conical or villifonn, not visible when mouth closed 4 b Gill rakers 17-21; premaxillary teetli elongate, thin, inwardly recurved, visible when mouth closed E. macrops 4a Pyloric caeca 7-14; first dorsal fin VII, rarely VIII; vertebral count 10 + 15; specimens not exceeding 220 mm SL 5 b Pyloric caeca 21-34; first dorsal fin VIII, rarely VII (DiVIII often small or rudi- mentary); vertebral count 11 -|- 14; specimens to 550 mm SL E. telescopus 5a Body long, shallow; depth 15.8-23.6% SL; peduncle length 25.4-32.2% SL; caudal peduncle ring absent 6 b Body short, deep; depth 22.4-29.6% SL; peduncle length 22.0-26.3% SL; caudal peduncle ring present on specimens shorter than 110-120 mm SL (Fig. IB) E. pandionis 6a Gill rakers 28-34; pyloric caeca 10-14; head length 31.2-38.6% SL; 2 pterygio- phores between neural spines 9 and 10, rarely 1 E. denticulatus b Gill rakers 25-26, pyloric caeca 7-8; head length 30.0-34.0% SL; 1 pterygio- phore between neural spines 9 and 10 E. fiagilis 7a Body moderate to deep, 20.0-32.0% SL; dorsal fins VII-1,9, rarely VII-1,10; gill rakers 26-35 8 b Body shallow, 14.0-19.5% SL; dorsal fins VII-1,10, rarely VII-1,9; gill rakers 22-27 — . E. occidentalis 8a Gill rakers of lower arch simple, awl- shaped 9 b Gill rakers of lower arch pectinate ( Fig. IC) - E. pectinifer 9a Tongue edentulous 10 h Tongue covered with tooth patches E. trewavasac 10a Head length 28.0-36.6% SL; head height 14.7-18.8% SL; gill filaments moderate or short 11 b Head length 36.8-41.9% SL; head height 18.9-21.1% SL; gill filaments long ,_ E. crassicaiidiis 11a Fin spines long, DJ 14.8-18.7% SL, All 13.0-20.8% SL; interorbital width 8.7-10.2% SL; eyes large, 40.0-51.1% HL E. leinmen b Fin spines moderate, D2I 10.0-12.6% SL, All 9.2-13.3% SL; interorbital width 6.5-8.2% SL; eyes moderate to small, 37.4-42.2% HL E. robustus Epigonus telescopus (Risso, 1810) Figure 2 Pomatomus telescopus Risso, 1810: 301, plate IX, fig. 31 (original description; Nice; holotype examined, MNHN B862); Lowe, 1841: 173; Capello, 1868: 160; Moreau, 1881: 386, fig. 125; Vaillant (in part), 1888: 376. Epigonus macwphthalmus Rafinesque, 1810: 64 ( original description; no type locality; holotype lost). Pomatomus telescopium Cuvier, 1828: 171 (in- correct emendation of Pomatomus telescopus Risso, 1810); Valenciennes, 1830: 495; Valenciennes, 1837-1844: 6, plate I; Giinther, 1859: 250; Cocco, 1885: 85; Holt and Calder- wood, 1895: 405, plate LXIl. Pomatomus cuvieri Cocco, 1829: 143 (original description; seas of Messina; holotype not examined ) . Pomatotnus cuvicrii \'alenciennes, 1830: 501 (in- correct emendation of Pomatomus cuvieri Cocco, 1829). ?Pomatomichthys constanciae Giglioli, 1880: 20 ( original description; Straits of Messina; holo- type not examined, MZF 3089); Goode and Bean, 1896: 234. Epigonus telescopus Goode and Bean, 1896: 232; Cligny, 1903: 9; Barnard, 1927: 523; Gall, 1931: 1, fig. 1; Fowler, 1936: 736, fig. 326; Smith, 1949b: 206, fig. 474. Scepterias lenimen, Whitley ( in part ) ( not Whit- ley, 1935), 1968: 56. Diagnosis. E. telescopus is the largest species of the genus, growing to over 550 mm SL. Specimens are characterized by 21-34 pyloric caeca and eight first dorsal EriaoNvs Systematics • Mayer 153 B j^i^' ^^.^fffi^^'^h'"**'^^' Figure 1. A. Tongue of E. oligolepis. Stippled areas indicate tooth patches; shape and size of tooth patches may vary among specimens. B. Caudal peduncle of young E. pandionis showing anterior ring and posterior band. C. Gill raker of E. pectinifer showing nub-like processes. fin spines. The opercular spine is blunt and poorly ossified and distinguishes the species from E. occidentalis, E. trewavosae, E. pectinifer, E. rohiistus, E. lenimen, and E. crassicaudus, which have pungent oper- Y12yc cular spines. Unlike remaining congeners, E. telescopus possesses 11 + 14 vertebrae. Description. Meristic data presented in Table 1; regression data for morpho metric traits presented in Table 2. Body thickset, shortened; anterodorsal profile slightly convex, rising most steeply from tip of snout to interorbital region; body moderate to deep, 21.2-26.3%^ SL; caudal peduncle short, 19.0-26.5% SL. Head moderate to deep, height 13.3- SL; length 30..5-37.9% SL; snout blunt; angle of gape moderate to large; lower jaw equalling or protruding slightly beyond upper jaw. Maxilla rarely exceed- ing %-% eye length, posterior margin of maxilla broad, posteriormost point near ventral surface of bone; maxillae of large specimens scaled. Eye round, 49.5-58.9% HL; circumorbital tissues scaled, scale pockets particularly apparent in large spec- imens; anterodorsal rim of orbit projecting into profile in small forms, reaching profile in larger forms; interorbital width 9.0- 10.9% SL. Dentition variable with age (see Onto- genetic change); premaxillae, mandibles, vomer, and palatines dentigerous; tongue edentulous. Opercle bearing short, poorly ossified spine ventral to 1-8 membranous or poorly ossified spinelets; spine and spinelets sepa- rated by shallow gap; spinelets occasionally obscured by underlying membranes. Pre- opercle variable with age; angle rounded, slightly produced in specimens shorter than Figure 2. Epigonus telescopus, 220.0 mm SL, ISH 70/63. 154 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 1. Epigonus telescopus meristic data. X = mean; SD = stan- dard DEVIATION; n = NUMBER OF SPECIMENS. Range SD Pectoral fin rays 20.85 19-23 0.71 54 Gill rakers 24.40 23-26 0.85 52 Lateral line scales 48.14 46-50 1.09 50 Pyloric caeca 25.25 21-34 3.59 16 200 mm SL, broadly produced in larger forms; minute serrations on angle and ven- tral surface of bone, rarely along posterior surface dorsal to angle; striations radiating from inner edge of angle. Interopercles and subopercles without stiiations, occa- sionally bearing minute serrations on pos- tero ventral surfaces. Gill rakers simple, awl-like. First dorsal fin VII (7), VIII (46), eighth spine small or rudimentary, lack- ing membranous connection to preceding spines; second dorsal fin 1,9 (1), 1,10 (52), 1,11 (1); anal fin 11,9 (56); D,I long, 3.5-6.3% SL; DJ, All short, 5.3-9.5%, 5.7-10.6%o SL respectively; Pol moderate, 6.5-11.9% SL. Vertebrae 11 + 14 (18); epipleural ribs Table 2. Epigonus telescopus regression data, b = regression coeffi- cient ± 95%o CONFIDENCE INTERVAL; a = Y INTERCEPT; n = NUMBER OF specimens. All regressions on SL. b a n HL 0. 35 + 0. 01 1. 60 50 Body depth 0. 25 + 0. 01 -2, 43 48 Head height 0. 19 + 0. 00 0. 45 45 Eye diameter 0. 13 + 0. 01 6. 57 44 Snout length 0. 10 + 0. 00 -2. 19 49 Interorbital width 0. 10 + 0. 00 -0, 32 52 Maxillary length 0. 16 + 0. 00 0. 00 48 Lower jaw length 0. 19 + 0. 00 -0. 61 50 Caudal peduncle de pth NONLINEAR Caudal peduncle length 0. 2 1 + 0. 01 3. 35 51 D2I 0. 06 + 0. 02 3. 89 1 1 All 0. 06 + 0. 01 4. 18 31 P2I NONLINEAR Ei'iaoNus Systematics • Mayer 155 Table 3. Ontogenetic changes in the dentition of E. telescopus. A. PREMAXILLARY DENTITION < 200 mm SL Extent 1/2-2/3 of ventral surfoce Pottern I row 200-400 mm SL 2/3-7/8 of ventrol surface 1-2 irregular rows tapering to I row > 400 mm SL 2/3-7/8 of ventral surface Multiple irregular rows B. MANDIBULAR DENTITION Extent < 150 mm SL Entire coronoid surface 150-250 mm SL Entire coronoid surface > 250 mm SL Entire coronoid surface Pottern row 2-3 irregular rows tapering to |-2 rows 3, 4, or 5 irregular rows C. VOMERINE DENTITION Extent < I 75 mm SL Center of vomer > I 75 mm SL Entire face of vomer Pattern Scattered teeth in few irregular rows Numerous teeth in multiple irregular rows D. PALATINE DENTITION Extent < 150 mm SL Length of ventral surface > I 50 mm SL Length of ventral surface Pottern 1-2 irregular rows tapering to I row 2-5 irregular rows tapering to I row 7 (11), 8 (2), inserting on vertebrae 1-7 or 1-8 respectively; pleural ribs 9 (17), inserting on vertebrae 3-11. Large specimens black or brown-violet, iridescent in life ( Risso, 1810; Steindachner, 1891; Dons, 1938). Color in alcohol vari- able with mode of collection and preser- vation; skin often abraded, revealing under- lying white-orange tissue; scale pockets mottled with black or brown, melanophores more densely packed near caudal edges; pigment darker in larger fish; skin oily, cutaneous fat deposits adding rust-colored tint; opercular area black. Guanine de- posits occasionally occurring on opercular, tlioracic, and abdominal regions; iris black with silver highlights; branchial membranes black; mouth darkening with age (see Ontogenetic change ) . Description based on 54 specimens 68.1- 553 mm SL. Ontogenetic change. Several marked ontogenetic changes occur in E. telescopus, the most noticeable involving dentition patterns. Tooth-bearing bones of young specimens exhibit relatively prominent con- ical teeth. Teeth become more numerous witli growth but appear smaller and form weak conical or villiform bands. As Table 3 illustrates, older specimens have more complex tooth patches with larger numbers of tooth rows. 156 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Epigonus Systematics • Mayer 157 A second change involves oral pigmenta- tion. Young individuals have white or pale yellow mouths; melanin is present only in the vicinity of the pharynx. By the time specimens reach 175-225 mm SL, black pigment extends anteriorly to cover the entire tongue. Shortly thereafter, the palate becomes totally blackened, and by 300 mm SL, the entire mouth is dark. The above changes are associated with alterations in intestinal length. Measure- ments of fourteen specimens ranging from 90.7-553 mm SL indicate that intestinal length increases from 66-737^ SL in small specimens (90.7-128.5 mm) to 98-108% SL in moderate-sized individuals (220-250 mm). Thereafter, intestines grow more slowly, reaching 110-115% SL in the largest specimens. The coincidence of rapid in- testinal growth, dentition changes, and de- velopment of oral pigment suggests that E. telescopus modifies its feeding habits with growth. Distribution. E. telescopus has an anti- tropical distribution in the Atlantic, oc- curring from Iceland to the Canary Lslands and reappearing along the western coast of South Africa (Fig. 3). Specimens have also been taken in the Subtropical Con- vergence region east of New Zealand. The species is well known in the western Med- iterranean and has been captured once off the eastern coast of North America. A single specimen is known from shallow water off Norway (Dons, 1938). Adults are taken by bottom trawl or long- line and are most abundant from 300 to 800 meters; however, specimens have been captured from water as shallow as 75 to 80 meters to as deep as 1000 to 1200 meters. Koefoed (1952) reports four pelagic ju- veniles from the Azores; Bertolini (1933) mentions the presence of juveniles in the Tyrrhenian Sea. Earlier workers reported the range of E. telescopus to include St. Helena (Val- enciennes, 1837-1844; Giinther, 1868; Bauchot and Blanc, 1961), tropical west Africa (Osorio, 1898; Poll, 1954; Bauchot and Blanc, 1961), and the Indian Ocean (Steindachner, 1907; Fowler, 1935). These accounts are based on misidentified or tenuously identified material. The .speci- mens described by Giinther, Poll, and Bauchot and Blanc are E. pandionis, while that examined by Fowler is Scomhrops- like. Valenciennes' identification is ba.sed on an impublished description and figure by a St. Helena resident and must be re- garded with suspicion. Reports by Stein- dachner and Osorio could not be evaluated, because neither includes a description or figure of the material studied. Geographic variation. The scarcity of material from South Africa and New Zealand makes it difficult to judge the degree to which Northern and Southern Hemisphere populations of E. telescopus have diverged. Comparisons of dorsal and pectoral fins, pleural and epipleural ribs, lateral line scale counts, gill rakers, and pyloric caeca reveal no subspecific dif- ferences (coefficients of difference ^ 0.44). Moiphometric characters, on the other hand, exhibit greater variability. Of eight traits successfully analyzed, three are sig- nificantly different at both the 95%, 98%, and 99% levels of confidence (Table 4). These differences suggest that northern and southern populations represent gemi- nate subspecies; however, additional ma- terial must be collected, especially from the Southern Hemisphere, before definitive statements can be made on intraspecific variability. Ta.xonomic notes. Pomatomichthtjs con- stanciae Giglioli, 1880 is pro\'isionaily con- sidered a junior synonym of E. telescopus on the basis of work by Tortonese and Queirolo (1970). These authors re-exam- ined and, for the first time, figured the holotype of P. constanciae. The latter species is known only from the type speci- men. The original description (Giglioli, 1880) is incomplete; no adequate rede- scription has ever been published. Data from the papers mentioned above indicate a similarit\' between P. constanciae 158 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 4. Comparison of regression coefficients from Northern and Southern Hemisphere populations of E. telescopus. Data evaluated at the 95%, 98%, and 99% levels of confidence. df = degrees of freedom; nb = regression coefficients of Northern Hemisphere specimens; Sb = regression coefficients OF Southern Hemisphere specimens; SD = significant difference between tab- ular AND calculated VALUES OF t; t ^ CALCtJLATED VALUES OF t. Significance irt CO 05 Nb Sb DF t a> CT> 05 HL 0. 35 0 36 46 1. 54 Body depth 0. 25 0 24 44 0. 82 Head height 0. 20 0 18 41 3. 88 SD SD SD Eye diameter 0. 13 0 12 40 1. 75 Snout length 0. 09 0 10 45 3. 19 SD SD SD Interorbital width 0. 10 0 10 48 1. 05 Maxillary length 0 15 0 . 16 12 0 . 16 Lower jaw len gth 0 19 0 . 19 14 0. 71 Caudal pedunc depth le NONLINEAR Caudal pedunc length le 0 21 0 . 22 47 0. 58 ^2^ INSUFFICIENT DATA All 0 06 0 . 04 23 3. 80 SD SD SD ^2^ NONLINEAR Table 5. Comparison of dorsal and pectoral fin counts from E. telescopus, P. CONSTANCIAE, AND E. TREWAVASAE. DaTA FOR P. CONSTANCIAE FROM GiGLIOLI (1880) AND TORTONESE AND QuEIROLO (1970); REMAINING DATA FROM PRESENT STUDY. E. telescopus P. constanciae E. trewavasae VII VII I, 9. 1,9 rarely I, 10 18 16-18 First VIII, dorsal rarely fin VII Second I, 10, dors a 1 r ar ely fin I, 9 Pectoral fin 19 - 2: Ei'iGONus Systematics • Mayer 159 Figure 4. Epigonus macrops, 154.6 mm SL, USNM 207679. and E. telescopus but also suggest an af- finity between P. constanciae and E. tre- wavasae Poll, 1954. As is shown in Table 5, dorsal and pectoral fin counts fall within the range of E. treicavasae rather than E. telescopus. Tortonese and Queirolo's figure similarly shows the holotype to possess a sharp opercular spine, short DJ, and long PJ — all characteristics of E. treicavasae. Mensural data fail to differentiate P. constanciae from either species. Unlike E. treicavasae but like E. telescopus, the holo- type lacks lingual teeth (Giglioli, 1880). In view of the uncertainty surrounding P. constanciae, a closer study of this form must be undertaken. The problem is all the more pressing, because E. treicavasae is recorded from the Mediterranean for the first time in this paper. Common names. Comprehensive lists of common names for E. telescopus are pro- vided by Doderlein (1889), Nobre (1935), and Bini (1968). Three names not re- corded in these works are "Mejluza" — Gran Canaria ( Steindachner, 1891), "Devil-fish" — North Sea area (Ehrenbaum, 1928), and "Big-eyed cardinal fish" — New Zealand ( Anonymous, 1961 ) . Epigonus macrops (Brauer, 1906) Figure 4 Oxijdon macrops Brauer, 1906: 288, fig. 172 (original description; Indian Ocean, land-locked sea on west coast of Sumatra, VALDIVIA Sta. 186, 03°21'01"S, 101°11'05"E, 903 m; syntype examined, ZMB 17678); Weber and de Beau- fort, 1929: 351, fig. 81; Nomian, 1939: 60. Diagnosis. E. macrops may be distin- guished from all congeners by its low gill raker counts ( 17-21 ) . It is further char- acterized by eight fully developed first dorsal fin spines and eight pyloric caeca, one of which may function as a lumin- escent organ. Description. Meristic values presented in Table 6; regression data for morpho- metric traits presented in Table 7. Body elongate; anterodorsal profile rising steeply to occipital area; thereafter, weakly convex, almost horizontal to first dorsal fin; body depth 19.7-24.1% SL; caudal pe- duncle length 22.0-26.7% SL. Head length 34.1-38.5% SL; head licight 17.2-21.9% SL; snout blunt; angle of gape large; lower jaw protruding beyond upper jaw. Maxilla rarely exceeding Vs-% eye length; posterior margin of maxilla broad, bearing posteriormost point at ventral sur- face of bone. Eye round to oval, 39.7- 48.3%' HL; anterodorsal rim of orbit pro- jecting strongly into dorsal profile; inter- orbital region wide, 9.5-11.7% SL. Teeth conical, frequently recurved. Pre- maxillary and mandibular teeth prominent, needle-like, arranged in single row along length of jaws; mandibular teeth occa- sionally forming double row near sym- physis; vomerine teeth few, moderate, arranged in 2-4 irregular rows or in a triangular or diamond-shaped patch; pala- tin(\s bearing 2-6 teeth, arranged in single row covering anterior half or second quar- ter of bone; tongue edentulous. Opercular spine short, weak, bony, ven- 160 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 6. Epigonus macrops mebistic data. X = mean; SD = standard DEVIATION; n = NLTNIBER OF SPECIMENS. X Range SD Pectoral fin rays 18.87 18-19 0.35 30 Gill rakers 18.63 17-21 0.87 32 Lateral line scales 48.61 46-50 0.83 28 Pyloric caeca 8.00 8 0.00 15 tral to 3-10 spinelets; spine and spinelets separated by shallow, occasionally narrow gap. Preopercular angle weakly produced, rounded, serrate; serrations occasionally ex- tending to posterior and ventral surfaces of bone, rarely absent; striations radiating from inner edge of angle. Subopercle and interopercle generally serrated, occasionally striated. Gill rakers short, awl-like. First dorsal fin VII (1), VIII (29); second dorsal fin 1,9 (1), 1,10 (31); anal fin 11,9 (30), 11,10 (1). D^I, DJ, All short, equalhng 1.2-2.9%, 5.3-7.7%, 5.9- 9.9% SL respectively; PJ moderate, 11.7- 14.1% SL. Vertebrae 10 + 15 (25); epipleural ribs 6 (23), inserting on vertebrae 1-6; pleural ribs 8 (24), inserting on vertebrae 3-10. Table 7. Epigonus macrops regression data, b = regression coeffi- cient ± 95% confidence interval; a = Y intercept; n =: number of specimens. All regressions on SL. b a n HL 0. 35 + 0. 02 1 ± • 48 26 Body depth 0, 22 + 0. 02 0. 51 31 Head height 0. 18 + 0. 02 1. 74 19 Eye diameter 0, 14 + 0. 02 3. 41 29 Snout length 0. 08 + 0. 01 -0. 18 22 Interorbital wi dth 0, 11 + 0. 01 0. 36 30 Maxillary leng th 0. 14 + 0. 01 1. 31 22 Lower jaw leu gth 0, 18 + 0. 01 1. 18 31 Caudal peduncl e dep th 0. 12 + 0. 01 - 1. 53 29 Caudal peduncl e len gth 0. 24 + 0. 02 0. 70 30 D2I 0. 02 + 0. 02 7. 05 13 All 0. 04 + 0. 01 4. 73 22 P2I 0. 13 + 0. 02 0. 49 16 Ei'icoNus Systematics • Mayer 161 Figure 5. Caudal peduncle of young E. macrops bear- ing anterodorsally canted ring. Specimens probably black in life. Color in alcohol variable with preservation; scale pockets covered with black melanophores near posterior edges; skin trecjiiently abraded, revealing pink-yellow muscnla- ture; opercular bones transparent, colored black by underlying branchial membranes; iris black; mouth black in adults. Young bearing anterodorsally canted caudal pe- duncle ring (see Ontogenetic change). First pyloric caecum modified into lumin- escent organ ( see Remarks ) . Description based on 32 specimens 77.8- 206.0 mm SL. Ontogenetic change. The transition from juvenile to adult in E. macrops is marked by changes in pigmentation and body shape. Pelagic juveniles 15-37.9 mm SL and young demersal forms 77.8-79.8 mm SL bear a thin, black, anterodorsally tilted ring circling the center portion of the caudal peduncle (Fig. 5). Specimens larger than 90 mm SL lack this marking. Melanophores forming the rings are deeply embedded in the peduncle musculature and cannot be obliterated by abrading the surface of the fish. Adult E. macrops arc characterized by black oral and branchial membranes. Al- though these areas are colorless or poorly pigmented in specimens smaller than 40 mm SL, the former surfaces darken and the latter become covered with brown melano- phores by the time fish reach 80 nun SL. Juvenile E. macrops appear longer and shallower than adults. Ratio-on-size diagrams for interorbital width (i.e., in- terorbital width/ SL vs. SL) indicate al- lometric growth takes place in small specimens. Similar statements are probably- true for head height, eye length, and caudal peduncle measurements but could not be tested because of damage to juvenile specimens. DistriJnition. E. macrops adults are taken exclusively by bottom trawls between 550 and 1100 meters in the Lidian Ocean, Gulf of Mexico, Caribbean Sea, and West- ern Atlantic. Specimens are most abundant between 640 and 920 meters. Pelagic ju- veniles are known from the Caribbean at depths of 120 to 550 meters (Fig. 6). GcograpJiic variation. No investigation made because of inadec^uate Indian Ocean samples. Taxonomic notes. Brauer's description of Oxyodon macrops ( 1906 ) is based on two syntypes from the eastern Indian Ocean ( 172 and 212 mm total length ) . Of these, only the larger is in the Zoologisches Mu- seum der Humboldt Universitiit; the smal- ler has been lost. The misplaced type may have been deposited in the Zoologisches Institut der Universitiit Leipzig and may reappear when portions of this collection, presently stored in Berlin, are sorted and catalogued (Karrer, personal communica- tion ) . Remarks. Specimens of E. macrops bear eight pyloric caeca; one of these appears modified into a bioluminescent organ. The luminescent caecum arises from the mid- ventral surface of the pylorus just before the duodenum and main body of pyloric appendages (Fig. 7). It extends ventrally until it reaches the floor of the abdominal cavity, bends anteriorly and inserts into a pouch formed by the black peritoneal lining of the body cavity. At the posterior edge of the pelvic girdle, the caecal pouch lies over a thin, translucent portion of the body wall which may function as a biolu- minescent window. Externally the biolumi- nescent window is covered by a single large scale. The caecal pouch is lined with silver or silver-gray pigment. Guanine deposits appear most concentrated anter- odorsally. .\lthouah there is no direct evidence to 162 Bulletin Museum of Coiiiparative Zoology, Vol. 146, No. 3 m in ^— V E T3 O 3 ■D OS TO (0 <0 E (U ■D H— o "5 CO JC "cO 3 T3 > T3 C If) 3 to Q. O O *^ TO o m 130 mm SL) have as many as 3-4 tooth rows on palatines and anterior segments of dentaries and premax- illae. Vomerine teeth may become suf- ficiently numerous to cover the entire face of the bone. Distribution. E. pandionis is amphi-At- lantic, occurring primarily in the Caribbean, Gulf of Mexico, and Gulf of Guinea (Fig. 9). The species has been taken as far north as New Jersey and as far south as French Guiana in the western Atlantic. It occurs between Portuguese Guinea and Angola in the eastern Atlantic. Adults are captured exclusively by bottom trawls be- tween 210 and 600 meters. American forms are most numerous from 300 to 500 meters, while African populations are most abun- dant between 260 and 450 meters. A single pelagic juvenile (35.5 mm SL, MCZ 48839) was taken at 275 to 300 meters in the Caribbean. Geo<:,raphic variation. Statistical analyses provide conflicting assessments of the similarity of African and American popula- tions. Meristic characters reveal little vari- ability. Coefficients of difference calculated for standard counts are always less than or equal to 0.49 — far below conventional levels of subspecies recognition. Mensural data, on the other hand, suggest there are considerable differences between the pop- ulations. Of thirteen traits analyzed, seven separate eastern and western populations at the 95% level of confidence, five separ- ate them at the 98% level, and two separate them at the 99% level (Table 10). A closer examination of the characters exhibiting signilicant differences reveals that regression coefficients of American E. pandionis are always greater than those of African forms. Since regression coef- ficients are a measure of relative growth, observed intraspecific variation may reflect environmental factors. Water temperature is a major parameter determining growth rates in fishes. If other factors are conti'olled, rates of growth in- crease proportionally with temperature (Brown, 1957: 391). With this in mind, it is interesting that temperatures are gener- ally higher and superficial warm-water layers thicker in the western tropical At- lantic (Ekman, 1953). At 300 meters Gulf of Mexico and Caribbean temperatures vary from 10 to 18° C while west African tem- peratures range between 9 and 11° C. At 500 meters the difference is slightly less pronounced — 8-13° C as opposed to 6-8° C (from temperatin-e profiles in Fuglister, 1960; Wiist, 1964; and Nowlin and McLellan, 1967). One would therefore expect western Atlantic E. pandionus to grow more rapidly and exhibit larger re- gression coefficients than eastern Atlantic forms. In view of these findings, the two morphs are not considered to represent separate subspecies. Remarks. See E. trewavasae: Remarks for discussion of E. pandionis .sensu Lozano (1934), Navarro et al. (1943), and Maurin (1968). Specimens of doubtful identity. Five spec- imens were examined that resembled E. pandionis but could not, with certainty, be placed in the species. Four were taken in the Atlantic, the fifth in the Gulf of Oman (see Mayer, 1972: Appendix II for complete data). These fishes were not considered when preparing the description of E. pandionis, nor were they used in morphometric, meristic, or distribution analyses. Tlie Atlantic specimens include three 168 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 10. Comparison of regression coefficients from eastern and %vestern Atlantic populations of E. pandionis. Data evaluated at the 95%, 98%, and 99% LE\rELS OF confidence. DF =: DEGREES OF FREEDOM; Eb = REGRESSION COEFFI- CIENTS OF EASTERN ATLANTIC SPECIMENS; SD = SIGNIFICANT DIFFERENCE BETWEEN TABULAR AND CALCULATED VALUES OF t; t=3 CALCULATED VALUES OF t; Wb = REGRES- SION COEFFICIENTS OF WTESTERN ATLANTIC SPECIMENS. Significance Wb Eb DF t S S S HL 0. 36 0. 37 60 0. 97 Body depth 0. 29 0. 29 71 0, 31 Head height 0. 22 0. 20 63 2. 41 SD SD Eye diameter 0. 17 0. 15 76 2. 14 SD Snout length 0. 09 0. 08 69 3. 00 SD SD SD Interorbital width 0. 11 0, 11 70 2. 13 SD Maxillary leng ^th 0. 18 0. 16 70 3. 35 SD SD SD Lower jaw len gth 0. 20 0. 18 74 2. 63 SD SD SD Caudal pedunc depth le 0. 13 0. 12 76 2. 39 SD SD Caudal pedunc length le 0. 23 0. 24 77 1. 16 D2I 0. 05 0. 06 32 1 83 All 0. 05 0. 05 52 0 91 P2I 0. 10 0. 10 71 0 83 fishes from St. Helena. The most recently for only the Caribbean form, which was collected (UZM P45148) was incorrectly taken at relatively shallow depths. Exact identified as E. telescopus by Banchot and determination of the variants' status must Blanc (1961). The two older fonns (BMNH await the capture of additional material. 1868.3.11.14/15) are probably the fish The Indian Ocean form (BMNH discussed by Giinther (1868). The re- 1889.4.15.24) is distinguished from E. maining specimen (USNM 207703) was pandionis by its shallow body (22.5% SL), taken in the Caribbean. narrow interorbital region (8.3% SL), den- The four Atlantic individuals are basi- tigerous glossohyal, numerous weak oper- cally similar to E. pandionis but exhibit cular spinelets, and elongate gill filaments, shallower heads ( 17.4-19.8% SL), narrower The last trait suggests the fish may have interorbital regions (8.6-9.4% SL), fewer inhabited an oxygen minimum layer. As pyloric caeca (8-9), and fewer gill rakers with the Atlantic variants, additional ma- ( 25-27 ) . In these respects they resemble terial must be collected before the status of E. fragilis. Little is known about the habits the form can be determined, of the variants; station data are available Common names. None. Epigonus Systematics • Mayer 169 Z^A ^ y y x» Figure 10. Epigonus fragilis, HOLOTYPE, 89.1 mm SL, CM 3900/FMNH 55204 (from Jordan and Jordan, 1922). Epigonus fragilis (Jordan and Jordan, 1922) Figure 10 Scepterias fragilis Jordan and Jordan, 1922: 45, plate II, fig. 2 (original description; Honolulu market; holotype examined, CM 3900/FMNH 55204). ?Hynnodus fragilis Pietschmann, 1930: 13. Diagnosis. E. fragilis most closely re- sembles E. pandionis but may be dis- tinguished by its shallow body (18.8-21.1% SL) and short, shallow head (length 31.7- 34.0% SL, height 16.0-17.4% SL). Unlike £. pandionis, E. fragilis lacks peduncle rings on specimens smaller than 100-120 mm SL. In the past E. fragilis has been confused with Hijnnodus atherinoides, a junior syn- onym of E. occidentalis. E. fragilis may be distinguished on the basis of body depth ( see above ) , pectoral fin counts ( 16-17 ) , and the absence of a pungent, bony oper- cular spine. Weak opercular armor, to- gether with second dorsal fin counts of 1,10 differentiate E. fragilis from E. treivavasae, E. pectinifer, E. rohustus, E. lenimen, and E. crassicaudus. Gill raker counts of 25-26 separate E. fragilis from all remaining con- geners except E. telescopiis. E. fragilis may be distinguished from the latter by the presence of 7-8 pyloric caeca. Description. E. fragilis is known from only five specimens. Of these, the holotype is of little descriptive value. The specimen is severely dehydrated and has become discolored, brittle, and shrunken. The fol- lowing account is based primarily on two recently captured specimens of E. fragilis (LACM 32668-6 and USNM 207704) and two forms collected by D. S. Jordan in 1921 (SU 23246). The latter are mentioned in the original description of E. fragilis but are not designated as types. All meristic and mensural data are pre- sented in the text. Detailed statistical analyses were not undertaken because of small sample size. Body elongate; anterodorsal profile con- vex, rising without interruption from tip of snout to first dorsal fin. Body depth 18.8-21.1% SL; caudal peduncle length 25.4-26.9% SL. Head short, 31.7-34.0% SL; head height 16.0-17.4% SL; snout blunt, 7.2-7.9% SL; angle of gape moderate; jaws equal. Max- illa reaching % eye length; posteriormost point of maxilla at ventral edge of bone. Eye round, 38.1—41.5% HL; anterodorsal rim of orbit reaching profile; interorbital width 8.8-9.4% SL. Dentition variable with age. Teeth con- ical; premaxillary teeth in irregular double rows anteriorly, tapering to single row posteriorly, occupying anterior %-% of bone. Mandibular dentition more promin- ent than that of premaxilla; teeth recurved, occupying from % to entire length of dentary, arranged in single or double rows near symphysis and single row posteriorly. Vomerine teeth recurved, arranged in oval or diamond-shaped patch, covering entire face of bone in adults. Palatine teeth 170 Bulletiti Museum of Comparative Zoology, Vol. 146, No. 3 Figure 11. Epigonus occidentalis, 152.7 mm SL, MCZ 48840. medially recurved, arranged in single-triple rows anteriorly, tapering to single row posteriorly; tongue edentulous. Opercular spine weak, ventral to 7-9 small serrae; angle of preopercle produced, rounded, ornamented with striations and weak serrations; subopercle and inter- opercle unornamented. Gill rakers 25 (3), 26 (1), simple, awl-like. Pyloric caeca 7(1), 8(2). First dorsal fin VII (4), VIII (1); second dorsal fin 1,10 (5); anal fin 11,9 (5); pectoral fin 16 (1), 17 (3); DJ moderate to long, 5.9-8.9% SL; DJ short, 6.9% SL; PJ long, 10.1-10.2% SL; All broken. Vertebrae 10 + 15 (4); epipleural ribs not visible on radiographs; pleural ribs 8 (4), inserting on vertebrae 3-10. Pored lateral line scales 49 ( 2 ) . Color in alcohol yellow-brown; fin mem- branes dark; iris silver-black; mouth light; branchial membranes light, darkening with age. Distribution. E. fragilis is endemic to the Hawaiian Islands (Fig. 12). The spe- cies is demersal and has been taken between 120 and 125 meters. Taxonomic notes. Six years after E. fragilis was described, Fowler ( 1928 ) syn- onymized the species with a second Ha- waiian apogonid, Hijnnodus atherinoides Gilbert, 1905. The synonymy achieved moderate acceptance and appeared in sev- eral publications (e.g., Matsubara, 1936; Tinker, 1944; Gosline and Brock, 1960). Fowler's conclusions were based on a 33-mm specimen (BPBM 3914) obtained by the Tanager Expedition. The specimen is in extremely poor condition. All colora- tion has been lost, most of the muscle tissue has decomposed, and much of the skeleton has become decalcified. Although it is impossible to identify the fish because of its condition, the following traits suggest it is neither E. fragilis nor H. atherinoides: dorsal fin elements — VIII-1,8; anal fin ele- ments— 11,6; vertebrae — 11 + 14. These data differ from Fowler's report of VI-I,8 dorsal elements, no anal spines, and 7 anal rays. As was discussed in the diagnosis, E. fragilis is distinct from H. atherinoides. Fowler's synonymy appears to have been based on inaccurate data taken from an incorrectly identified fish. Common names. None. Epigonus occidentalis Goode and Bean, 1896 Figure 11 Epigonus occidentalis Goode and Bean, 1896: 233, plate LXVI, fig. 236 (original description; Steamer BLAKE, off Barbados, 237 fms.; holo- type examined, MCZ 28032 ) . Hijnnodus atherinoides Gilbert, 1905: 618, plate 79 (original description; ALBATROSS Sta. 3867, Pailolo Channel, Hawaii, 284-290 fms.; holotype examined, USNM 51601); Jordan and Jordan, 1922: 44; Fowler and Bean, 1930: 121. Hijnnodus megalops Smith and Radcliffe, 1912 {in Radcliffe, 1912): 445, plate 38, fig. 3 (original description; ALBATROSS Sta. 5388, 12°51'30"N, 123°26T5"E, between Bnrias and Luzon, Philippines, 226 fms.; holotype ex- amined, USNM 70255). Ei'iGONus Systematics • Mayer 171 Table 11. Epigonus occidentalis meristic data. X = mean; SD STANDARP nEVIMION; n = NUMBER OF SPECIMENS. X Range SD Pectoral fin rays Gill r aker s Lateral line scales Pyloric caeca 20. 21 19-2 1 0. 59 56 24. 68 22-27 1. 08 60 48. 15 46-51 0. 97 46 9. 27 8-13 1. 05 45 Table 12. Epigonus occidentalis regression data, b = regression COEFFICIENT ± 95% CONFIDENCE INTERVAL; a = Y INTERCEPT; n = NUMBER OF SPECIMENS. AlL REGRESSIONS ON SL. b a n HL 0. 34 + 0. 02 0. 72 48 Body depth 0, 19 + 0. 02 -1. 72 48 Head height 0. 15 + 0. 01 0. 53 49 Eye diameter 0. 16 + 0. 01 0, 66 49 Snout length 0. 08 + 0. 00 0. 06 49 Interorbital width 0. 08 + 0, 01 0. 83 39 Maxillary length 0. 13 + 0, 01 0. 88 51 Lower jaw length 0, 15 + 0. 01 1. 26 51 Caudal peduncle de pth 0, 10 + 0. 01 -0. 90 54 Caudal peduncle len igth 0. 23 + 0. 01 1. 50 53 D2 I 0. 05 + 0. 00 1. 42 34 All 0. 05 + 0. 01 2. 18 42 P2I 0. 09 + 0. 01 0. 67 47 Diagnosis. E. occidentalis is distin- guished from all other congeners by the combination of shallow body depth (14.1- 19.57t SL), reduced gill raker counts (22- 27), and the presence of a pungent, bony opercular .spine. It is frequently confused with E. denticulatus. Description. Meristic values presented in Table 11; regression data for morpho- metric traits presented in Table 12. Body elongate, cigar-shaped; anterodor- sal profile weakly convex, flattened, rising gradually from tip of snout to interorbital region, leveling off toward occipital region, and rising gradually to base of first dorsal fin. Body depth 14.1-19.5% SL, body 172 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 width subequal to or greater than body depth; caudal peduncle narrow, length 22.4-28.1% SL. Head length 30.5-37.9% SL; head height 13.3-17.2% SL; angle of gape moderate to small; lower jaw equalling or protruding slightly beyond upper jaw. Maxilla reach- ing Vi-% eye length; posterior margin of maxilla moderate to narrow, posterior- most point at ventral edge of bone. Eye long, oval, 40.6-52.3% HL; anterodorsal rim of orbit reaching or projecting into dorsal profile; interorbital region narrow, 5.6-8.5% SL. Teeth conical; premaxillary and man- dibular teeth frequently recurved, arranged in simple single row or single row widening to double or triple rows near symphysis; teeth covering % to entire length of pre- maxilla and % to entire length of dentary; vomerine teeth arranged in 1-4 irregular rows; palatines rarely edentulous, teeth 1-10, arranged in single row, covering anterior Vi-V^ of bone; tongue edentulous. Opercular spine pungent, bony, ventral to 1-3 poorly ossified spinelets; spine sep- arated from spinelets by shallow indentation. Preopercular angle produced, rounded or pointed, bearing serrations and striations; subopercle serrate, occasionally striate; interopercle variable, frequently serrate. Gill rakers short, awl-like. First dorsal fin VII (59); second dorsal fin 1,10 (59); anal fin 11,8 (1), 11,9 (59); DJ, DJ, All, PJ short, equahing 1.1^.2%, 4.8-7.8%, 4.8-9.2%, and 8.0-11.3% SL re- spectively. Vertebrae 10 -I- 15 (35); epipleural ribs 6 (19), 7 (5), inserting on vertebrae 1-6 or 1-7 respectively; pleural ribs 7 (31), 8(1), inserting on vertebrae 2-9 or 3-9. Color in alcohol variable with preser- vation; skin frequently removed by trawl- ing; underlying tissue pale yellow, yellow- pink, occasionally marked with rust brown; scale pockets and fin membranes black; opercular area black-slate gray, occasion- ally tinged with silver; lower jaw, bran- chiostegal membranes, and thoracic and abdominal regions occasionally silvered; guanine most prevalent on specimens from old collections. Mouth color variable with age (see Ontogenetic change); iris and branchial region dark. Description based on 62 specimens 58.2- 178.9 mm SL. Ontogenetic change. The most striking age-related change in E. occidentalis is the development of oral pigmentation. As in E. telescopus and E. macro'ps, immature forms bear pigmentless or slightly pig- mented mouths, while adults have black- ened oral membranes. Pigmentation first appears in specimens 80-110 mm SL. Melanophores develop just anterior to the pharynx and spread rostrally, covering a third of the roof and floor of the mouth and half of the tongue by the time specimens reach 115-130 mm SL. By 150 mm SL the tongue is completely black, and by 175- 180 mm the entire mouth is dark. Branchial membranes undergo an analogous trans- formation before specimens reach 58 mm SL. A faint black ring circling the middle of the caudal peduncle was observed on three small E. occidentalis (< 65 mm SL). Similar markings were absent from larger individuals. The rings are reminiscent of markings observed on young E. macrops and E. pandionis and probably represent a juvenile feature that is lost with growth. Distribution. E. occidentalis has been taken in the Caribbean, Gulf of Mexico, and western tropical Atlantic. It is also known from the Philippine and Hawaiian Islands (Fig. 12). The species is caught by bottom trawls between 360 and 735 meters. Adults are most abundant in the Caribbean from 500 to 550 meters. Geographic variation. E. occidentalis, as here defined, includes two nominal species — Hijnnodus atherinoides Gilbert, 1905 and H. megalops Smith and Radcliffe, 1912. The former originally represented a Ha- waiian endemic; the latter represented a Philippine form. In 1930 Fowler and Bean synonymized the Pacific morphs. In the Epigonus Systematics • Maxjer 173 Figure 12. Distribution of E. fragilis and E. occidentalis. Map A shows localities in the Caribbean and Gulf of Mexico. Map B shows localities in the western Pacific. J^ E. fragilis, individual haul of demersal adults; % E. occidentalis, indi- vidual haul of demersal adults; cross-hatching indicates areas where E. occiden- talis are frequently taken. 174 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 3 cm Figure 13. Epigonus denticulatus, 115.1 mm SL, UMML 12463. authors' opinions, characters separating the two forms were "simply minor discrepancies of portraiture and should never have been credited as specific distinctions [p. 122] ." Although descriptions and illustrations of H. atherinoides and H. megalops suggest a link with E. occidentalis, detailed com- parisons of the three forms were never made. To a large extent this was the re- sult of inadequate sampling. Until the initiation of the OREGON cruises in 1950, few E. occidentalis were available for study. Pacific forms are still poorly represented; only seven specimens have been collected. Reports of additional material by Fowler (1928), Matsubara (1936), Smith ( 1949a,b, 1961), Kamohara (1952), and Moreland ( 1957) are based on misidentifications. Comparisons of E. occidentalis and the H. atherinoides-H . megalops complex pro- vide no evidence to support their status as separate species. Analyses of head length, body depth, head height, eye di- ameter, snout length, interorbital and max- illary widths, caudal peduncle length and depth, and All and PJ lengths reveal no significant differences between the pop- ulations at either the 95%, 98%, or 99% levels of confidence. Meristic data also show considerable overlap for most char- acters; however, the coefficients of dif- ference for pyloric caeca and gill raker counts are above conventional levels of subspecies recognition (1.68 and 1.99, re- spectively). In addition, Atlantic and Pacific populations may be distinguished by minor qualitative characters such as: (1) short, rounded preopercular angles in Atlantic forms; longer, pointed angles in Pacific specimens; (2) fusion of uroneurals 1 and 2 in Atlantic forms (based on 3 alizarin preparations ) ; separate occurrence in Pacific forms (based on 1 alizarin preparation ) . On the basis of the above information, Atlantic and Pacific forms are placed in the same species but considered members of separate subspecies. Formal description of the subspecies must await the capture of additional Pacific specimens. Remarks. A single unripe female E. occidentalis (USNM 197353, 172.1 mm SL) was found carrying small egg masses in the anterior portion of its mouth (anterior to the tongue and vomer). The masses con- tained 125 oval eggs 0.40-0.55 mm in diameter. The presence of eggs in the mouth of an Epigonus is of interest, be- cause several shallow-water apogonids ex- hibit oral brooding. No such activity has ever been reported for deep-sea forms. Although it is difficult to say with certainty, the E. occidentalis eggs are prob- ably not incubating clutches, but rather non-apogonid ova ingested during trawling. Unlike the egg masses of typical oral brooding apogonids, those found in E. oc- cidentalis are broken, disrupted, and con- tain very few eggs. An 84.9-mm specimen of Cheilodipterus affinis was reported in- Epigonus Systematics • Mayer 175 Table 13. Epigonus denticulatus meristic data. X = mean; SD STANDARD DEVIATION; n = NUMBER OF SPECIMENS. X Range SD n Pectoral fin rays 19. 09 18-20 0. 56 54 Gill rakers 30. 98 28- 34 1. 10 58 Lateral line scales 48. 12 46- 49 0. 76 43 Pyloric caeca 11. 83 10- 14 0. 85 42 ciibating 21,000 eggs 0.35-0.4 mm in di- ameter (Smith et al., 1971: 8-9). The ova fully occupied the oral and branchial chambers and extensively distended the head. These conditions were not observed in E. occidentalis. It is possible that the eggs represent the remnants of a larger mass that was spit out and partially reingested. However, were this the case, one might expect to find eggs in the stomach (Sakomoto, 1930) or gill rakers. No eggs were found in either region. Finally, Breder and Rosen (1966) state that eggs of oral brooding apogonids are lield together by fibers attaching to one pole. The eggs of E. occidentalis are loosely embedded in an open matrix of fibers. Grape-like egg clusters character- istic of Apogon semilineatus (Ebina, 1931: 20 ) were not observed. Common names. None. Epigonus denticulatus Dieuzeide, 1950 Figure 13 Pomatomus telescoptis, Vaillant (in part) (not Risso, 1810), 1888: 376. Scepterias lenimcn, Whitley (in part) (not Whit- ley, 1935), 1935: 230; Whitley (in part), 1940: 420. Epigonus atherinoides, Matsubara (not Gilbert, 1905), 1936: 120, fig. lA; Smith, 1961: 378, fig. 3; Kamohara, 1952: 37, fig. 31. Hynnodus atherinoides. Smith (not Gilbert, 1905), 1949a: 101; Smith, 1949b: 210, fig. 495A. Epigonus denticulatus Dieuzeide, 1950: 89, figs. 1-2 (original description; Algerian Coast at 200-500 m; holotype not examined); Tortonese, 1952: 72, 1 fig.; Dieuzeide et al., 1953: 216, 2 figs.; Tortonese and Queirolo, 1970: 33, fig. 6. Diagnosis. E. denticulatus lacks a fully ossified opercular spine, bearing instead 3-7 membranous projections. This feature distinguishes it from E. occidentalis, E. treivavasae, E. pectinifer, E. rohustus, E. lenimen, and E. crassicaudtis, which have pungent, bony opercular spines. E. denti- culatus is differentiated from E. telescopus, E. macrops, and E. fragilis by the presence of 10-14 pyloric caeca and 28-34 gill rakers. It differs from E. oligolepis by bearing 46-51 lateral line scales. E. denti- culatus closely resembles E. pandionis but may be distinguished on the basis of the former's shallow body (15.8-23.67r SL), long caudal peduncle (25.9-32.2% SL), and short DJ (2.4-3.7% SL). Description. Meristic values presented in Table 13; regression data for morpho- metric traits presented in Table 14. Body fusiform, slightly compressed; an- terodorsal profile rising gradually above snout, becoming steeper and slightly con- vex over eyes, thereafter rising gradually to first dorsal fin; body moderate to shal- low, depth 15.8-23.6% SL; caudal peduncle narrow, length 25.9-32.2%o SL. Head moderate to short, 31.2-38.6% SL; head height 16.0-19.8% SL; snout short, blunt; angle of gap(> moderate to large; lower jaw protruding slightly beyond up- per jaw. Maxilla reaching %-% eye length, 176 Biilletifi Museum of Comparative Zoology, Vol. 146, No. 3 Table 14. Epigonus denticulatus regression data, b = regression COEFFICIENT ± 95% CONFIDENCE INTERVAL; a = Y INTERCEPT; 11 = NUMBER OF SPECIMENS. AlL REGRESSIONS ON SL. HL 0. 32 + 0. 01 2. 88 57 Body depth 0. 25 + 0. 01 -4. 09 54 Head height 0. 16 + 0. 01 0. 86 56 Eye diameter 0. 14 + 0. 01 1. 39 58 Snout length 0. 07 + 0. 00 0. 33 56 Interorbital width 0. 09 + 0. 00 0. 37 55 Maxillary length 0. 14 + 0. 01 1. 37 56 Lower jaw length 0. 15 + 0. 01 1. 61 57 Caudal peduncle d epth 0. 11 + 0. , 01 -0. 78 57 Caudal peduncle li ength 0. 28 + 0. , 01 0. 61 57 D2I 0. 05 + 0. , 01 2. 24 36 All 0. , 06 + 0, , 01 1. , 43 40 P2I 0. , 08 a. 0, , 01 0. , 92 41 posteriormost point near ventral surface of bone. Eye round or slightly oval, 40.3- 48.0% HL; anterodorsal rim of orbit reaching dorsal profile, projecting into pro- file in smaller specimens; interorbital width 8.2-10.4% SL. Teeth small, conical, occasionally re- curved; premaxilla bearing single row of teeth along anterior Vs-% (usually %) of bone. Mandibular teeth arranged along length of dentary in irregular single row, occasionally double near symphysis; larger specimens with 3-4 rows near symphysis. Vomerine teeth variable, arranged in 1-4 irregular rows. Palatine dentition occupy- ing length of bone, arranged in simple single row or double row tapering to single row posteriorly; large specimens bearing 3-4 rows of teeth anteriorly. Tongue gen- erally edentulous, rarely Ijearing isolated tooth patches on glossohyal or edges of tongue. Opercle lacking bony spine, bearing in- stead 3-7 (usually 5-6) jagged, mem- branous projections; projections often ob- scured by underlying tissues. Peropercular angle produced, broadly rounded, striations radiating from inner edge, angle occasion- ally serrate; subopercle and interopercle occasionally serrate. Gill rakers simple, awl-like. First dorsal fin VII (53); second dorsal fin 1,9 (1), 1,10 (56), 10 (1); anal fin 11,8 (1), 11,9 (57). DJ moderate, 2.4-3.7% SL; Dol, All, P,I short, 5.2-8.0%, 6.0-8.2%, 7.9-10.0% SL respectively. Vertebrae 10 + 15 (44); epipleural ribs 6 ( 32 ) , 7 ( 1 ) , inserting on vertebrae 1-6 or 1-7 respectively; pleural ribs 8 ( 44 ) , inserting on vertebrae 3-10. Color in alcohol variable with preserva- tion; skin frequently removed by trawling, underlying tissue pink-brown or yellow; scale pockets mottled with numerous brown-l)lack melanophores, dorsal surfaces of body and head more heavily pigmented. Epigonus Systematics • Mayer 177 Guanine deposits frequently occurring on gill cover, ventral surface of mandible, isthmus, thoracic region, and abdomen to anus; iris black; mouth light; branchial region dark. Description based on 58 specimens 57.0- 187.5 mm SL. Ontogenetic change. Two young spec- imens of E. dcnfictihifus (29.2 mm SL, MCZ 48846, and 49.7 mm SL, MCZ 48847) were examined in the course of this in- vestigation. These specimens were taken by midwater trawls made in the central North Atlantic and Gulf of Mexico and reveal that the life cycle of E. denticulatus includes a pelagic juvenile stage. The pelagic young resemble adults in most respects. For example, the juveniles bear diagnostic gill raker counts and opercular ornamentation. However, slight changes in body shape are associated with growth. The 29.2 mm specimen has a more shallow body, shorter head, narrower inter- orbital region, and smaller eyes than demersal adults. Similar trends are present but less apparent in the larger juvenile. Juvenile dentition patterns are basically like those of adults but involve fewer and relatively larger recurved teeth. Oral and branchial regions are light in young speci- mens. The latter areas darken with age. Distribution. E. denticulatus is the only cosmopolitan species in the genus (Fig. 14). Specimens have been taken from the southwest coast of Japan, the Gulf of Mexico, and the Caribbean. In addition, the species occurs continuously from the western Mediterranean, south along the western coast of Africa to the tip of the continent. It reappears south of the Great Australian Bight and southeast of New Zealand. Adults are generally taken by bottom trawls between 300 and 600 meters, al- though specimens have been captured from as shallow as 200 meters and as deep as 830 meters. Pelagic juveniles have been taken by IKMT between 130 to 145 meters and 350 to 425 meters. Geographic variation. E. denticuhitus may be divided into North Atlantic, South- ern Hemisphere, and Japanese populations. North Atlantic forms include material from the Mediterranean, northeast Atlantic, Caribbean, and Gulf of Mexico. Southern Hemisphere populations contain specimens from the southeast Atlantic, Australia, and New Zealand. Statistical analyses reveal surprisingly little divergence between North Atlantic and Southern Hemisphere specimens. Co- efficients of difference for standard meristic characters are far below accepted levels for subspecies recognition (all are ^ 0.53), and regression coefficients for mensural data are virtually identical. Only maxil- lary lengths differ significantly at the 95% level of confidence. It is clear from the data that North Atlantic and Southern Hemisphere E. denticulatus do not repre- sent separate subspecies. Detailed analyses of the Japanese pop- ulation could not be undertaken because of inadequate sampling. Only one speci- men was available from the area. On the basis of this fish, the Japanese population appears closely allied to the rest of the species. With the exception of eye di- ameter, standard counts and measm-ements made on the Japanese morph fall within the 95% and 99% confidence intervals of remaining E. denticulatus. Eye diameter falls outside the 95% confidence interval but within the 99% confidence interval. The similarity of E. denficuhiius pop- ulations, despite the wide rangc> of the species, suggests ( 1 ) there may be con- siderable gene flow between populations, (2) the present distribution may have been achieved only recently, or (3) evolution is occurring very slowly. Discovery of a pelagic juvenile in the mid-North Atlantic gives credence to the first hypothesis and proN'ides a mechanism for the dis- persal of a species with demersal adults such as E. denticulatus. Common names. "Castagnera briina" in Monaco (Bini, 1968). 178 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 W E o o Q. a> m E O) Q. 3 CO CO 3 T3 T3 C o 3 CO V E 3 CO 3 •a '> ■D c to o c c o 3 10 b ii. Epigonus Systematics • Mayer 179 Epigonus oligolepis sp. nov. Figure 15 llolotype: One specimen, 90.8 nun SL, taken from the Straits of Florida bv M/V COMBAT, Sta. 436: 21 July 1957, 1319 to 1530 hrs.; 24°13'N, 81°42'W; 300 fnis., 10' flat trawl. USNM 207718. Parat>pes: One specimen, 126.7 mm SL, M/V OREGON, Sta. 4731: 27 February 1964; 27°35'N, 92°32'W; 250-300 fms.; 40' flat trawl. MCZ 48848. Three specimens, 52.7-72.7 mm SL, Steamer ALBATROSS, Sta. 2643: 9 April 1886; 25°25'00"N, 79°55'15"W; 211 fms. USNM 109430. Three specimens, 53.7-84.2 mm SL, M/V OREGON, Sta. 5043: 26 September 1964; 12°01'N, 6P53.5'W; 210-250 fms.; 40' shrimp trawl. USNM 207719. One specimen (cleared and stained), 62.0 mm SL, locality data identical with those of preceding lot. USNM 207720. One specimen, 117.1 mm SL, M/V OREGON, Sta. 3741: 26 August 1962; 29°10'N, 88°01.5' W; 300-340 fms.; 100' flat trawl. USNM 207721. Diagnosis. E. oligolepis is distinguished from all congeners by lateral line scale counts of 33-36 and the presence of lingual and endopterygoid teeth. Description. Meristic values presented in Table 15; regression data for morphometric traits presented in Table 16. Body elongate, moderately compressed; anterodorsal profile rising gradually from tip of snout to interorbital region, rising more steeply and becoming slightly convex to occiput, thereafter rising gradually to base of first dorsal fin; body depth 19.8- 24.5% SL; caudal peduncle length 23.9- 27.2% SL. Head moderate to long, 34.4-43.0% SL; head height 16.6-18.8% SL; snout pointed; angle of gape moderate; lower jaw pro- truding beyond upper jaw. Maxilla reach- ing %-% eye length; posterior margin of maxilla rounded, posteriormost point between midline and ventral margin of bone. Eye round to slightly oval, 40.1- 43.77o HL; anterodorsal rim of orbit reach- ing or projecting into dorsal profile; inter- orbital width 8.5-9.6% SL. Teeth small, conical; premaxilla edent- ulous or bearing few teeth on anterior Vi- -f. of bone; mandibular teeth arranged in single or double row antericnly, single row posteriorly; teeth covering anterior half of bone and occasionally extending along length of dentaiy. Vomer covered with irregular tooth patches, teeth extending posteriorly along midline of palate; pala- tine teeth arranged in single or multiple rows anteriorly, single row posteriorly, covering from half to entire length of bone; endopterygoid dentigerous; auxiliary tooth patches occurring between vomer, pala- tines, and endopterygoids; tongue den- tigerous, bearing lateral and glossohyal tooth patches (Fig. lA). Opercular spine weak, poorly ossified, ventral to 2-6 membranous spinelets; spine and spinelets separated by moderate gap; spinelets occasionally obscured by under- lying membranes. Preopercular angle rec- tangular or slightly produced; preopercle, subopercle and interopercle unserrated. Gill rakers simple, awl-like. First dorsal fin VII (10); second dorsal fin 1,10 (10); anal fin 11,8 (1), 11,9 (9). Fin spines moderate; DJ 2.7-4.0% SL; D,I 10.9-12.1% SL; All 10.3-12.2% SL; PJ 11.0-13.6% SL. Vertebrae 10 + 15 ( 10 ) , epipleural ribs 7 (4), 8 (1), inserting on vertebrae 1-7 or 1-8 respectively; pleural ribs 7 (10), inserting on vertebrae 3-9. Color in alcohol variable with preserva- tion; specimens frequently abraded reveal- ing underlying pale yellow or pink-purple tissue. Recently collected specimens bear scale pockets mottled with numerous melanophores; dorsal surfaces of head and trunk more heavily pigmented; iris black. Specimens from old collections devoid of melanin, bearing silver on opercular region, isthmus, thoracic region, and abdomen to anus; iris silver. Mouth light, dotted with brown or black melanophores; l)ranchial region light in small specimens, darkening with age. 180 Bulletin Museum of Comparative Zoology, Vol 146, No. 3 r- o CM z CO 3 CO E E 00 o c> ui Q. > \- O _i o Q. O to 3 C O S. Uj in 0) Epigonus Systematics • Mayer 181 Table 15. Epigonus oligolepis meristic data. X = mean; SD = stan- dard DEVIATION; n = NUMBER OF SPECIMENS. X Range SD Pectoral fin rays 17.20 16-18 0.79 10 Gill rakers 30.50 29-31 0.71 10 Lateral line scales 34.70 33-36 1.06 10 Pyloric caeca 8.83 8-10 0.75 6 Table 16. Epigonus oligolepis regression data, b = regression coef- ficient ± 95% confidence interval; a = Y intercept; n = number of specimens. All regressions on SL. b a n HL 0. 36 + 0. 1 1 0. 88 7 Body depth 0. 26 + 0. 02 -2. 45 10 Head height 0. 21 + 0. 06 -2. 56 5 Eye diameter 0. 15 + 0. 03 0. 48 9 Snout length 0. 08 + 0. 03 0. 97 5 Interorbital width 0. 10 + 0. 01 -0. 51 9 Maxillary length 0. 18 + 0, 02 -0. 92 6 Lower jaw length 0. 17 + 0. 02 1. 45 10 Caudal peduncle d epth 0. 11 + 0. 02 -1. 20 10 Caudal peduncle L e n g t li 0. 26 + 0. 04 0. 58 9 D2I 0. 13 + 0, 01 -1. 07 6 All 0. 12 + 0. 02 -0. 33 10 P2I 0. 12 + 0. 03 0. 13 9 Description based on 10 specimens 53.7- 126.7 mm SL. Ontogenetic change. Two juvenile E. oligolepis (32.0-32.2 mm SL, USNM 207722) were taken by bottom trawls from the Gulf of Mexico. These specimens exhibit many traits characteristic of adult forms but differ in head shape, meristics. and dentition. Unlike adults, young E. oligolepis have smaller eyes (38.2-39.4% HL) and wider interorbital regions (10.4% SL). Dorsal fin and gill raker counts are reduced to VI-I,10 and 26 respectively. Premaxillary, mandibular, and lingual tooth patterns are similar to those of mature individuals, but dentition associated with 182 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Figure 16. Distribution of E. oligolepis. ■ individual haul of demersal adults; □ individual haul of demersal juveniles. the roof of the mouth is strongly reduced. Vomers and palatines are edentulous or bear 1-4 teeth; auxiliary tooth patches have not developed. Endopterygoid teeth are present but few in number, relatively long, and medially recurved. Distribution. E. oligolepis is endemic to the Gulf of Mexico-Caribbean region (Fig. 16). Specimens have been taken by bot- tom trawls between 380 and 660 meters. Remarks. The type specimens of E. oligolepis exhibit two seemingly disparate color patterns. One lot, taken in 1886 by the ALBATROSS, is devoid of melanin but bears extensive guanine deposits. Remain- ing fish, all more recently collected, bear no silver but are dotted with numerous melanophores. These differences are arti- facts of preservation. Specimens collected by early workers were generally placed directly into ethanol, while material obtained today is fixed in 10 percent formalin (Hubbs and Lagler, 1958: 16-17). When ethanol is used as a fixative, it leaches out melanins but does not affect guanine deposits. Specimens become pale, but silver pigment is retained. Formalin has the opposite effect; it blackens melanophores but destroys gua- nine crystals. The appearance of preserved specimens is thus dependent on fixative composition, concentration, and immersion time. An alcohol-formalin mixture con- taining one tablespoon of full strength formalin per two gallons of 6.5-75 percent ethanol might be used instead of conven- tional fixatives to preserve both guanine and melanin deposits (Myers, personal communication ) . Etymology. Oligolepis (Greek), few scales, from oligos, few, and lepis, scale; a noun in apposition, refers to the reduced number of lateral line scales characterizing the species. Common names. None. Epigonvs Systematics • Mayer 183 Figure 17. Epigonus trewavasae, 98.6 mm SL, USNM 207723. Epigonus trewavasae Poll, 1954 Figure 17 Glossamia pandionis, Lozano (not Goode and Bean, 1881), 1934: 89; Navarro, 1942: 202; Navarro et al., 1943: 136, plate XXII, fig. A. Epigonus trewavasae Poll, 1954: 91, fig. 27 (original description; NOORDENDE III Sta. 52, 06°08'S, 11°30'E, 280-290 m; holotype examined, IRSN 209). Epigojitis pandionis, Maurin (not Goode and Bean, 1881), 1968: 69, fig. 36. Diagnosis. E. trewavasae is most likely to be confused with E. robustus, E. leni- men, E. crassicaudus, and E. pectinifer. It is distinguished from the first three species by vertebral counts of 10 + 15 and the presence of glossohyal and lateral lingual teeth. The fourth form, E. pectinifer, bears only glossohyal teeth or a totally eden- tulous tongue. E. trewavasae may be further differentiated from E. pectinifer on the basis of the former's 30-35 awl-like gill rakers and long, pungent Dol and All (12.7-16.5% SL, 13.8-16.8% SL respec- tively). E. trewavasae is unlike remaining congeners because it bears a pungent, bony opercular spine, second dorsal fin counts of 1,9, and pectoral fin counts of 16-18. Description. Meristic values presented in Table 17; regression data for morpho- mctric traits presented in Table 18. Body elongate; anterodorsal profile flat, rising without interruption from snout to base of first dorsal fin; body moderate to deep, 23.1-27.0% SL; caudal peduncle length 24.3-27.5% SL. Head length 33.7-38.1% SL; head height 16.6-18.7%^' SL; snout pointed; angle of gape small to moderate; lower jaw pro- truding beyond upper jaw, bearing two nubs on anterior surface of mandible. Maxilla reaching slightly less than % eye length; posterior margin of maxilla narrow, rounded, or bearing posteriormost point near midline of bone; short, pungent mus- tache-like process projecting from postero- ventral surface of maxillary head. Eye round, slightly oval in younger specimens, 41.1-49.1%f HL; anterodorsal rim of orbit reaching profile; interorbital width 8.8- 10.8% SL. Dentition variable with age (see Onto- genetic change); teeth conical, small, fre- quently microscopic, present on premaxiL lae, mandibles, and vomer; palatines occasionally edentulous; tongue bearing lateral and glossohyal tooth patches. Opercular spine pungent, bony, sur- mounted by 2-3 horny spinelets; spine and spinelets separated by large gap; spinelets often obscured by underlying opercular membranes. Preopercular angle narrowly produced, unserrated or bearing serrations on angle and ventral surface of bone; in- teropercle and subopercle unserrated or weakly serrated. Gill rakers simple, awl- like. First dorsal fin VII (14); .second dorsal fin 1,9 (13), 1,10 (1); anal fin 11,9 (14); DJ moderate, 2.4-3.2% SL; DJ, All, PJ, 184 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 17. Epigonus trewavasae meristic data. X = mean; SD STANDARD DEVIATION; n =: NUMBER OF SPECIMENS. X Range SD Pectoral fin rays Gill rakers Lateral line scales Pyloric caeca n 17. 54 16- 18 0. 66 13 33. 15 30- 35 1. 46 13 47. 69 47- 49 0. 75 13 7. 00 6- 8 0. 60 12 Table 18. Epigonus trewavasae regression data, b = regression co- efficient ±95% confidence interval; a ^ Y intercept; n =: number of specimens. All regressions on SL. b a n HL 0. 38 + 0. 03 -2. 03 13 Body depth 0. 29 + 0. 02 -4. 43 12 Head height 0. 19 + 0. 01 -1, 26 12 Eye diameter 0. 17 + 0. 02 -0, 49 13 Snout length 0. 07 + 0. 02 1. 19 13 Interorbital width 0. 09 + 0, 01 1. 41 13 Maxillary length 0. 15 + 0. 02 0. 69 13 Lower jaw length 0. 16 + 0. 01 0. 39 13 Caudal peduncle de pth 0. 13 + 0. 01 -1. 79 1? Caudal peduncle len igth 0, 26 + 0. 02 -0. 08 13 D2 I 0, 15 + 0. 03 -0. 32 12 All 0. 18 + 0. 03 -0. 65 11 P2I 0, 14 + 0. 01 0. 36 13 long, pungent, 12.7-16.5%, 13.(8-16.8%, 13.8-16.27^ SL respectively. Vertebrae 10 + 15 (12); epipleural ribs 6 (9), 7 (2), inserting on vertebrae 1-6 or 1-7 respectively; pleural ribs 7 (8), 8 (4), inserting on vertebrae 3-9 or 3-10 respectively. Color variable with presei'vation; speci- mens abraded, revealing underlying yel- low to yellow-pink tissue; fin membranes dark; scale pockets covered with dense brown or black melanophores; dorsal sur- face of trunk more heavily pigmented than ventral; opercles brown, black, or slate gray; guanine deposits occurring occasion- ally on opercular region and from isthmus to bases of paired fins; iris black with sil- ver highlights; mouth light; branchial re- Epigonus Systematics • Mayer 185 JO' SO" Figure 18. Distributions of E. trewavasae and E. pectinifer. Large map shows localities in the Atlantic; insert shows localities off Japan. E. trewavasae: ^ individual haul of adults; Q individual haul of juveniles; cross- hatching indicates areas of capture cited in the literature. £. pectinifer: ■ individual haul of adults; □ individ- ual haul of juveniles; A report from the literature. gion light in smiill specimens, becoming l)lack with age. Description based on 13 specimens 70.9- 153.9 mm SL. Ontogenetic change. The most striking ontogenetic changes in E. trewavasae are associated with the development of adult tooth patterns. Large specimens bear ir- regular double or triple rows of premaxil- lary and mandibular teeth that taper to a single row posteriorly. Vomers are covered with minute conical teeth, while palatines are either edentulous or bear single to double rows of teeth. Dentition patterns are simple in small specimens but become more complex as teeth are added during growth. A 29.8- mm juvenile lacks both premaxillary and mandibular teeth. By 70-75 mm SL teeth are present in single rows on the jaws, and by 145 mm SL adult tooth patterns pre- vail. As premaxillary tooth patches widen, they extend posteriorly and eventually cover the first half of the bone. Analogous expansion occurs in vomerine tooth patches. Distribution. E. tretcavasae is known from equatorial west Africa, northwest Africa, and the western Mediterranean (Fig. 18). It has been taken by bottom trawls between 200 and 600 meters. Geographic variation. Statistical com- parisons of African and Mediterranean E. trewavasae were not made because of small sample size. As additional material is collected, the following intraspecific differences should be examined: ( 1 ) vomerine and palatine teeth more strongly developed in Mediterra- nean forms; (2) chin nubs more strongly developed in African forms; (3) preopercular serrations more strongly developed in Mediterra- nean forms. Although the significance of thc\se features 186 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 is unknown, they suggest that African and Mediterranean forms may represent sepa- rate subspecies. Taxonomic notes. Pomatomichthys con- stonciae GigHoh, 1880 may be a synonym of E. trewavasae Poll, 1954. See E. tele- scopus: Taxonomic notes, for a discussion of this possibility. Remarks. Dieuzeide (1950: 104-105) reported that specimens designated as Glossamia pandionis hy hozano (1934) and NavaiTO et al. (1943) were actually mis- identified E. denticulatus. This is incor- rect. Lozano's report is based on a single specimen (131 mm total length) taken from the Catillian coast. Among the char- acters cited for this fish are dorsal fin counts of VII-1,9, pectoral counts of 16, and an All subequal to the eye diameter (p. 89). All of these are characters diag- nostic of E. trewavasae. E. denticulatus bears 10 rays in the second dorsal fin, 18- 20 pectoral rays, and an All equalling half the eye diameter. Navarro et al.'s specimens also appear to be E. trewavasae. Altliough no descrip- tion is provided, the account includes a photograph (plate XXII, fig. A) that shows the fish have deep bodies, pungent oper- cular spines, and long D2I, All, and P2rs. All of these features are characteristic of E. trewavasae. More recently, Maurin (1968) mistook E. trewavasae for E. pandionis. Propor- tional measurements of body depth, head height. All, and P-I made on Maurin's figure 36 (p. 69) fall within ranges char- acteristic of E. treioavasae; however, pub- lished gill raker counts of 28-30 (p. 70) are lower than expected. Common names. None. Epigonus pectin! fer sp. nov. Figure 19 Ilolotype: A 114.3-mm SL specimen taken from the Caribbean west of Grenada by M/V OREGON, Sta. 5043: 26 September 1964, 12°01'N, 61°53.5'W, 210-250 fms., 40' shrimp trawl. USNM 207725. Paratypes: One specimen, 97.4 mm SL, 16 September 1964, Suruga Bay, commercial trawl. ABE 64-2085. One specimen, 100.6 mm SL, 14-31 October 1964, Suruga Bay, commercial trawl. ABE 64-2245. One specimen, 99.8 mm SL, 14-31 October 1964, Suruga Bay, commercial trawl. ABE 64-2248. Two specimens, 95.2-117.1 mm SL, station data identical with those of holotype. MCZ 48850. One specimen (cleared and stained), 108.1 mm SL, station data identical with those of holotype. MCZ 48851. One specimen, 94.8 mm SL, R/V PILLS- BURY, Sta. P-582: 23 May 1967; 21°10'N, 86°18'W; 250-155 fms.; 10' otter trawl. UMML 30378. One specimen, 111.2 mm SL, M/V OREGON, Sta. 4405: 27 September 1963; 11°53'N, 69°28"W; 215 fms.; 40' flat trawl. USNM 207726. Ten specimens, 101.8-120.6 mm SL, station data identical with those of holotype. USNM 207727. Nine specimens 81.5-118.9 mm SL, station data identical with those of holotype. USNM 207728. Two specimens (cleared and stained), 94.8- 98 mm SL, station data identical with those of holotype. USNM 207729. Epigonus rohiistiis, Matsubara (not Barnard, 1927), 1936: 121, fig. IB; Kamohara, 1952: 37. Diagnosis. E. pectinifer is characterized by comb-like gill rakers on the lower half of the first gill arch. This feature, together with glossohyal dentition (present in most specimens) and vertebral counts of 10 + 15, differentiate E. pectinifer from E. ro- htistus, E. lenimen, and E. crassicaudus. E. pectinifer most closely resembles E. tre- ioavasae but is distinguished by less exten- sive lingual dentition, fewer gill rakers (26-30), and shorter DJ and All (11.2- 12.7% SL and 11.9-14.0% SL respectively). E. pectinifer may be separated from re- maining congeners by its pungent, bony opercular spine, second dorsal fin counts of 1,9, and pectoral fin counts of 15-18. Description. Meristic values presented in Table 19; regression data for morpho- metric traits presented in Table 20. Epigonus Systematics • Mayer 187 CM o CM 3 E E UJ Q. >- !^ < Q. O V> Q. in C O o> 188 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 19. Epigonus pectinifer meristic data. X = mean; SD = stan- dard DEVIATION; n = NUMBER OF SPECIMENS. Range SD Pectoral fin rays 16.03 15-18 0.57 29 Gill rakers 27.59 26-30 0.98 29 Lateral line scales 48.14 47-49 0.58 29 Pyloric caeca 6, 10 5- 7 0. 41 29 Table 20. Epigonus pectinifer regression data, b = regression coef- ficient ± 95% confidence interval; a = Y intercept; n = number of specimens. All regressions on SL. b a n HL 0. 35 + 0. 05 -2. 21 27 Body depth 0. 28 + 0. 03 -4. 95 28 Head height 0. 18 + 0. 03 -2. 83 19 Eye diameter 0, 17 + 0. 03 -3. 18 28 Snout length 0. 09 + 0. 03 0. 03 26 Interorbital width 0. 10 + 0. 03 -1, 84 28 Maxillary length 0. 17 + 0. 04 -2. 10 26 Lower jaw length 0. 15 + 0. 03 0. 55 28 Caudal peduncle de pth 0. 84 + 0, 03 1. 88 28 C audal peduncle length 0. 27 + 0. 04 -0. 22 28 D2I 0. 11 + 0. 02 0. 62 24 All 0. 12 + 0. 02 0. 32 24 P2I 0. 11 + 0. 02 1. 30 28 Body elongate; anterodor.sal profile flat or slighdy convex, rising withont interrup- tion from snout to base of first dorsal fin; body depth 21.1-24.6% SL; caudal pedun- cle narrow, length 2.5.1-28.7% SL. Head short to moderate, 31.3-35.7% SL, shallow, 14.2-16.9% SL; snout wide, pointed; angle of gape small; lower jaw proti-uding slightly beyond upper jaw; nubs at anterior end of mandible paired, barely discernible, or absent. Maxilla reacliing %-y2 eye length, posterior margin narrow, rounded, or bearing posteriormost point near midline of bone; short, pun- gent, mustache-like process projecting from posteroventral surface of maxillary head. Eye round or slightly oval, 38.7- 45.4% HL; anteiodorsal rim of orbit not Epigonus Systematics • Maijer 189 reaching profile; intcrorl)ital widtli 7.7- 9.4% SL. Teeth small, conical; premaxilla edentii- lons or bearing teeth anteriorly; when present, teeth 1-15, arranged in single row. Mandibular teeth covering all or part oi dentary, arranged in single row. Vomer- ine teeth strong, arranged in tightly packed ()\'al patch. Palatines edentulous or bear- ing teeth anteriorly; when present, teeth 1-6, arranged in single row; tongue with glossohyal teeth, rarely edentulous. Opercular spine pungent, bony, ventral to 2-3 horny spinelets; spine and spinelets separated by large gap; spinelets occasion- ally obscured by underlying membranes. Preopercular angle narrowly produced, serrated; subopercle and interopercle un- serrated or weakly serrated. Gill rakers pectinate, bearing nub-like projections proximally along mesial surfaces (Fig. IC); pectinate structure variable in extent, most prominent on ventral portions of gill arch. First dorsal fin VII (28); second dorsal fin 1,9 (29); anal fin 11,9 (29); D,I short, 1.6-2.8% SL; D,I moderate, 11.2- 12.7% SL; All, PJ, 11.9-14.0% SL. Vertebrae 10 + 15 (29); epipleural ribs 6 (17), 7 (13), inserting on vertebrae 1-6 or 1-7 respectively; pleural ribs 8 (29), inserting on vertebrae 3-10. Color in alcohol brown-black; fin mem- branes black; scale pockets covered with densely packed melanophores; skin often abraded, revealing underlying yellow-pink tissue; iris black; branchial region white to dark gray; mouth light. Description based on 30 specimens 81.5- 120.6 mm SL. OntO(!,enetic change. A 33.8-mm E. pec- tinifer was taken by bottom trawl in the Gulf of Mexico (USNM 207731). The specimen appears similar to adults and pro- vides little evidence of ontogenetic change. The major difference is the presence of six rather than seven first dorsal fin spines. Distribution. E. pectinifer is known from the Caribbean Sea, Gulf of Mexico, and eastern coast of Japan (Fig. 18). Specimens were taken between 280 and 550 meters. GeograpJiic variation. Definitive com- parisons of Japanese and American E. pec- tinifer were not undertaken, because only three oriental specimens were available for study. The latter forms were, however, in- dividually compared with Amcnican fish. The analyses revealed virtually no differ- ences between the populations aside from a slight tendency toward broader caudal peduncles and shorter maxillae and man- dibles by the Japanese specimens. Remarks. A teratological specimen of E. pectinifer was taken from the Yucatan Channel (109 mm SL, UMML 30379). The fish was captured at depths characteristic of E. pectinifer and bears diagnostic traits such as 27 gill rakers ( many are pectinate ) , VII + I dorsal fin spines, 16 pectoral fin rays, and 10 + 15 vertebrae. The tongue is edentulous. Unlike the condition in typical forms, opercles are not fully ossified and lack spines and spinelets. Similarly, the lateral line is incomplete on the right side and bears only 43 pored scales on the left. Other differences include enlarged teeth and chin nubs, 10 rather than 9 dor- sal rays, and 8 rather than 5-7 pyloric caeca. The aberrant specimen was not consid- ered in preparing the species description. Etymology. Pectinifer (Latin), comb- bearer, from pecten, comb, and ferare, to bear; a noun in apposition, refers to the comb-like gill rakers characterizing this species. Common names. None. Epigonus robustus (Barnard, 1927) Figure 20 Epigonus macrops Gilchrist and von Bonde, 1924: 14, plate I, fig. 3 (oiij^inal description; S..S. PICKLE Sta. 344, 30°12'00"S, 14°25'()()"E, 510 fms.; Sta. 347, 31°58'00"S, 16°00'00"E, 670 fms.; syntype examined, RUSI 669; name suppressed, junior homonym of Oxi/odot} macrops Brauer, 1906); Barnard, 1927:' 523; Smith, 1961: 377, fig. 2. 190 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Figure 20. Epigonus robustus, 154.6 mm SL, LACM 11449-7. Parahynnodiis robustus Barnard, 1927: 525, plate XXII, fig. 4 (original description; off Cape Point, 460 fms.; holotype in poor condition, not examined, SAM 13080). Hymwdus robustus Smith, 1949b: 210, fig. 495. Diagnosis. E. robustus sti-ongly resem- bles E. pectinifer, E. trewavasae, and E. lenimen. It may be distinguished from the former two species by the absence of hngual teeth. In addition, mihke E. pec- tinifer, it has awl-Hke gill rakers. E. ro- hu^us differs from E. lenimen by having a narrow interorbital region (6.5-8.2% SL), short DJ (10.0-12.6% SL) and short All (9.2-13.3% SL). E. robustus may be distinguished from E. crassicaudus by the former's short head (28.0-34.0% SL) and shallow body (20.3-24.6% SL). It differs from remaining congeners by bearing a pungent, bony opercular spine, vertebral count of 11 + 14, and nine rays in the second dorsal fin. Description. Meristic values presented in Table 21; regression data for morpho- metric traits presented in Table 22. Body elongate, moderately compressed; anterodorsal profile weakly convex, rising without interruption from tip of snout to base of first dorsal fin; body depth 20.3- 24.6% SL; caudal peduncle moderate to long, 25.3-30.7% SL. Head short, shallow, length 28.0-34.0% SL, height 14.8-16.3% SL; snout short, pointed; angle of gape moderate to large; lower jaw protruding beyond upper jaw, bearing two nubs of variable prominence on anterior surface of mandible. Maxilla reaching ¥3-^/4 eye length; posterior margin of maxilla narrow, rounded or bearing posteriormost point near midline of bone; small, weak mustache-like process project- ing from j)osteroventral surface of maxil- lary head. Eye round to oval, small, 37.4- 42.4% HL; anterodorsal rim of orbit not reaching dorsal profile; interorbital region narrow, 6.5-8.2% SL. Teeth small, conical; premaxilla edentu- lous or bearing single row of teeth on an- terior half of bone; mandibular dentition covering all or part of dentary, arranged in double row anteriorly, tapering to single row posteriorly; vomer bearing 1-6 irregu- lar rows of teeth; palatines edentulous or bearing teeth on anterior half of bone; tongue edentulous. Opercular spine pungent, bony, ventral to 2-3 membranous or horny spinelets; spine separated from spinelets by wide gap; spinelets often obscured by underly- ing membranes. Preopercular angle not produced, serrations on posterior and/ or ventral surfaces of bone rarely absent; sub- opercle and interopercle serrated. Gill rakers simple, awl-like. First dorsal fin VI (1), VII (27), VIII (1); second dorsiil fin I, 9 (28), II, 8 (1); anal fin II, 9 (29). DJ short, 1.4-2.5% SL; DJ, All, PJ moderate to long, 10.0- 12.6%, 9.2-13.3%, 11.7-15.3% SL respec- tively. Vertebrae 11 + 14 (29); epipleural ribs Epigonus Systematics • Mayer 191 Table 21. Epigonus robustus meristic data. X = mean; SD = stan- dard DEVIATION; n = NUMBER OF SPECrMENS. X Range SD n Pectoral fin rays 16.79 16-18 0.55 29 Gill rakers 31.68 30-33 0.93 29 Lateral line scales 48.76 47-50 0.91 29 Pyloric caeca 6.36 5- 8 0.78 28 Table 22. Epigonus robustus regression data, b = regression coef- ficient ± 95% confidence interval; a = Y intercept; n = number of specimens. All regressions on SL. b a n HL 0. 28 + 0. 02 5, 35 28 Body depth 0. 28 + 0. 02 -7. 80 28 Head height 0, 17 + 0. 02 - 1. 65 20 Eye diameter 0. 11 + 0. 01 44 28 Snout length 0. 06 + 0. 02 80 23 Interorbital widtl: 0. 09 + 0. 01 - 1. 97 28 Maxillary length 0. 12 + 0. 01 14 24 Lower jaw length 0, 14 + 0. 01 27 27 Caudal peduncle dep ith 0. 13 + 0. 01 - 1. 69 28 Caudal peduncle len gth 0. 25 + 0. 02 5. 32 28 D2I 0. 08 + o„ 02 5. 49 2 1 All 0. 07 + 0, 02 8„ 18 15 P2I 0. 09 + 0. 02 6. 74 24 6 (2), 7 (8), inserting on vertebrae 1-6 brown or black niclanopliores; brancliial or 1-7 respectively; pleural ribs 9 (29), in- region black. Body very oily; body cavity serting on vertebrae 3-11. filled with rust brown fat globules; viscera Color variable with preservation, pale and swinibladder often completely envel- yellow to rust brown; scale pockets out- oped in fat. lined by small black or brown melano- Description based on 29 .specimens phores; opercular region tinged with 121.1-198.0 mm SL. black; iris black; mouth light, mottled with Distribution. Most specimens of E. ro- 192 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 EriGONus Systematics • Mayer 193 Figure 22. Epigonus lenimen, 139.0 mm SL, UZM P45165. biistus liave been taken by bottom trawLs between 800 and 1225 meters off south- eastern South Ameriea, South Africa, and AustraHa (sec Fig. 21). One specimen (ISH 430/71) was taken by a deep pelagic trawl. Geographic variation. No investigation was undertaken because insufficient ma- terial was available from South Africa and Australia. Taxonomic notes. Epigonus macrops Gilchrist and von Bonde, 1924 was des- cribed from two syntypes; the larger was 19(S mm (SL?). These specimens, together with manv others collected bv the Fish- cries and Marine Biological Survey, were lost while being transferred to the South African Museum. A portion of the ma- terial was subsequently rediscovered at Rhodes University, Grahamstown. From the contents Smith (1961: 378) described a specimen that he believed to be "Gil- christ and \'on Bonde's type of macrops from 600 fathoms off St. Helena Bay." This fish was re-examined during the present study. Smith's specimen measures 162.2 mm SL and thus cannot be the larger syntype; however, it conforms to the descriptions and proportions supplied by Gilchrist and von Bonde and probably represents the smaller type for which no length was pub- lished. An unusual aspect of the syntypes is that the locations at which they were captured will never be precisely known. The speci- mens were taken at different stations. Al- though these are recorded in both the orig- inal description of E. macrops and in the 1921 report of the Fisheries and Marine Biological Survey (Gilchrist, 1922), neither account specifies which data are associated with which syntype. Common names. None. Epigonus lenimen (Whitley, 1935) Figure 22 Scepterias lenimen WHiitley (in part), 1935: 230 (original description; Great Australian Biglit: south from Eucla, 350—450 fnis.; holot\-pe examined, AM E3368); Whitley, 1940: 420, fig. 33; Wliitley (in part), 1968: 56. Epigonus lenimen Scott, 1962: 191, 1 fig. Diagnosis. E. lenitnen is distinguished from E. robustus and E. crassicaudus by its broad interorbital region (8.7-10.2% SL), long DJ (14.9-18.7% SL), and large eyes (40.0-51.1% HL). It is further dif- ferentiated from E. crassicaudus by shorter head lengths (32.7-36.67^ SL) and .shal- lower head heights (16.2-18.8% SL). E. lenimen lacks lingual teeth but has 11 + 14 vertebrae and thus may be distinguished from E. trewavasae and E. pectinifer. Un- like remaining congeners, E. lenimen bears a pungent, bony opercular spine, nine second dorsal fin rays, and 16-18 pectoral fin rays. 194 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 23. Epigonus lenimen meristic data. X = mean; SD = standard deviation; n = nxtmber of specimens. X Range SD Pectoral fin rays 16.96 16-18 0.58 28 Gill rakers 30.29 28-34 1.27 24 Lateral line scales 48.12 47-50 0.91 26 Pyloric caeca 7.33 7- 9 0.56 24 Table 24. Epigonus lenimen regression data, b = regression coeffi- cient ± 95% confidence interval; a = Y intercept; n = nltmber of specimens. All regressions on SL. b a n HL 0. 35 + 0, 01 -0. 12 29 Body depth 0. 28 + 0. 02 -3. 32 27 Head height 0, 19 + 0. 01 -1. 08 27 Eye diameter 0. 18 + 0. 01 -1, 60 27 Snout length 0. 08 + 0. 01 0. 31 27 Interorbital width 0. 10 + 0, 01 -0. 59 26 Maxillary length 0. 16 + 0. 01 0. 08 27 Lower jaw length 0. 16 + 0. 01 0. 55 28 Caudal peduncle dep th 0. 11 + 0. 01 -0. 14 28 Caudal peduncle len gth 0. 24 + 0. 02 2. 26 26 D2I 0. 17 + 0. 02 -0. 79 18 All 0. 21 + 0. 02 -2. 36 22 P2I 0. 19 + 0. 01 -2, 13 28 Description. MerLstic values presented in Table 23; regression data for morphometric traits presented in Table 24. Body elongate; anterodorsal profile flat or weakly concave, rising without inter- ruption to first dorsal fin, more steeply inclined behind occiput in large specimens; body moderate to deep, 21.5-27.5% SL; caudal peduncle moderate to long, 23.6- 29.3% SL. Head length 32.7-36.6% SL; head height 16.2-18.8% SL; snout moder- ately pointed; angle of gape moderate, variable with age; lower jaw protruding slightly or not at all; no prominent nubs on anterior surface of mandible. Maxilla reaching Vs-V2 eye length; posterior margin Epigonus Systematics • Mayer 195 Table 25. Comparison of E. LENiMEy paratypes with specimens of E. lknimen and E. DENTICULATVS. PaRATYPE MERISTICS REPORTED AS VALUE, FOLLOWED IN PARENTHESES HY NUM- BER OF SPECLMENS EXHIBITING THAT VALUE. RATIOS ARE EXPRESSED AS PERCENTAGES. E . 1 e n i m e n E_. 1 e n i m e n paratypes E^. denticulatus Dorsal fin rays 8-9 9(1), 10(11) 10 Pectoral fin rays 16—18 19(6), 20(6) 18—20 Vertebrae 11+14 10+15 10+15 10(1), 11(3) , „ , , Pyloric caeca 7 — 9 12(7) — BH/SL 21.5—27.5 18.4 — 21.7 15.8—23.6 D2 I/SL 14.9—18.7 6.0—7.6 5.3—8.0 AII/SL 13.0 — 20.8 6.2—7.1 6.0—8.2 P2 I/SL 12.5 — 18.7 8.5—9.9 7.9—10.0 of maxilla narrow, rounded, or bearing First dorsal fin VII (29); second dorsal posteriormost point near midline of bone; fin 1,8 (1), 1,9 (28); anal fin 11,8 (2), 11,9 weak mustache-like process projecting from (26); DJ moderate, 2.0-4. 17^ SL; DJ, posteroventral surface of maxillary head, All long, 14.9-18.7%, 13.0-20.8% SL re- process occasionally absent. Eye large, spectively; PJ moderate to long, 12.3- oval, 40.0-51.1% HL; anterodorsal rim of 18.7% SL. orbit reaching dorsal profile; interorbital Vertebrae 11 + 14 (29); epipleural ribs width 8.7-10.2% SL. 6 (6), 7 (12), 8 (2), inserting on vertebrae Teeth small, conical; premaxilla eden- 1-6, 1-7, or 1-8 respectively; pleural ribs tulous or bearing single row of teeth oc- 9 (28), inserting on vertebrae 3-11. cupying anterior half of bone. Mandible Color in alcohol variable; skin often edentulous or bearing single row of teeth abraded, revealing underlying pale pink- occupying up to % of dentary; tooth row yellow tissue; fin membranes and scale occasionally double near symphysis. Vo- pockets mottled with numerous black mer edentulous or bearing up to seventeen melanophores; head, opercular region, and teeth arranged in diamond-shaped patch fin bases deep rust brown. Guanine de- or in 1-3 irregular rows; palatines edentvi- posits variable, occurring on ventral por- lous or bearing 1-2 teeth anteriorly; tongue tions of opercular region, isthmus, pectoral edentulous. and pelvic fin bases, and abdomen to anus; Opercular spine pungent, bony, ventral silver chromatophores on dorsal, anal, pee- to 1-5 (usually 2) membranous or horny toral, or pelvic fin rays; iris black with spinelets; spine and spinelets separated by silver highlights; moutli light, dotted with wide gap; spinelets frequently obscured by melanophores; branchial region light in underlying membranes. Preopercular angle small specimcMis, blackening with age. narrowly produced, occasionally serrated; Description based on 32 specimens 40.0- subopercle and interopercle unserrated or 147.8 mm SL. weakly serrated. Gill rakers simple, awl- Distrihution. E. lenimen is known from like. three localities (Fig. 21). The liolotype 196 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 0 I 2 3 cm Figure 23. Epigonus crassicaudus, 259.0 mm SL, MCZ 48855. was taken south of Austi-alia between 622 and 823 meters. Remaining specimens were taken off New Zealand between 530 and 660 meters. Taxonomic notes. Although G. P. Whit- ley was the first to describe E. lenimen (1935), inaccuracies in his papers have produced several problems. The most serious involve the type series and type locality of the species. The original description of E. lenimen designates a holotype and nine paratypes. A figure of the new form was not included but was published in a subsequent paper (Whidcy, 1940: fig. 33). Both the de- scription and the illustration were based exclusively on the holotype. Whitley re- alized that the paratypes were different from the holotype but considered them to be poorly preserved specimens ( Whitley, 1935: 320). A re-examination of the type series re- veals that the paratypes are not conspecific with the holotype. They are, instead, mem- bers of E. denticulattis. As is shown in Table 25, counts and measurements from the paratypes always fall within ranges characteristics of E. denticulatus. Pectoral fin counts, vertebral number, pyloric caecum counts, and fin spine lengths are particularly noteworthy in this respect. The paratypes further resemble E. denti- culatus by bearing dentigerous palatines and weak opercular armor, E. lenimen, on the other hand, is characterized by eden- tulous palatines and pungent opercular spines. Confusion over the type locality stems from Whitley's 1940 paper. The locality is cited as "from 190-320 fathoms, S.W. from Eucla, Great Australian Bight [p. 420] ." This contradicts the data presented in the original description: "Great Aus- ti-alian Bight; south from Eucla, 350 to 450 fathoms [p. 231]." The 1940 citation is extremely similar to station data listed for paratypes AM E3581-3582 in 1935 ("Great Australian Bight; SW from Eucla, 190-320 fathoms. 126° 451/2'E long. [p. 231]"). In the absence of other information, it must be concluded that erroneous locality data were inserted in the 1940 publication through an editorial oversight. The most recent taxonomic questions arise from Whitley's check list of New Zealand fishes ( 1968 ) . This work includes two incorrect citations in the synonymy of E. lenimen. The first is based on a fish taken off the Chatham Islands and tenta- tively identified as Hynnodiis atherinoides (Moreland, 1957). This specimen was later re-identified as Grahamichthijs radiatus (Moreland, personal communication). The second misidentified .specimen is a "Big- eyed Cardinal Fish" captured off Cape Palliser, New Zealand (Anonymous, 1961). This fish is actually a specimen of E. telescopus and is presently in the collec- Ei'ic.oNus Systematics • Mayer 197 tions of the Dominion Museum (DM 3072, examined ) . Common names. None. Epigonus crassicaudus de Buen, 1959 Figure 23 Epiguinis crcis.'iicattdii.s de Buen, 1959: 196 (original description; preabysnial zone off Valparaiso, Chile; holotype not examined, EBM 10.183). Diagnosis. E. crassicaudus is strongly compressed. It reaches 260-270 mm SL and is the second largest species in the genus. E. crassicaudus may be distin- guished from E. trewavasae, E. pectinifer, E. rohustus, and E. lenimen by its deep head (18.9-21.2% SL) and deep body (24.3-32.0% SL). It differs from remain- ing congeners by bearing 9 rays in the second dorsal fin and 6-7 pyloric caeca. Description. Meristic values presented in Table 26; regression data for morpho- metric traits presented in Table 27. Body elongate, compressed; anterodorsal profile rising from tip of snout to occiput, becoming moderately convex from occiput to base of first dorsal fin. Body deep, 24.3- 32.07f SL; caudal peduncle broad, moderate to short, 21.6-26.4% SL. Head long, deep, postorbital portion greatly expanded, length 36.8-41.9% SL; height 18.9-21.27r SL; snout moderately pointed in small specimens, blunt in adults; angle of gape moderate to small; mandible long, strongly protuberant, young bearing two weak nubs on anterior surface of lower jaw. Maxilla reaching %-% eye length; posterior margin of maxilla broad, rounded or bearing posteriormost point between midline and ventral surface of bone. Eye round, small, 34.2-39.6% IlL; surrounded by numerous small scale pockets; antero- dorsal rim of orbit reaching dorsal profile^; interorbital region narrow, 6.2-8.5% SL. Teeth small, conical, occasionally villi- form, larger in small specimens; premax- illary teeth arranged in irregular single or double rows tapering to single row posteriorly and covering from % to entire length of bone; mandibular teeth arranged in multiple rows, tapering to single row posteriorly, covering from V2 to entire IcMigtli of dentary; vomer edentulous or bearing up to six irregular rows of minute teeth; palatines edentulous or bearing 1-3 teeth anteriorly; tongue edentulous. Opercular spine pungent, bony, ventral to 3-5 flat, horny spinelets; spine separated from spinelets by narrow gap; spinelets often obscured by underlying membranes. Preopercular angle slightly produced, pos- terior and/ or venti'al surfaces serrated; subopercles and interopercles serrated. Gill rakers awl-like, short; gill filaments long. First dorsal fin VII (22); second dorsal fin 1,9 (20), 1,10 (2); anal fin II.8 (1), 11,9 (21); DJ 2.0-3.6% SL; DJ 9.-8-13.2% SL; All 10.3-14.0% SL; PJ 13.0-15.5% SL. Vertebrae 11 + 14 (25); epipleural ribs 6 (2), 7 (16), inserting on vertebrae 1-6 or 1-7 respectively; pleural ribs 9 (25), inserting on vertebrae 3-11. Color in alcohol variable with preserva- tion; skin frequently abraded, exposing underlying pink tissue and orange-rust fat deposits; skin exti'emely oily; fin membranes black; scale pockets mottled with numerous black melanophores; dorsal portion of body darker than ventral; forehead, snout, an- terior half of mandible, and circumorbital area heavily invested with black pigment; opercles black or slate gray. Guanine de- posits occasionally on opercles, isthmus, pectoral and pelvic fin bases, and al)do- men to anal fin; iris variable — black, siher, or black with silver highlights; mouth and branchial region light, darkening with age. Description based on 27 specimens 80.3- 262.5 mm SL. Ontogenetic change. Two juvenile E. crassicaudus (12.2 mm SL, MCZ 48857, and 21.7 mm SL, MCZ 48858) were taken off the Chilean coast by midwater trawl. Although these forms bear characteristics diagnostic of the species, they differ con- 198 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Table 26. Epigonus crassicaudus meristic data. X = mean; SD STANfDARD DEVIATION; n = NUMBER OF SPECIMENS. X Range SD Pectoral fin rays 18.05 17-19 0.58 22 Gill rakers 32.27 31-34 0.70 22 Lateral line scales 47.86 46-49 0.85 21 Pyloric caeca 6.87 6- 7 0.35 15 Table 27. Epigonus crassicaudus regression data, b = regression COEFFICIENT ± 95% CONFIDENCE INTERVAL; a = Y INTERCEPT; n = NUMBER OF SPECIMENS. AlL REGRESSIONS ON SL. b a n HL 0. 39 + 0. 02 0. 47 20 Body depth 0. 30 + 0. 04 -3. 75 21 Head height 0. 21 + 0. 02 -1. 62 18 Eye diameter 0. 14 + 0. 01 0. 74 21 Snout length 0. 08 + 0. 01 0. 08 19 Interorbital width NONLINEAR Maxillary length 0, 17 + 0. 01 -0. 49 20 Lower jaw length 0. 20 + 0, 01 -1, 17 22 Caudal peduncle dep th 0, 12 + 0. 01 - 0, 65 20 Caudal peduncle len gth 0. 22 + 0, 03 4, 32 20 D2 I 0. 10 + 0. 02 3. 34 12 All 0. 11 + 0. 02 2. 63 17 P2I 0. 14 + 0. 01 0. 05 16 sideral)ly in appearance and habit from adnlts. Most .striking is the juvenile pigment pattern. Pelagic specimens are basically pale yellow with large, brown patches covering most of the caudal peduncle. Caudal peduncle rings, like those found on E. pandionis young, are absent, although myotomes are outlined by thin brown bands. Brown pigment extends anteriorly as a band from the caudal peduncle to the frontal region of the head. A poorly de- fined black stripe extends across the snout to the anterior rim of the orbit. In general, juvenile E. crassicaudus resemble E. teles- copiis young figured by Koefoed (1952: plate IIA). The midwater capture of E. crassicaudus Epigonus Systematics • Mayer 199 juveniles suggests that tlie life cycle of tlK> specit^s includes a pelagic stage. Unfortu- nately, the data available are not sufficient to determine the duration of this stage. Distribution. E. crassicaudus is endemic to the waters off central Chile (Fig. 21). Adults have been captured by bottom trawls made between 200 and 400 meters; juveniles were taken by midwater trawls fishing from 200 to 270 meters. Common mimes. None. Species Incertae Sedis Micwichtlnjs coccoi Riippell, 1852: 1 (original description; "Mare siculum"; holotype not ex- amined, SMF 1069). The original description of M. coccoi provides only a superficial account of the holotype. Subsequent papers either para- phrase Riippell's work {e.g., Canestrini, 1860; Doderlein, 1889) or are based on material not compared to the holotype (i.e., Facciola, 1900; Caporaicco, 1926; Gonzales, 1946). It is questionable whether the latter specimens are conspecific with the holotype. Most recent revisers (e.g., Schultz, 1940; Norman, 1957) have synonymized Micro- ichthys with Apogon; however, the data are inconclusive and also suggest an affinity with Epigonus (Eraser, 1972: 5). A re- examination of the holotype must be under- taken to clarify the status of M. coccoi. A second species of Microichthys — M. sonzoi Sparta, 1950 — does not appear to be an Epigonus on the basis of vertebral and dorsal fin counts. The only known speci- men of this species has been lost (Torton- ese, personal communication). ACKNOWLEDGMENTS This work would not have been possible without the assistance and support of niunerous people. I wish to thank the following scientists and institutions for material used in this study: J. R. Paxton and D. Hoese, Australian Museum; A. W. Wheeler and G. Palmer, British Museum (Natural History); J. Randall, Bemice P. Bishop Museum; W. Eschmeyer, California Academy of Sciences; E. Bertclsen, Carls- bergfondets; J. Moreland, Dominion Mu- seum; L. P. Woods, Eield Museum of Natural History; P. Struhsaker, National Marine Fisheries Service, Honolulu; R. Raymond, Instituto de Fomento Pesquero; X. Missonne, Institut Royal des Sciences Naturelles de Belgique; G. Krefft, Institut fiir Seefischerei; M. M. Smith, J. L. B. Smith Institute of Ichthyology; I. Naka- mura, Kyoto University; R. J. Lavenberg, Los Angeles County Museum of Natural History; M. Bauchot, Museum National d'Histoire Naturelle; M. Poll, Musee Royal de I'Afrique Centrale; E. A. Lachner and T. H. Eraser, National Museum of Natural History; M.-L. Penrith, South African Mu- seum; George R. Vliller, Tropical Atlantic Biological Laboratory; R. S. Gaille, Texas Parks and Wildlife Department; M. Leible, Universidad Catolica de Chile; C. R. Robins, University of Miami; J. Nielsen, Universitetets Zoologiske Museum; B. Nafpaktitis, University of Southern Cali- fornia; T. Abe, University of Tokyo; R. Backus and J. Craddock, Woods Hole Oceanographic Institution; and C. Karrer, Zoologisches Museum, Berlin. W. Klause- witz of the Natur-Museimi Senckenberg provided invaluable information on the holotype of Microichthys coccoi, and E. Tortonese of the Museo Civico di Storia Naturale, Genoa, answered numerous ques- tions about problematical forms such as Pomatomichthys constanciae and Micro- icJithys sanzoi. I am greatly indebted to Ernst Mayr, Giles W. Mead and Karel F. Liem for their guidance, criticism, and support of my work. I am also grateful to Richard L. Haedrich for reading the manuscript and assisting in the planning of this research. Special thanks are extended to G. S. Myers for assistance with taxonomic problems. I wish to thank the staffs of the Fish Department, Museum of Comparative Zoology, and Department of Natinal Sci- 200 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 ences, Boston University, for their useful comments and practical help. Karen Green- leaf and Pat Allen typed the final draft of this manuscript. Illustrations of eleven of the twelve species of Epigonus were prepared by L. Laszlo Meszoly. Jordan and Jordan's il- lustration of E. fragilis (Fig. 10) was made available through the courtesy of the Carnegie Museum. Finally, a hearty vielen Dank to my wife for her patience, encouragement, and ed- itorial assistance. Support for this work was provided by NSF Graduate Fellowships during 1966 to 1971 and a grant from Harvard Uni- versity's Committee on Evolutionary Bi- ology (GB7346). LITERATURE CITED Anonymous. 1961. Caught only 3 times before. Wellington Evening Post. Bailey, N. T. 1959. Statistical Methods in Biology. New York: John Wiley and Sons, Inc. 200 pp. Barnard, K. H. 1927. A monograph of the marine fishes of South Africa, Part II. Ann. South African Mus., 12(2): 419-1065. Bauchot, M. L., and M. Blanc. 1961. Poissons marins de I'Est Atlantique Tropical, II. Percoidei (Teleosteens Perciformes) lere partie. Atlantide Rep. No. 6: 65-100. Bertolini, F. 1933. Apogonidae. Fauna e flora del Golfo di Napoli. Uova, larve e stadi giovanili di Teleostei. Pubbl. Staz. Zool. Napoli, 38: 306-309. BiNi, G. 1968. Atlante dei Pesci delle Coste Italiane, Vol. IV. Roma: Mondo Sommerso Editrice. 163 pp. Bleeker, p. 1876. Systema Percarum revisum. Pars I et II. Arch. Neerl. Sci., 11: 274-340. Braxjer, a. 1906. Die Tiefsee-Fische. I. Syste- matischer Teil. Wiss. Ergeb. Deut. Tiefsee- Exped. "Valdivia," 1898-1899, 15: 1-420. Breder, C. M., and D. E. Rosen. 1966. Modes of Reproduction in Fishes. Garden City, N. Y.: Natural History Press. 941 pp. Brown, M. E. 1957. Experimental studies on growtli. In The Physiology of Fishes, Vol. I, M. E. Brown (ed. ). New York: Academic Press, Inc., pp. 361-400. Buen, F. de. 1959. Notas preliminares sobre la fauna marina preabismal de Chile, con descripcion de una familia de rayas, dos generos y siete especies nuevos. Bol. Mus. Nac. Hist. Natur., 27(3): 171-201. Canestrini, J. 1860. Zur Systematik der Per- coiden. Verb. Zool. Bot. Ver. Wien, 10: 291-314. Capello, F. de B. 1868. Peixes novos de Portugal e da Africa occidental e caracteres distinctivos d'outras especies ja conhecidas. J. Sci. Math. Phys. Natur. Acad. Real Sci. Lisboa, 1(2): 154-169. Caporaicco, L. dl 1926. II cranio de "Micro- ichthvs coccoi" Riipp. Monit. Zool. Ital., 37(6): 127-132. Cligny, a. 1903. Poissons des Cotes d'Espagne et de Portugal ( Ocean Atlantique ) . Premiere partie. Boulogne. 30 pp. Cocco, A. 1829. Su di alcuni nuovi pesci de' mari di Messina. Giom. Sci. Lett. Art. Sicil., 7(77): 138-147. . 1885. Indice Ittiologico del Mare di Messina. Nat. Sicil., 4(4): 85-88. CuvrER, G. 1828. Des pomatomes. In Histoire Naturelle des Poissons, Tome Second, G. Cuvier and A. Valenciennes. Paris: F. G. Levrault, pp. 169-174. Dieuzeide, R. 1950. Sur un Epigonus nouveau de la Mediterranee (Epigonus denticulatus, nov. sp.). Bull. Sta. Aquic. Peche CastigUone, N.S., No. 2: 89-105. , M. Novella, and J. Roland. 1953. Catalogue des poissons des Cotes algeriennes. II. Osteopterygiens. Bull. Sta. Aquic. Peche Castiglione, N.S., No. 5: 1-258. Dlxon, W. J. (ed.). 1967. BMD Biomedical Computer Programs. Univ. Calif. Publ. Auto- matic Computation No. 2. Berkeley: Univ. Cahf. Press. 600 pp. DoDERLEiN, p. 1889. Manuale Ittiologico del Mediterraneo. Fascicolo IV. Palermo: Giornali di Sicilia. 188 pp. Dons, C. 1938. Notes on fishes III. Epigonus telescopiis (Risso), new to Norway. Det Kgl. Nor. Vidensk. Selsk. Forh., 11(35): 141-142. Eblna, K. 1931. Buccal incubation in the two sexes of a percoid fish, Apogon semiUneatus T. & S. T. Imp. Fish. Inst. Tokyo, 27(1): 19-21. Ehrenbaum, E. 1928. Rare fishes in the North Sea. Nature (London), 121(3053): 709. Ekman, S. 1953. Zoogeography of the Sea. London: Sidgwick and Jackson, Ltd. 417 pp. Facciola, L. 1900. Sul Microichthys coccoi Rapp. Monit. Zool. Ital., 11(5): 188-194. Fowler, H. W. 1928. The fishes of Oceania. Mem. Bernice P. Bishop Mus., 10: 1-540. . 1935. South African fishes received from Mr. H. W. Bell-Marley in 1935. Proc. Acad. Nat. Sci. Philadelphia, 87: 361-408. . 1936. The marine fishes of West Africa Ei'iGONus Systematics • Mayer 201 based on tlie collection of the American Musemn Congo Expedition, 1909-1915. Bull. Amer. Mus. Nat. Hist., 70(2): 607-1493. — , AND B. A. Bean. 1930. The fishes of the British Museum, Vol. I. London. .524 pp. 1868. Report on a c(jllection of fishes the families Amiidae, Chandidae, Duleidae, and Serranidae obtained by the United States Bureau of Fisheries Steamer "Albatross" in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Bull. U.S. Nat. Nhis., 10(100): 1-334. FuASER, T. H. 1972. Comparative osteology of the shallow water cardinal fishes IPerci- formes: Apogonidae] with reference to the s>'steniatics and exolution of the family. Ichthvol. Bull. J. L. B. Smith Inst. Ichthyol., Rhodes Univ., No. 34: 1-105. FucJLLSTER, F. C. 1960. Atlantic Ocean atlas of temperature and salinity profiles and data from the International Geophysical Year of 1957-1958. Woods Hole Oceanogr. Inst. Atlas Ser., 1 : 1-209. Gall, J- le. 1931. Epigomts telescopus Risso 1810. In Faune Ichthyologique. Copenhagen, Conseil Permanent International pour 1' Ex- ploration de la Mer, 2 pp. GiGLiOLi, E. H. 1880. Elenco dei Mammiferi, degli Uccelli e dei Rettili Ittiofagi Ap- partenenti alia Fauna Italica e Catalogo degli Anfibi e dei Pesci Italiani. Firenze: Staniperia Reale. 85 pp. Gilbert, C. H. 1905. The deep-sea fishes of the Hawaiian Islands. In The Aquatic Re- sources of the Hawaiian Islands. II, D. S. Jordan and B. W. Evermann (eds. ). Bull. U. S. Fish. Comm., 23: 575-716. Gilchrist, J. D. F. 1922. Report No. 2 for the year 1921. Fish. Mar. Biol. Surv. South Africa, Rep. No. 2: 1-84. , AND C. vox BoxDE. 1924. Deep-sea fishes procured by the S. S. "Pickle" (Part II). Fish. Mar. Biol. Surv. South Africa, Rep. No. 3, Spec. Rep. VII: 1-24. GoxzALES, T. 1946. Contributo alia conoscenza dello s\iluppo post-embrionale in Micro- ichtJit/s coccoi, Riippell. Boll. Pesca Piscicolt. Idrobiol., Ser. 22, 1 ( 1 ) : 39-46. GooDE, G. B., AXD T. H. Beax. 1881. De- scription of a new species of fish, Apogon ))andionis, from deep water off the mouth of Chesapeake Bav. Proc. U. S. Nat. Mus., 4: 160-161. , AXD . 1896. Oceanic Ichthy- ology. Wa.shingtou, D. C: Smithsonian In- stitution. 553 pp. GosLixE, \V. A. 1961. The perciform caudal skeleton. Copeia, 1961(3): 265-270. , AXD \'. E. Brock. 1960. Handbook of Hawaiian Fishes. Honolulu: Univ. Hawaii Press. 372 pp. Giinther, A. 1859. Catalogue of the Fishes in made at St. Helena by J. C. Melli.ss, Esq. Proc. Zool. Soc. London, pp. 22.5-228. Haxeda, Y., F. H. Johxsox, and O. Shi.mo.mura. 1966. The origin of luciferin in the luminous ducts of Pampriacmithus ransonncti, Pem- plicris kltiiizingcii, and Apogon cllioti. In Bioluminescence in Progress, F. H. Johnson and Y. Haneda (eds.). Princeton: Princeton Univ. Press, pp. 533-545. Holt, E. W., AND W. L. Calderwood. 1895. Survey of fishing grounds, west coast of Ireland, 1890-1891. Report on the rarer fishes. Scient. Trans. R. Dublin Soc, Ser. 2, 5: 361-524. HuHBS, C. L., AND K. F. Lacler. 1958. Fishes of the Great Lakes Region. Ann Arbor: Univ. Michigan Press. 213 pp. I\\Ai, T. 1959. Notes on the luminous organ of the apogonid fish, Siphaiiiia incijiinoi. Ann. Mag. Natur. Hist., Ser. 13, 2(21): 545-550. Jordan, D. S. 1917. Notes on Glossamia and related genera of cardinal fishes. Copeia, No. 44: 46-47. , AND C. H. Gilbert. 1882. Synopsis of the fishes of North America. Bull. U. S. Nat. Mus., 16: 1-1018. , AND E. K. Jordan. 1922. A hst of the fishes of Hawaii, with notes and de- scriptions of new species. Mem. Carnegie Mus., 10(1): 1-92. Kamohara, T. 1952. Revised descriptions of the offshore bottom-fishes of Prov. Tosa, Shikoku, Japan. Rep. Kochi Univ. Nat. Sci., No. 3: 1-22. Koefoed, E. 1952. Zeomorphi, Percomorphi, Plectognathi. Rep. Sci. Res. Michael Sars N. Atl. Deep-Sea Exped. 1910, 4, Pt. 2(2): 1-26. Lowe, R. T. 1841. A synopsis of the fishes of Madeira; with principal synonyms, Portu- guese names, and characters of the new genera and species. Trans. Zool. Soc. London, 2(14): 173-200. LozANO, L. 1934. Algunos peces pelagicos o de profundidad procedentes del Mediterraneo occidental. Bol. Soc. E.span. Hist. Natur., 34: 85-92. Matsubara, K. 1936. Biometry of two species of Japanese cardinal-fishes, with special reference to their taxononn. J. Imp. Fish. In.st. Tokyo, 31: 119-1.30. Maurin, C. 1968. Ecologie ichthyologiques des fonds Chalutables atlantiques (de la baie ibero-marocaine a la Mamitanie) et de la Mediterranee occidentale. Theses presentees a la Faculte des Sciences de I'Universite de Nancy. No. d'Ordre: A. O. 2 182. 145 pp. 202 Bulletin Museum of Comparative Zoology, Vol. 146, No. 3 Mayer, G. F. 1972. Systematics, functional anatomy, and ecology of the caidinalfish genus Epigonus ( Apogonidae ) . Ph. D. Thesis, Hanard University. 190 pp. Mayr, E. 1969. Principles of Systematic Zo- ology. New York: McGraw-Hill, Inc. 428 pp. Mead, G. W., and M. G. Bradbury. 1963. Names of bones. In Fishes of the Western North Atlantic, H. B. Bigelow and W. C. Schroeder (eds.). Mem. Sears Found. Mar. Res., 1(3): 20-23. MoNOD, T. 1968. Le complexe urophore des poissons teleosteens. Mem. Inst. Fondam. Afr. Noire, No. 81. 705 pp. MoREAU, E. 1881. Histoire Naturelle des Pois- sons de la France. Vol. II. Paris: G. Masson. 572 pp. MoRELAND, J. 1957. Appendix 6: Report on the fishes. New Zealand Dep. Sci. Ind. Res. Bull., 122: 34. Navarro, F. de P. 1942. Nota preliminar sobre los paces de la costa de Africa, desde el Cabo Bojador a la Bahia de Tanit ( Resultados de una compana industrial de pesca d'arrastre). Bol. Real Soc. Espan. Hist. Natur., 40(5-6): 189-214. , F. LozANO, J. M. NovAz, E. Otero, AND J. Sainz Pardo. 1943. La pesca d' arrastre en los fondos del Cabo Blanco y del Banco Arguin (Africa Sahariana). Trab. Inst. Espaii. Oceanogr. Numero 18. 225 pp. Nohre, a. 1935. Descricao dos Peixes de Portugal. Vertebrados (mamiferos, reptis e peixes). I. Fauna Marinha de Portugal. Porto. 574 pp. Norman, J. R. 1939. Fishes. Sci. Rep. John Murray Exped., 7(1): 1-116. . 1957. A Draft Synopsis of the Orders, Families and Genera of Recent Fish and Fish-like Vertebrates. London: British Mu- seum (Natural History). 649 pp. Nowlin, W. D., jr., and H. J. McLellan. 1967. A characterization of the Gulf of Mexico waters in winter. ]. Mar. Res., 25(1): 29-59. Osomo, B. 1898. Da distribviicao geographica dos peixes e crustaceos colhidos nas pos- sessoes Portuguezos d'Afrique occidental e existentes no Museu Nacional de Lisboa. J. Sci. Math. Phys. Natur. Acad. Real Sci. Lisboa, Ser. 2, 5: 185-202. PiETSCHMANN, V. 1930. Rcuiarks on Pacific fishes. Bull. Bernice P. Bi.shop Mus., 73: 1-24. Poll, M. 1954. Poissons IV. Teleosteens acanthopterygiens (premiere partie). Re- sidtats scientifiques. Expedition Oceano- graphique Beige dans les Eaux Cotieres Africaines de I'Atlantique Sud (1918-1949), 4(3A): 1-390. Radcliffe, L. 1912. Descriptions of fifteen new fishes of the family Gheilodipteridae, from the Pliilippine Islands and contiguous waters. Proc. U. S. Nat. Mus., 41(1868): 431-446. Rafinesque, G. S. 1810. Indice d' Ittiologia Siciliana; Ossia, Gatalogo Metodico dei Nomi Latini Italiani, e Siciliani dei Pesci, che si Rinvengono in Sicilia: Disposti Secondo un Metodo Naturale e Seguito da un Appendice che Gontiene la Descrizione di Alcuni Nuovi Pesci Siciliani. Messina: Presso Giovanni del Nobolo. 70 pp. Risso, A. 1810. Ichthyologie de Nice ou Histoire naturelle des poissons du department des Alpes Maritimes. Paris: Chez F. Schoell. 388 pp. Royce, W. R. 1957. Statistical comparison of morphological data. In Contributions to the Study of Subpopulations of Fishes, J. G. Marr (coordinator). U. S. Fish. Wildl. Serv. Spec. Sci. Rep. Fish., No. 208: 7-28. RuppELL, W. 1852. \'erzeichniss der in dem Museum der Senckenbergisclien naturfor- schenden Gesellschaft aufgestellten Samm- lungen. Vierte Abteilung. Fische und deren Skelette. Frankfurt-a.M.: J. D. Sauerliinder. 40 pp. Sakamoto, K. 1930. Buccal incubation in the percoid fish, Apogon lineatus T. & S. J. Imp. Fish. Inst. Tokyo, 26( 1 ) : 9-10. ScHULTZ, L. p. 1940. Two new genera and three new species of cheilodipterid fishes, with notes on the other genera of the familv. Proc. U. S. Nat. Mus., 88(3085): 403-423. Scott, T. D. 1962. The Marine and Fresh Water Fishes of South Australia. Adelaide: South Australian Branch of the British Sci- ence Guild. 338 pp. Simpson, G. G., A. Roe, and R. C. Lewontin. 1960. Quantitative Zoology. New York: Har- court, Brace and World, Inc. 440 pp. Smith, G. L., E. H. Atz, and J. C. Tyler. 1971. Aspects of oral brooding in the cardinalfish Cheilodiptcius af finis Poey (Apogonidae). Amer. Mus. Novitates, No. 2456: 1-11. Smith, J. L. B. 1949a. Forty-two fishes new to South Africa, with notes on others. Ann. Mag. Natur. Hist., Ser. 12, 2: 97-111. — . 1949b. The Sea Fishes of Southern Africa. South Africa: Central News Agenc>-, Ltd. 550 pp. 1961. Fishes of the family Apogonidae of the western Indian Ocean and the Red Sea. Ichth%ol. Bull. Rhodes Univ., No. 22: 373-418. Sparta, A. 1950. Su di una nuova specie di Micwichthi/s: M. sanzoi (n. sp.). Boll. Pesca, Piscicolt. Idrobiol., Ser. 26, 5(2): 202-206. EricoNus Systematics • Mayer 203 Stein'dachner, F. 1891. lehthNologischc Bei- triiKt' (X\^). i'her ciiiige scltenc iiiul ucnc Fisc'luiitc'ii aus ck'in caiuiiisclu'ii Aicliipcl. Sitzber. Akad. VViss. Wien, 100(1): 343-374. . 1907. Fi.sche aiis Siidaiabicn imd Sokotra. Denkschr. Akad. Wiss. Wien, 71: 123-168. Taylok, W. R. 1967. An enz\-matic method of elearing and staining small \eitel)iates. Proe. U.S. Nat. Mns., 122(3596): 1-17. Tinker, S. W. 1944. Hawaiian Fishes. Ilono- kdii: Tongg PnbHshing Company. 404 pp. ToRTONESE, E. 1952. Un percoide marino e batifilo nno\o per Tittiofauna itaHana (Epi- ^oinis clciilicuUitiis Dienz.). Boll. Pesca Pi.scicolt. Idrobiol. Ser. 28, 7(1): 72-74. , AND L. C. QuEiROLO. 1970. Contribute alio studio dell ittiofauna del Mar Ligure orientale. Ann. Mus. Civ. St. Natur. Genova, 78: 21-46. Vaillaxt, L. 1888. Poissons. E.xpeditions Sci- entifiques du Travailleur et du Talisman pendant les Annees 1880, 1881, 1882, 1883. Paris: G. Masson. 406 pp. V^ALENCiENNES, A. 1830. Additions et correc- tions anx tomes II, III, IV, et V. In Historic Naturelle des Poissons, Tome Sixieme, G. Cuvier and A. Valenciennes. Paris: F. G. Levrault, pp. 495-559. . 1837-1844. Ichthyologie des iles Can- aries, on Histoire naturelle des poissons raportes par M. M. Webb et Berthelot. In Histoire Naturelle des lies Canaries, P. B. Webb and S. Berthelot (eds.). Vol. 2, Pt. 2. Paris, pp. 1-109. [pp. 1-8 published Dec. 1842; plate 1 published May 1837.] Weuer, M., and L. F. de Beaufort. 1929. The Fishes of the Indo-Australian Archipelago. y. Anacanthini, Allotriognathi, Heterosomata, Berycomorphi, Percomorplii: Families Kuhli- idae, Apogonidae, Plesiopidae, Pseudo- plesiopidae, Priacanthidae, Centropomidae. Leiden: E. J. Brill, Ltd. 458 pp. Whitley, G. P. 1935. Studies in ichthyology. No. 9. Rec. Aust. Mus. Sydney, 19(4): 215-250. -. 1940. Illustrations of some Australian fishes. Aust. Zool., 9(4): 397-428. . 1968. A check-list of the fishes re- corded from the New Zealand region. Aust. Zool., 15(1): 1-102. Williams, F. 1968. Report on the Gnincan Trawling Survey, \'ol. I. General Report. OAU Scientific, Technical and Research ('oninn'ssion, Ful)licali<)n 99. 828 pp. W'iisi, (;. 1961. Slratitication and Circulati(m in the Antillean-Caribbcan Basins. New York: C]()luml)ia l'ni\ . Press. 201 pp. APPENDIX The following chart lists all meristic and niorphometric data for the holotypcs of E. oUiiolepis sp. nov. and E. pectinifer sp. nov. Measurements are given in milli- meters. E. oligolepis E. pectinifer USNM 2 077 18 USNM 207725 MERISTIC DATA Dorsal fin VII-I, 10 VII -I, 9 Anal fin II, 9 II, 9 Pectoral fin 18 15 Pelvic fin I, 5 I, 5 Lateral line sc ale s 34 47 Gill r ake r s 31 27 Pyloric caeca 10 6 Ve r tebr ae 10 + 15 10 + 15 Pleural ribs 7 8 E p i p 1 e u r a 1 rib s 7 6 MORPHOMETRIC DATA SL 90. 8 114. 3 HL 33. 2 40. 5 Body depth 21.2 28. 1 Head height 17. 1 18. 8 Eye diameter 14. 5 16. 1 Snout 1 e n g t li 7. 8 10. 9 I n t e r 0 r b i t a 1 w i idth 8. 4 10. 0 Maxillary leng th 15. 6 18. 2 Lower jaw len gth 16. 3 18. 5 Caudal p c nid-]ike tracheal system, but also the lamelliform claw tufts so characteristic of anyphaenids. Further, the genitalia are close to those of the anyphaenid genus Oxysouui, and the body form is similar to that of several species of anyphaenids known from Chile, Peru, and Argentina. For these reasons, the family Amauro- bioididae is newly synonymized with Anyphaenidae in the taxonomic section of this paper. Thus the problem of the correct macro- taxonomic placement of Anyphaenidae has been clarified but not solved by this study of the groups with which the family has been associated in the past. Futiu-o work should start with an examination of the family Miturgidae (as construed by Leht- inen ) . Although it was necessary to limit the scope of the detailed revision to the man- ageable number of species occurring north of Mexico, all available specimens from other areas were examined to gain an overview of the family. Preliminary im- pressions indicate that the family probably originated in the southern half of South America with subsequent radiations north- ward. As indicated by the ability of Amaurohioides to withstand prolonged sub- mersion, it is likely that early anyphaenids were able to survive hydrochore dispersal by rafting, etc., across considerable ex- panses of water. GENERIC PROBLEMS IN THE ANYPHAENIDAE The generic taxonomy of anyphaenids is currently chaotic. Every author who has worked with the group, including Petrunke- vitch (1930), Bryant (1931) and Chicker- ing (1937), has expres.sed frustration at the confusion and ambiguity in the use of many of the most common generic names. One of the principal causes of this con- fusion is the interesting e\olutionary pat- tern encountered time and again witliin 210 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 this family: species tend to occur in groups that are remarkably homogeneovis in genitalic structure but quite distinct from other such groups. Often many of the species in these groups are sympatric, are found in a rather limited area and are clearly the result of radiation within that area. An excellent example of this is the occurrence of nine closely related species of the Anyphaena celer species group in the mountains of southeastern Arizona. It is tempting to consider each of these groups a genus, as unambiguous key char- acters are then available to distinguish genera. Such an approach would at least double the number of genera found in the United States, and, if applied to the Central and South American fauna, would neces- sitate the creation of a vast number of new genera. If, instead, characters refer- ring to the general body form are used, a more workable classification in terms of both number and size of genera results. Unfortunately, this makes the unambiguous definition of genera much more difficult and makes keys to genera awkward and cumbersome. With either approach, how- ever, reliable genera composed of mo- nophyletic groups of species can be estab- lished. The second approach to anyphaenid clas- sification has been taken by the majority of former authors, and is continued in this work. Thus the European genus Any- phaena is used for the bulk of the any- phaenids occurring in the United States, even though only one of our species, Any- phaena aperta, is actually a close relative of the European Anyphaena accentuata, type species of the genvis. Nonetheless, all the species here included in Anyphaena share a basic body form. The neotropical genus Wulfila is used for all the pale, long-legged species, even though they are genitalically quite diverse; the other genera used here are similarly construed. Although this system is not wholly satisfactory, it seems decidedly better than creating a host of new generic names that are likely to fall into synonymy when a detailed ge- neric revision of the group as a whole can be carried out. METHODS Tracheae were examined by dissecting away the dorsal cuticle of the abdomen and boiling the spider in ten percent sodium hydroxide for ten minutes. By this method, all the soft structures in the abdomen are digested away, leaving the tracheae intact. Types of the new species are being deposited in the American Museum of Natural History, New York City, and the Museum of Comparative Zoology, Harvard University. Type depositories are abbrevi- ated as follows: AMNH — American Mu- seum of Natural History, BMNH — British Museum, Natural History, MCZ — Museum of Comparative Zoology. Measurements and drawings were made with a standard ocular grid. Measurements of gross morphological featiu-es are ac- curate to ~ 0.04 mm; measurements of ocular featiu'es are accurate to ~ 0.01 mm. Rather than selecting a small number of measurements and providing means and standard deviations for these on the basis of a small series of specimens, one male and one female of each species were measured in detail. As only one of the species included here shows any significant variation in size, this procedure was deemed more informative. Actual measurements are given rather than ratios since in many cases (e.g., Anyphaena catalina and A. arhida) closely related species differ sig- nificantly in size but not in their relative proportions. Most of the measurements taken are self-explanatory, though a few need furdier comment. Cephahc width refers to the width of the carapace at a point just behind the posterior median eyes, and thus provides an indication of the degree to which the carapace is nar- rowed in front. The difficult problem of accurately de- scribing die eye relationships has been Spider Family Anyphaenidae • Platnick 211 solved by providing a set of measurements from wliich it is possible to reconstrnct, using grapli paper, the exact eye arrange- ment. Diameters are given using the con- ventional abbreviations (AME = anterior median eye, ALE = anterior lateral eye, PME = posterior median eye, PLE = posterior lateral eye). The length of each eye row is measured from the lateral edge of one lateral eye to the lateral edge of tlie other lateral eye. Curvature of the eye rows is described as viewed frontally, not dorsally. This was accomplished by posi- tioning the spider in sand, a technique found most useful for making all the measurements. The dimensions of the median ocular quadrangle (MOQ) are given, as well as the distances between each of the eyes. The latter measurements extend between the edges of the lenses of the eyes under consideration (not just between the dark circles surrounding each eye). The relative length and thickness of each leg is indicated by the tibial length index — the tibial width divided by the tibial length, with the result multiplied by 100 to obtain a whole number. All tibial measurements were taken from a dorsal view and refer to the maximum lengths and widths. The lower the tibial index, the longer and thinner the leg; conversely, the higher the index, the shorter and thicker the leg. In practice the index varies from around 3 to 35. Ventral spination of the leg segments is indicated by the standard formula in which the number of spines on the proximal, median and distal thirds of the leg segment are given. Only ventral spines, not lateral ones, are included, and any even number in the formula may be taken to represent a pair of spines. Unless the last number is followed by an asterisk, the last pair of spines is terminally located. Thus, for example, the formula 2-2-2* indicates that the segment bears three pairs of ventral spines, the last pair of which is not termi- nally located. The term "spine" is used in its conventional arachnological sense and rc'fers to the moxable macrosetae found on the legs. Similarly, the term "clypeus" is used to refer to the area between the anterior eye row and the anterior edge of tlie carapace and not to the small sclerite folded under the carapace. Since neither usage of tht> term reflects certain knowledge of homology with the insect clypeus, the old and established usage should be main- tained. Scale lines for the drawings always equal 0.1 mm. Each scale line applies to all consecutively numbered drawings imtil a new scale line appears. Exceptions are noted in the captions. TAXONOMY Anyphaenidae Anyphaenidae Bertkau, 1878, Arch. Naturg., 44: 358, 379. Tvpe genus Anyphaena Sundevall, 1833. Amaurobioididae Hickman, 1949, Pap. Proc. Roy. Soc. Tasmania, 1948: 31. Tvpe genus A77iauro- hioides O.P.-Cambridge, 1883. NEW SYN- ONYMY. Diagnosis. The combination of the ad- vanced tracheal spiracle and the lamelli- form claw tufts will serve to distinguish the anyphaenids from all other families. Description. Chelicerae diaxial, not fused together at base. Labium free. Without cribellum or calamistrum. With one pair of book lungs and a tracheal spiracle lo- cated considerably anterior to the spin- nerets, most often midway between spin- nerets and epigastric furrow, sometimes closer to one or the other. Eight eyes in two rows. Six spinnerets, anterior spin- nerets approximate, colulus represented only by hairs, anal tubercle unmodified. Legs prograde, metatarsi and tarsi I and II scopulate, tarsi with two toothed claws and claw tufts composed of lamelliform setae. Key to Genera IN America north of Mexico la. Tracheal spiracle luucli closer to epigastric furrow tlian to spinnerets ..Aysha 212 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 lb. Tracheal spiracle roughly midway between epigastric furrow and spinnerets 2 2a. Legs very long and thin. Leg I greatly elongated, tibial index (width/length X 100) usually 5 or less Wulfila 2b. Legs normal, tibial index of leg I usually 8 or more 3 3a. Chelicerae with 2 retromarginal teeth Oxysoma 3b. Chelicerae with 4-9 retromarginal denticles 4 4a. Carapace usually with two dark paramedian longitudinal bands; chelicerae not produced forward; femora not much darker than other leg segments Anijphaena 4b. Carapace without dark paramedian longi- tudinal bands; either chelicerae produced forward or femora much darker than other leg segments Teudis Anyphaena Sundevall Amjphaena Sundevall, 1833, Conspectus Arachn., 28. Type species by monotypy Aranea ac- centuata Walckenaer, 1802. Diagnosis. The combination of the fol- lowing characters will serve to distinguish the genus in America north of Mexico: trachael spiracle roughly midway between epigastric furrow and spinnerets, leg I not greatly elongated, chelicerae with 4-9 retromarginal denticles and not produced forward, femora not much darker than other leg segments. The carapace usually has two dark paramedian longitudinal bands. The genus is used here in a very broad sense; this prevents simple diagnosis, and makes detailed descriptions of each species group more meaningful than a description of the whole genus. Uncertain names. Types of the follow- ing species were unavailable and are too poorly described to permit identification: Cluhiona agresiis Hentz, 1847, type de- stroyed; Chihiona fallens Hentz, 1847, type destroyed, Cluhiomi suhlurida Hentz, 1847, type destroyed; Amjphaena argentata Becker, 1879, type lost; and Amjphaena striata Becker, 1879, type lost. The three Hentz Cluhiona species were transferred to Anijphaeiui by Marx (1890), but there is little justification for this in the vague descriptions. All the above names are regarded as nomina cluhia. Species groups. Although there seem to be several species groups of Amjphaena in the Neotropic region, only four occur north of Mexico. The celer group is the largest; it has representatives at least as far south as Panama and probably contains over thirty species. The pectorosa and pacifica groups are closely related and occur com- monly in Mexico as well as the United States; it is difficult to place females in one group or the other unless the male is also known; they probably contain together at least twenty species. The accentuata group is predominantly Palearctic and prob- ably contains at least five species. Key to Species Groups la. Metatarsi I and II with one pair of ventral spines accentuata group lb. Metatarsi I and II with two pairs of ventral spines 2 2a. Retrolateral tibial apophysis of males bifid, with ventral prong elongated ( Figs. 18-20, 25-32). Epigynum of females with a hood (Figs. 21, 23, 33, 36, 37, 39-42) ._ celer group 2b. Retrolateral tibial apophysis of males not bifid or elongated (Figs. 55-58, 69-71). Epigynum without a hood (Figs. 66, 67, 72, 74, 77, 79) 3 3a. Eastern United States. Coxae III and IV of males with pointed spins (Figs. 59-62). Female epigyna on broad sclerotized plates (Figs. 74, 77, 79); internal genitalia lacking long ducts (Figs. 75, 78, 80) pectorosa group 3b. Western United States. Coxae III and IV of males without pointed spurs, though rounded knobs may be present. Female epig>'na not on broad sclerotized plates (Figs. 66, 67, 72); internal genitalia with long, sometimes coiling, ducts (Figs. 68, 73, 76) _._. pacifica group Anyphaena celer Group Diagnosis. Males of the celer group may be recognized by their retrolateral tibial apophysis, which is usually bifid with an elongated ventral prong (Figs. 18, 20, 26). Females have a characteristic epigynum consisting of a hood, two sidepieces and a midpiece (Figs. 9, 33), though the mid- SpiDEii Family Axyphakxidae • Pkilnick 213 piece is reduced in A. crebrispimi ;ind A. (lixiana (Figs. 21, 23). Description. Total length 3-7 nnn, with males of most species between 3.3-4.6 mm, females of most species between 4.1-5.9 mm. Carapace longer than wide, narrowed in front to less than half its maximum w idth. Clypens height greater than anterior median eye diameter. Posterior median, posterior lateral and anterior lateral eyes subeqiial in size, larger than anterior medians. Procurved posterior eye row longer than recin'\'ed anterior row. Median ocular ({nadrangle longer than wide in front, wider than long in back. Anterior median eyes separated by their diameter, by their radius from anterior laterals. Posterior medians separated by their diam- eter, slightly closer to posterior laterals than to each other. Anterior laterals separated by their radius from posterior laterals. Sternum longer than wide, un- modified. Chelicerae with 4-5 promarginal teeth and 6-9 reti-omarginal denticles. Abdomen longer than wide, tracheal spiracle midway between epigastric fur- row and base of spinnerets. Leg formula 1423. Metatarsi I and II with two pairs of \'entral spines. Males often with femur III thickened distally, set with stiff short setae ventrally; tibia III ventral spines thickened, cone-like; coxae set with clumps of stiff short setae. Palpus with an elon- gated median apophysis, retrolateral teg- ular apophysis, conspicuous curving em- bolus and conductor. Retrolateral tibial apophysis bifid, with dorsal prong reduced in some species. Epigynum with hood, two sidepieces and midpiece; two simple spermathecae. Variation. None of the species in this group show any significant individual or geographic intraspecific variation in struc- tin-e, size or coloration. Key to Species la. Dorsal and \entral prongs of retrolateral tibial apophysis ( RTA ) roughly equal in length (Figs. 18, 19); epigynal hood wide, more than four times the minimum width of epigynal sidepiece ( Figs. 9, 11); east- ern U.S 2 II). Ventral prong of retrolateral tibial apophy- sis ( RTA ) nuieh longer than dorsal prong (as in Figs. 26, 27); epigynal hood narrow, less than four times the minimum width of epigynal sidepiece ( as in Figs. 33, 36); western U.S. 3 2a. Dorsal prong of RTA broad, with a trans- lucent ridge (Fig. 18); epigynal hood a thick oval, sidepieces straight (Fig. 9) — - - — _.. celer 21). Dorsal prong of RTA narrow, without a translucent ridge (Fig. 19); epigynal hood a thin oval, sidepieces rounded ( Fig. 1 1 ) - - _ inaculata 3a. Base of RTA expanded into a broad triangle (Fig. 20); retrolateral tegular apophysis prolonged medially (Fig. 3); epigynal sidepieces more than three times the width of epigynal hood (Fig. 21) - crebrispina 3b. Base of RTA not expanded; retrolateral tegular apophysis not prolonged medialK ; epigynal sidepieces less than three times the width of epigynal hood 4 4a. Dorsal prong of RTA bearing a sharp spur (Fig. 25); epigynal midpiece greatly re- duced, sidepieces widely separated pos- teriorly (Fig. 23) dixiana 4b. Dorsal prong of RTA without a spur; epigynal midpiece conspicuous, sidepieces approximate posteriorly 5 5a. Males '. 6 5b. Females 14 6a. Dorsal prong of RTA with two triangular processes separated by a concave notch ( Fig. 26 ) judicata 6b. Dorsal prong of RTA witliout triangular processes „. 7 7a. Dorsal prong of RTA with a long recurved hook (Fig. 29) autumna 7b. Dorsal prong of RTA without a long re- curved hook 8 8a. Dorsal prong of RTA witli a basal hook (Figs. 31, 38) _.-.. 9 8b. Dorsal prong of RTA without a basal hook 10 9a. Conductor and retrolateral tegular apophy- sis recurved (Fig. 15) catalina 9b. Conductor and retrolateral tegular apoph- ysis not recurved ( Fig. 17 ) arhida 10a. Dorsal prong of RTA a shaiply pointed spike ( Fig. 32 ) liespar 101). Dorsal prong of RTA not a sharply pointed spike 1 1 11a. Fmbolus with a conspicuous enlargement (Figs. 7, 13) .12 111). Embolus without a con.spicuous enlarge- ment (Figs. 21, 33) 13 214 Bulletin Museum of Comparative Zoolog,ij, Vol. 146, No. 4 J? I ^J'', ,^-^ m-p. Anyphaeno maculota \ q\ —>> o- I -* J \ \A- 1 1 ^~« ', >-:-: Anyphoena crebrispino /— \\v Anyphaena dixiana j»- \ v^\ \ Anyphaena rita ' V f 1 ~T 1 1 \h~^ ' 1 Anyphaena cochise V Map 1. Distributions of Anyphaena arbida, A. autumna, A. catalina, A. celer, A. cochise, A. crebrispina, A. dixi- ana, A. gibboides, A. hespar, A. judicata, A. maculata, A. marginalis and A. rita. 12a. Dorsal prong of RTA more than half the length of ventral prong ( Fig. 35 ) cochise 12b. Dorsal prong of RTA less than half the length of ventral prong (Fig. 28) rita 13a. Median apophysis sharply pointed; con- ductor short, bent (Fig. 14); Oregon and Utah gibboides 13b. Median apophysis rounded; conductor long, straight (Fig. 6); Arizona and New Mexico marginalis 14a. Epigynal hood wider than long; midpiece not wider than hood, without constric- tions; sidepieces very wide (Fig. 40); Oregon and Utah gibboides 14b. Epigynal hood as long as wide or midpiece wider than hood or sidepieces narrow; Arizona and New Mexico 15 15a. Epigynal midpiece a very broad triangle ( Fig. 37 ) rita 15b. Epigynal midpiece othenvise 16 16a. Spermathecae much further apart pos- teriorly than anteriorly ( Fig. 49 ) _-, hespar 16b. Spemiathecae as far apart anteriorly as posteriorly 17 17a. Epigynal hood much wider than long ( Fig. 39 ) _ autumna 17b. Epigynal hood as long as wide —18 18a. Epigynal midpiece less than twice the length of epigynal hood (Fig. 41) catalina 18b. Epigynal midpiece more than twice the length of epigynal hood ...19 19a. Epigynal midpiece a short triangle (Fig. 33 ) judicata 19b. Epigynal midpiece an elongate triangle (Fig. 36) marginalis Anyphaena celer (Hentz) Map 1; Figures 1, 9, 10, 18 Chibiona celer Hentz, 1847, J. Boston Soc. Natur. Hist., 5: 452, pi. 23, fig. 20 ( 9 ). Male holo- type, female allotype from Alabama and North Carolina in the Boston Soc. Natur. Hist. (Boston Museum of Science), destroyed by beetles. Anyphaena incerta Keyserling, 1887, Verb. zool. hot. Ces. Wien, .37: 452, pi. 6, fig. 22 ( $ ). Female holotype from Cambridge, Massachu- setts, in MCZ, examined. Emerton, 1890, Trans. Connecticut Acad. Sci., 8: 186, pi. 6, figs. 2-2d, $,9. Anyphaena celer, Simon, 1897, Hist. Natur. Araign., 2: 96. Bryant, 1931, Psyche, 38: 111, pi. 6, fig. 9, pi. 8, figs. 25, 28, $, 9. Chickering, 1939, Pap. Michigan Acad. Sci., 24: 51, figs. Spider Family Axyphakxidae • PJaliiick 215 Plate 1 Figures 1-8. Left palpi, ventral view. Figures 9. 11. Epigyna, ventral view. Figures 10, 12. Internal genitalia, dorsal view. 1,9,10. /Anyphaena ce/er (Hentz). 2,11,12. Anyphaena maculata (Banks). 3. Anyphaena crebri- sp'ma Chamberlin. 4. Anyphaena dixiana (Chamberlin and Woodbury). 5. Anyphaena judicata O. P. -Cambridge. 6. Anyphaena marginalis (Banks). 7. Anyphaena rita new species. 8. Anyphaena autumna new species. 216 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 t>' 1-4, $, 9. Comstock, 1940, Spicier Book, rev. ed., p. 577, figs. 634-6.35, $,9. Kaston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 407, figs. 1471-1476, $,9. Roewer, 1954, Katalog der Araneae, 2:528. Bonnet, 1955, Bibliographia Araneorum, 2: 343. Gatjenna celer, Comstock, 1912, Spider Book, p. 563, figs. 634-635, $,9. Diaxternally visi- ble midpiece, is totally unlike that of any other species in this group ( Fig. 21 ) . Male (Los Angeles Co., California). Coloration as in AnypJiaena celer. Total length 4.61 mm. Carapace 2.00 mm long, 1.57 mm wide, cephalic width 0.74 mm, clypeus height 0.10 mm. Eyes: diameters (mm): AME 0.07, ALE 0.10, PME 0.10, PLE 0.10; anterior eye row 0.43 mm long, recurved; posterior eye row 0.56 mm long, prociuved; MOQ length 0.25 mm, front width 0.18 mm, back width 0.29 mm; eye interdistances (mm): AME-AME 0.04, AME-ALE 0.02, PME-PME 0.09, PME- PLE 0.09, ALE-PLE 0.04. Sternum 1.10 mm long, 0.S8 mm wide. Chelicerae 0.55 mm long with 5 promar- ginal teeth and 8 retromarginal denticles. Abdomen 2.65 mm long, 1.58 mm wide. Epigastric furrow 0.79 mm from tracheal spiracle, spiracle 0.68 mm from base of spinnerets. Tibial lengths (mm) and indices: I L69, 13; II 1.51, 15; III 1.06, 22; IV 1.69, 14. Ventral spination: tibiae I 2-2-2*, II 1-2- 2\ III 1-2-2, IV 2-2-2; metatarsi I, II 2- 2-0, III 2-0-2, IV 2-2-2. Modifications of third leg as in A. celer. Palpus as in Figures 3, 20. Female (Los Angeles Co., California). Coloration as in male of A. celer. Total length 4.39 mm. Carapace 1.85 mm long, 1.37 mm wide, cephalic width 0.77 mm, clypeus height 0.08 mm. Eyes: diameters (mm): AME 0.07, ALE 0.09, PME 0.09, PLE 0.09; anterior eye row 0.41 mm long, recurved; posterior eye row 0.56 mm long, procurved; MOQ length 0.23 mm, front width 0.18 mm, back width 0.28 mm; eye interdistances (mm): AME-AME 0.04, AME-ALE 0.02, PME-PME 0.10, PME- PLE 0.08, ALE-PLE 0.04. Sternum 1.08 mm long, 0.86 mm wide. Chelieerac> 0.64 mm long with 4 promar- ginal teeth and 9 retromarginal denticles. Abdomen 2.99 nun long, 1.98 mm wide. Epigastric furrow 0.95 mm from tracheal spiracle, spiracle 0.90 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.39, 15; II 1.31, 16; III 0.75, 28; IV 1.44, 14. Ventral spination: tibiae I 2-2-2*, II 1-2-0, III 1-1-0, IV 1-1-2; metatarsi as in male. Epigynum as in Figure 21, internal geni- talia as in Figure 22. Natural history. Mature males have been taken in November, mature females from early December through late April. Speci- mens have been taken by Berlese funnel sampling of grape bark. Distribution. Central and southern Cali- fornia ( Map 1 ) . Anyphaena dixiana (Chamberlin and Woodbury), new combination Map 1; Figures 4, 23, 24, 25 Gaijcnna dixiana Chamberlin and Woodbun', 1929, Proc. Biol. Soc. Washington, 42: 138, pi. 1, fig. 3 ( 9 ). Female holotype from St. Ceorge, Utah, in AMNH, examined. Roewer, 1954, Katalog der Araneae, 2: 540 ( G. dixima [sic]). Bonnet, 1957, Bibliographia Araneorum, 2: 1977. Anyphaena coloradensis Bryant, 1931, Psyche, 38: 112, pi. 6, figs. 9, 10, pi. 7, figs. .30, 33 { $, 9 ). Male holotype, female allotNpc from Boulder, Colorado, in MCZ, examined. Roewer, 1954, Katalog der Araneae, 2: 528. Bonnet, 1955, Bibliographia Araneorum, 2: 343. NEW SYNONYMY. Diafinosis. This distinctive species is closest to AmjpJuiena crebrispina, but may be quickly recognized by the spur borne on the dorsal prong of the RTA of males (Fig. 25) and the greatly reduced epigynal midpiece of females (Fig. 23). Male (Cochise Co., Arizona). Coloration as in Anyphaena celer except that posterior .spiimerets have dorsal surface sharply di- vided into dark brown lateral and pale orange median halves. Total length 3.85 mm. Carapace 1.67 222 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 mm long, 1.44 mm wide, cephalic width 0.65 mm, clypeus height 0.07 mm. Eyes: diameters (mm): AME 0.06, ALE 6.09, PME 0.09, PLE 0.10; anterior eye row 0.39 mm long, recurved; posterior eye row 0.51 mm long, procurved; MOQ length 0.25 mm, front width 0.16 mm, back width 0.26 mm; eye interdistances (mm): AME- AME 0.04, AME-ALE 0.03, PME-PME 0.08, PME-PLE 0.08, ALE-PLE 0.05. Sternum 0.96 mm long, 0.76 mm wide. Chelicerae 0.53 mm long with 4 promar- ginal teeth and 6 reti'omarginal denticles. Endites slightly invaginated at middle. Abdomen 2.56 mm long, 1.49 mm wide. Epigastric furrow 0.76 mm from tracheal spiracle, spiracle 0.76 mm from base of spinnerets. Tibial lengths (mm) and indices: I 1.69, 11; II 1.37, 14; III 0.81, 28; IV 1.44, 16. Ventral spination: tibiae I, II 2-2-2*, III, IV 1-2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2-2-2. Modifications of third leg as in A. celer. Palpus as in Figures 4, 25. Female (Cochise Co., Arizona). Colora- tion as in male. Total length 4.14 mm. Carapace 2.03 mm long, 1.57 mm wide, cephalic width 0.86 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.05, ALE 0.08, PME 0.09, PLE 0.10; an- terior eye row 0.43 mm long, recurved; posterior eye row 0.60 mm long, procurved; MOQ length 0.26 mm, front width 0.20 mm, back width 0.32 mm; eye interdis- tances (mm): AME-AME 0.09, AME- ALE 0.05, PME-PME 0.15, PME-PLE 0.09, ALE-PLE 0.07. Sternum 1.15 mm long, 0.86 mm wide. Chelicerae 0.71 mm long with 5 promar- ginal teeth and 8 retromarginal denticles. Abdomen 2.50 mm long, 1.69 mm wide. Epigastric furrow 0.60 mm from tracheal spiracle, spiracle 0.67 mm from base of .spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.46, 16; II 1.33, 17; III 0.94, 24; IV 1.49, 17. Ventral spination as in male. Epigynum as in Figure 23, internal geni- talia as in Figure 24. Natural history. Mature males have been taken from mid- August through mid-May, mature females from late September through late April. Specimens have been taken from 5400 to 9000 feet (1650-2750 m), in yellow pine/ oak and montane for- ests, in alfalfa, under dead agave and fre- quently in houses. Distribution. Northcentral Colorado south to western Texas, west to southern Cali- fornia (Map 1 ) . Anyphaena judicata O. P.-Cambridge Map 1; Figures 5, 26, 33, 34 Anijphaena iiidicata O. P. -Cambridge, 1896, Biologia Central! Americana, Aran., 1: 203, pi 26, fig. 4 { S ). Male holotype from Omiltemi, Guerrero, Mexico, in BMNH, examined. F. O. P.-Cambridge, 1900, Biologia Centrali Ameri- cana, Aran., 2: 96, pi. 7, fig. 9, $. Roewer, 1954, Katalog der Araneae, 2: 525. Bonnet, 1955, Bibliographia Araneormn, 2: 345. Diagnosis. Anyphaena judicata is most closely related to an unnamed Mexican species (or group of species) and has no close relatives among the species occur- ring north of Mexico. Males may be easily recognized by the distinctive form of the dorsal prong of the RTA (Fig. 26). The female epigynum is closest to that of A. niarginalis, but the midpiece is proportion- ately shorter and wider and the sidepieces are narrower and diminish in width an- teriorly smoothly, without the sharp de- crease in width shown by A. marginalis (Fig. 33). Male (Cochise Co., Arizona). Coloration as in Anyphaena celer, except that pos- terior spinnerets have entire dorsal surface dark brown. Total length 3.46 mm. Carapace 1.76 mm long, 1.44 mm wide, cephalic width 0.68 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.06, ALE 0.10, PME 0.09, PLE 0.10; anterior eye row 0.40 mm long, recurved; posterior eye row 0.52 mm long, procurved; MOQ length 0.26 mm, front width 0.17 mm, back width Spider Family Anyphaenidae • Plotnick 223 0.28 mm; eye interdistanees (mm): AME- AME 0.05, AME-ALE 0.03, PME-PME 0.11, PME-PLE 0.06, ALE-PLE 0.04. Sternum 0.95 mm long, ().6S mm wide. Chelicerae 0.56 mm long with 4 promar- ginal teeth and 7 retromarginal denticles. Abdomen 1.80 mm long, 1.15 mm wide. Epigastric furrow 0.61 mm from tracheal spiracle, spiracle 0.63 mm from base of .spinnerets. Tibial lengths (mm) and indices: I 2.25, 6; II 1.93, 8; III 1.01, 21; IV 1.66, 11. Ven- tral spination: tibiae I 4-2-2*, II 3-2-2', III 1-2-0, IV 1-1-2; metatarsi I, II, 2-2-0, III 2-0-2; IV 1-2-2. Femur III unmodi- fied. Tibia III ventral spine 1 on retrolat- eral side missing, ventral spine 2 thickened, cone-like. Coxae I, II and III ( but not IV ) with a small number of short, thick setae. Coxae III with a tiibercule. Palpus as in Figures 5, 26. Female (Cochise Co., Arizona). Colora- tion as in male. Total length 4.72 mm. Carapace 1.76 mm long, 1.37 mm wide, cephalic width 0.81 mm, clypeus height 0.06 mm. Eyes: diameters (mm): AME 0.07, ALE 0.10, PME 0.10, PLE 0.10; anterior eye row 0.44 mm long, recurved; posterior eye row 0.60 mm long, procurved; MOQ length 0.29 mm, front width 0.21 mm, back width 0.32 mm; eye interdistanees (mm): AME- AME 0.07, AME-ALE 0.03, PME-PME 0.13, PME-PLE 0.09, ALE-PLE 0.05. Sternum 0.97 mm long, 0.77 mm wide. Chelicerae 0.58 mm long with teeth as in male. Abdomen 3.13 mm long, 2.09 mm wide. Epigastric furrow 1.21 mm from tracheal spiracle, spiracle 1.31 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.69, 11; II 1.31, 14; III 0.88, 23; IV 1.66, 12. Ventral spination as in male. Epigynum as in Figure 33, internal geni- talia as in Figure 34. Natural history. Mature males have been taken from mid-June through mid-August, mature females from late March to Novem- ber, most ill July and August. Specimens have been taken from 5100 to 8000 feet (1550-2450 m), by sweeping and under rocks. Distribution. Arizona south to Guerrero, Mexico (Map 1). Anyptiaena marginalis (Banks), new combination Map 1; Figures 6, 27, 36, 43 Gayeima marginalis Banks, 1901, Proc. Acad. Natur. Sci. Philadelphia, 53: 574, pi. 2.3, fig. 22 ( 9 ). Female holotype from Beulali, San Miguel Co., New Mexico, was probabl>' de- posited in the MCZ along with the other types from this paper but was not found by Bryant when the MCZ t>pes were cataloged; lost, presumed destroyed. Roewer, 1954, Katalog der Araneae, 2: 540. Bonnet, 19.57, Biblio- graphia Araneorum, 2: 1978. Diagnosis. Amjphaena marginalis is most closely related to A. hespar, both species having a simple embolus and elongated conductor. Males of A. marginalis (Fig. 27), however, do not have the spine-like dorsal prong of the RTA of A. hespar, and females of A. marginalis (Fig. 36) do not have the conspicuous bulge in the epigynal midpiece which characterizes A. hespar females. Male (Graham Co., Arizona). Colora- tion as in Anyphaena celer. Total length 3.78 mm. Carapace 1.98 mm long, 1.60 mm wide, cephalic width 0.72 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.06, ALE 0.10, PME 0.08, PLE 0.10; anterior eye row 0.40 mm long, straight; posterior eye row 0.55 mm long, prociuved; xVlOQ length 0.20 mm, front width 0.17 mm, back width 0.28 mm; eye interdistanees (mm): AME- AME 0.05, AME-ALE 0.02, PME-PME 0.11, PME-PLE 0.08, ALE-PLE 0.04. Sternum 1.13 mm long, 0.81 mm wide. Chelicerae 0.54 mm long with 5 promar- ginal teeth and 6 retromarginal denticles. Abdomen 2.00 mm long, 1.33 mm wide. Epigastric furrow 0.52 mm from traclieal 224 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 spiracle, spiracle 0.59 mm from base of spinnerets. Tibial lengths (mm) and indices: I 1.67, 13; II 1.35, 16; III 1.03, 26; IV 1.62, 14. Ventral spination: tibiae I 4-2-2*, II 2-2- 2*, III, IV 1-2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2-2-2. Femur III unmodified. Tibia III ventral spine 1 on retrolateral side missing. Coxae unmodified. Palpus as in Figures 6, 27. Female (Graham Co., Arizona). Colora- tion as in male of A. celer. Total length 4.26 mm. Carapace 2.11 mm long, 1.55 mm wide, cephalic width 0.86 mm, clypeus height 0.08 mm. Eyes: diameters (mm): AME 0.07, ALE 0.10, PME 0.11, PLE 0.10; anterior eye row 0.44 mm long, straight; posterior eye row 0.64 mm long, procurved; MOQ length 0.30 mm, front width 0.19 mm, back width 0.33 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.03, PME-PME 0.12, PME-PLE 0.09, ALE-PLE 0.06. Sternum 1.05 mm long, 0.80 mm wide. Chelicerae 0.65 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 2.52 mm long, 1.53 mm wide. Epigastric furrow 0.67 mm from tracheal spiracle, spiracle 0.68 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.44, 18; II 1.21, 21; III 0.99, 25; IV 1.60, 17. Ventral spination as in male except tibia III 1-1-2. Epigynum as in Figure 36, internal geni- talia as in Figure 43. 'Natural history. Mature males have been taken from late August through late May, mature females in all months except Janu- ary and October. Specimens have been taken from 6000 to 9300 feet (1850-2850 m), in yellow pine/ oak forests and under rocks. I found this species in great abun- dance by sorting pine litter at Rustler's Park in the Chiricahua Mountains of south- eastern Arizona in August 1972. Distribution. Arizona, New Mexico and Colorado (Map 1). Anyphaena hespar new species Map 1; Figures 16, 32, 42, 49 Types. Male holotype, female paratype from Bear Canyon, Santa Catalina Moun- tains, Pima Co., Arizona, 8 December 1968 (Karl Stephan), deposited in AMNH. Male and female paratypes from Pima Co., Ari- zona, deposited in MCZ. The specific name is an arbitrary combination of letters. Diagnosis. Anyphaena hespar is most closely related to A. marginalis. Males of the former may be distinguished by the spine-like dorsal prong of their RTA (Fig. 32), females by the conspicuous bulge in their epigynal midpiece (Fig. 42). Male (Pima Co., Arizona). Coloration as in Anyphaena celer. Total length 3.13 mm. Carapace 1.62 mm long, 1.31 mm wide, cephalic width 0.59 mm, clypeus height 0.08 mm. Eyes: diameters (mm): AME 0.05, ALE 0.08, PxME 0.08, PLE 0.08; anterior eye row 0.33 mm long, straight; posterior eye row 0.45 mm long, procurved; MOQ length 0.19 mm, front width 0.14 mm, back width 0.24 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.03, PME-PME 0.08, PME-PLE 0.07, ALE-PLE 0.04. Sternum 0.95 mm long, 0.79 mm wide. Chelicerae 0.39 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 1.80 mm long, 1.10 mm wide. Epigastric furrow 0.56 mm from tracheal spiracle, spiracle 0.56 mm from base of spinnerets. Tibial lengths (mm) and indices: I 1.31, 17; II 1.08, 21; III 0.81, 28; IV 1.39, 18. Ventral spination: tibiae I 4-2-2*, II 3- 2-2*, III 1-2-2, IV 2-2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2-2-2. Femur III un- modified. Tibia III ventral spine 1 on retro- lateral side missing, spine 2 thickened, cone-like. Coxae unmodified. Palpus as in Figures 16, 32. Female (Pima Co., Arizona). Colora- tion as in male of A. celer. Total length 3.06 mm. Carapace 1.55 mm long, 1.26 mm wide, cephalic width 0.67 mm, clypeus height 0.06 mm. Eyes: Spider Family Anvphaenidak • Plafuick 225 diameters (mm): AME 0.05, ALE O.OS, PME 0.08, PLE O.OS; anterior eye row 0.33 mm long, .straight; posterior eye row 0.49 mm long, procurved; MOQ length 0.20 mm, front width 0.14 mm, back width 0.26 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.03, PME-PME 0.10, PME-PLE 0.07, ALE-PLE 0.04. Sternum 1.04 mm long, 0.70 mm wide. Chelicerae 0.47 mm long with teeth as in male. Abdomen 1.85 mm long, 1.08 mm wide. Epigastric furrow 0.49 mm from tracheal spiracle; spiracle 0.41 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.13, 20; II 0.92, 25; III 0.72, 33; IV 1.26, 18. Ventral spination: tibiae I, II 4-2-2^ III 1-1-2, IV 1-2-2; meta- tarsi as in male. Epigynum as in Figure 42, Internal geni- talia as in Figure 49. Natural history. Mature males and fe- males have been taken from late October through early April. Specimens have been taken from leaf litter and under rocks. DistriJmtion. Southeastern Arizona (Map 1)- Anyphaena rita new species Map 1; Figures 7, 28, 37, 44 Types. Male holotype, female paratype from Bear Canyon, Santa Catalina Moun- tains, Pima Co., Arizona, 8 December 1968 (Karl Stephan), deposited in AMNH. Male and female paratypes from Pima Co., Ari- zona, deposited in MCZ. The .specific name is a noun in apposition derived from the Santa Rita Mountains, where the species is abundant. Diagnosis. Anypliaemi rita is most closely related to A. cochise, both species having a conspicuously enlarged region of the embolus and a slightly recurved tip of the median apophysis. Males of A. rita (Fig. 28) may be distingui.shed by their smaller size and by the differences in the dorsal prong of the RTA. Females of A. cochise are unknown, l)ut the epigynum of A. rita. with its extremely broad midpiece, is quite distinctive (Fig. 37). Male (Pima Co., Arizona). Colorati(;n as in Anyphaena celer. Total length 4.10 mm. Carapace 1.94 mm long, 1.60 mm wide, cephalic width 0.67 mm, clypeus height 0.07 mm. Eyes: diameters (mm): AME 0.05, ALE 0.08, PME 0.09, PLE 0.09; anterior eye row 0.36 mm long, recurved; posterior eye row 0.53 mm long, procurved; MOQ length 0.22 mm, front width 0.15 mm, back width 0.27 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.03, PME-PME 0.09, PME-PLE 0.09, ALE-PLE 0.05. Sternum 1.13 mm long, 0.77 mm wide. Chelicerae 0.50 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 2.30 mm long, 1.26 mm wide. Epigastric furrow 0.67 mm from tracheal spiracle, spiracle 0.65 mm from base of spinnerets. Tibial lengths (mm) and indices: I 1.55, 14; II 1.39, 16; III 0.97, 23; IV 1.62, 14. Ventral spination: tibiae I 4-2-2*, II 3-2-2*, III, IV 2-2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2-2-2. Femur III un- modified. Tibia III ventral spines not thickened. Coxae III and IV with only a few short thick setae. Palpus as in Figures 7, 28. Female (Pima Co., Arizona). Coloration as in male of A. celer. Total length 5.04 mm. Carapace 2.05 mm long, 1.53 mm wide, cephalic width 1.03 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.05, ALE 6.10, PME 0.10, PLE 0.11; anterior eye row 0.41 mm long, recurved; posterior eye row 0.58 mm long, procurved; MOQ length 0.32 mm, front width 0.18 mm, back width 0.29 mm; eye interdistances (mm): AME- AME 0.07,' AME-ALE 0.03, PME-PME 0.09, PME-PLE 0.10, ALE-PLE 0.08. Sternum 1.13 mm long, 0.81 mm wide. Chelicerae 0.67 mm long with teeth as in male. Abdomen 2.75 mm long, 1.94 mm wide. Epigastric furrow 1.06 mm from tracheal 226 Bulletin Museum of Comparative Zoologij, Vol. 146, No. 4 spiracle, spiracle 0.95 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.48, 15; II 1.26, 18; III 1.03, 21; IV 1.58, 17. Ventral spination as in male except tibiae III, IV 1-2-2, meta- tarsi IV 2-1-2. Epigynum as in Figure 37, internal genitalia as in Figure 44. Natural history. Mature males have been taken from mid-October through late March, mature females from early June tlirough early February. Specimens have been taken from 4000 to 6800 feet. ( 1200- 2075 m), in oak/ grassland and under rocks. Distribution. Arizona to Chihuahua, Mexico (Map 1). Anyphaena cochise new species Map 1; Figures 13, 35 Types. Male holotype from Rustlers Park, 8600 ft. (2625 m), Chiricahua Moun- tains, Cochise Co., Arizona, 9 September 1950 (W. J. Gertsch), deposited in AMNH. Male paratype from Cochise Co., Arizona, deposited in MCZ. The specific name is a noun in apposition and refers to the type locality. Diapiosis. Anypliaena cochise is most closely related to A. vita, but the dorsal prong of the RTA is relatively longer in A. cochise (Fig. 35). Females of this species are unknown. Male (Cochise Co., Arizona). Colora- tion as in Anyphaena celer. Total length 5.44 mm. Carapace 2.52 mm long, 2.09 mm wide, cephalic width 0.88 mm, clypeus height 0.14 mm. Eyes: diameters (mm): AME 0.09, ALE 6.13, PME 0.13, PLE 0.13; anterior eye row 0.53 mm long, straight; posterior eye row 0.75 mm long, procurved; MOQ length 0.30 mm, front width 0.23 mm, back width 0.40 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.04, PME-PME 0.14, PME-PLE 0.11, ALE-PLE 0.06. Sternum 1.44 mm long, 1.08 mm wide. Chelicerae 0.75 mm long with 4 promar- ginal teeth and 7 retromarginal denticles. Abdomen 3.38 mm long, 1.94 mm wide. Epigastric fvuTow 0.92 mm from tracheal spiracle, spiracle 1.03 mm from base of spinnerets. Tibial lengths (mm) and indices: I 2.32, 12; II 2.05, 13; III 1.39, 20; IV 2.14, 14. Ventral spination: tibiae I 4-2-2*, II 2-2- 2*, III 1-2-2, IV 2-2-2; metatarsi I, II 2- 2-0, III 2-0-2, IV 2-2-2. Femur III un- modified. Tibia III ventral spine 1 on ret- rolateral side thickened slightly. All coxae with a few scattered short thick setae. Palpus as in Figures 13, 35. Female. Unknown. Natural history. Mature males have been taken in early September at 8600 feet (2625 m). Distribution. Known only from the type locality (Map 1). Anyphaena autumna new species IVIap 1; Figures 8, 29, 39, 45 Types. Male holotype, female paratype from Rustler Camp, Chiricahua Mountains, Cochise Co., Arizona, 9 September 1950 (W. J. Gertsch), deposited in AMNH. Male and female paratypes from Cochise and Graham Co., Arizona, deposited in MCZ. The specific name refers to the season of collection. Diagno.sis. Anyphaena autumna is un- likely to be confused with any other spe- cies. The long recurved hook on the RTA and the peculiar form of the tip of the median apophysis are mil ike any other species (Figs. 8, 29). The epigynum is closest to that of A. gibboides, but the mid- piece has a characteristic constriction near its midpoint ( Fig. 39 ) . Male (Cochise Co., Arizona). Colora- tion as in Anyphaena celer, though the paramedian bands on the carapace are darker and wider than in that species. Total length 5.51 mm. Carapace 2.50 mm long, 1.98 mm wide, cephalic width 1.03 mm, clypeus height 0.12 mm. Eyes: Spider Family Anyphaenidae • Platnick 227 diameters (nini): AME 0.09, ALE 0.12, PME 0.12, PLE 0.13; anterior eye row 0.55 mm long, recnrved; posterior eye row 0.75 mm long, proeun'ed; MOQ length 0.30 mm, front width 0.26 mm, back width 0.38 mm; eye interdistances (mm): AME- AME 0.08, AME-ALE 0.05, PME-PME 0.15, PME-PLE 0.11, ALE-PLE 0.06. Sternnm 1.46 mm long, 1.08 mm wide. Chelicerae 0.79 mm long with 4 promar- ginal teeth and 9 retromarginal denticles. Abdomen 3.20 mm long, 2.16 mm wide. Epigastric furrow 1.04 mm from tracheal spiracle, spiracle 1.06 mm from base of spinnerets. Tibial lengths (mm) and indices: I 2.16, 13; II 1.93, 15; III 1.39, 22; IV 2.16, 14. Ventral spination: tibiae I 2-2-2, II, III, IV 1-2-2; metatarsi I, II 2-2-0, III 2-0-2, I\^ 2-2-2. Third legs unmodified. Palpus as in Figures 8, 29. Female (Cochise Co., Arizona). Colora- tion as in male. Total length 6.41 mm. Carapace 2.34 mm long, 1.87 mm wide, cephalic width 1.12 mm, clypeus height 0.12 mm. Eyes: diameters (mm): AME 0.10, ALE 0.13, PME 0.13, PLE 0.13; anterior eye row 0.59 mm long, recurved; posterior eye row 0.70 mm long, procurved; MOQ length 0.33 mm, front width 0.27 mm, back width 0.42 mm; eye interdistances (mm): AME- AME 0.06,' AME-ALE 0.03, PME-PME 0.17, PME-PLE 0.12, ALE-PLE 0.07. Sternum 1.42 mm long, 1.08 mm wide. Chelicerae 0.99 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 3.96 mm long, 2.63 mm wide. Epigastric furrow 1.33 mm from tracheal spiracle, spiracle 1.33 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.75, 16; II 1.60, 18; III 1.10, 2.5; IV 1.89, 15. Ventral spination: tibiae I 4-4-2, II 2-4-2, III 1-1-2, IV 1-2-2; metatarsi as in male. Epigynum as in Figure 39, internal geni- talia as in Figure 45. Natural history. Mature males and fe- males have been taken in August and Sep- tember. Specimens have be(Mi taken at 8200 fec>t (2500 m). I collected a few im- matiue males (which matured in the labo- ratory) of this .species in pine litter in the Chiricahua Mountains, Arizona, where ma- ture A. mar^inalis were extremely abun- dant. Dustribution. Southeastern Arizona (Map 1)- Anyphaena gibboides new species Map 1; Figures 14, 30, 40, 46 Types. Male holotype, female paratype from City Creek Canyon, Salt Lake Co., Utah, 22 May 1943 (Wilton Ivie), depos- ited in AMNH. Male and female para- t)^es from Lake Co., Oregon, deposited in MCZ. The specific name is an arbitrary combination of letters. Diagnosis. Anyphaena gihJ)oi(Ies is a distinctive species. Males have a sharply pointed median apophysis and serrate RTA which will separate them from the other known species (Figs. 14, 30). The epigynum is closest to that of A. autumna, but lacks the constriction of the midpiece found in that species ( Fig. 40 ) . Male (Salt Lake Co., Utah). Coloration as in Anyphaena celer. Total length 3.31 mm. Carapace 1.60 mm long, 1.28 mm wide, cephalic width 0.54 mm, clypeus height 0.07 mm. Eyes: diameters (mm): AME 0.05, ALE 0.08, PME 0.08, PLE 0.08; anterior eye row 0.34 mm long, straight; posterior eye row 0.48 mm long, procurved; MOQ length 0.23 mm, front width 0.15 mm, back width 0.24 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.08, PME-PME 0.08, PME-PLE 0.07, ALE-PLE 0.05. Sternum 0.85 mm long, 0.72 mm wide. Chelicerae 0.49 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 1.94 mm long, 1.24 mm wide. Epigastric furrow 0.58 mm from tracheal spiracle, spiracle 0.59 mm from base of spinnerets. 228 BuUetm Museum of Comparative Zoology, Vol. 146, No. 4 Tibial lengths (mm) and indices: I 1.33, 17; II 1.24, 19; III 0.99, 23; IV 1.47, 16. Ventral spination: tibiae I 2-2-0, II 1-2-0, III 2-2-0, IV 2-2-2; metatarsi I, II 2-2-0, III 2-1-2, IV 2-2-2. Modifications of third leg as in A. celer save that all coxae have clumps of short thick setae. Palpus as in Figures 14, 30. Female (Salt Lake Co., Utah). Colora- tion as in male of A. celer. Total length 3.74 mm. Carapace 1.75 mm long, 1.35 mm wide, cephalic width 0.83 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.06, ALE 0.09, PME O.OS, PLE 0.08; anterior eye row 0.41 mm long, recurved; posterior eye row 0.57 mm long, procurved; MOQ length 0.24 mm, front width 0.18 mm, back width 0.28 mm; eye interdistances (mm): AME- AME 0.06, AME-ALE 0.03, PME-PME 0.12, PME-PLE 0.09, ALE-PLE 0.06. Sternum 1.19 mm long, 0.83 mm wide. Chelicerae 0.62 mm long with 4 promar- ginal teeth and 6 retromarginal denticles. Abdomen 2.36 mm long, 1..39 mm wide. Epigastric furrow 0.72 mm from tracheal spiracle, spiracle 0.70 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.39, 18; II 1.24, 20; III 0.75, 27; IV 1.39, 18. Ventral spination: tibiae I 2-2-0, II, III 1-2-0, IV 1-2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2-2-2. Epigynum as in Figure 40, internal geni- talia as in Figure 46. Natural history. Mature males and fe- males have been taken in late May and June. Habitat data is lacking. Distribution. Northern Utah west to southeastern Oregon ( Map 1 ) . Anyphaena catalina new species Map 1; Figures 15, 31, 41, 48 Types. Male holotype, female paratype from Mt. Lemon, Santa Catalina Moun- tains, Pima Co., Arizona, 13 July 1916 ( F. E. Lutz), deposited in AMNH. Male and female paratypes from Pima Co., Arizona, and Mexico, Mexico, deposited in MCZ. The specific name is a noun in apposition and refers to the type locality. Diagnosis. Anyphaena catalina is most closely related to A. arbida, though males of A. catalina may be readily distinguished by their recurved retrolateral tegular apophyses (Figs. 15, 31). Females of A. arbida are unknown; those of A. catalina may be recognized by the epigynal hood being roughly equal in size to the epigynal midpiece ( Fig. 41 ) . Male (Pima Co., Arizona). Coloration as in Anyphena celer. Total" length 3.53 mm. Carapace 1.78 mm long, 1.42 mm wide, cephalic width 0.72 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.05, ALE 0.09, PME 0.08, PLE 0.09; anterior eye row 0.40 mm long, recurved; posterior eye row 0.51 mm long, procurved; MOQ length 0.21 mm, front width 0.17 mm, back width 0.26 mm; eye interdistances (mm): AME- AME 0.07, AME-ALE 0.04, PME-PME 0.09, PME-PLE 0.08, ALE-PLE 0.04. Sternum 0.90 mm long, 0.70 mm wide. Chelicerae 0.56 mm long with 4 promar- ginal teeth and 6 retromarginal denticles. Abdomen 1.85 mm long, 0.90 mm wide. Epigastric furrow 0.61 mm from tracheal spiracle, spiracle 0.65 mm from base of spinnerets. Tibial lengths (mm) and indices: I 2.07, 8; II 1.94, 9; III 1.08, 23; IV 1.80, 10. Ven- tral spination: tibiae I 4-2-2*, II 2-2-2*, III, IV 2-2-2; metatarsi I, II 2-2-0, III 2- 0-2, IV 2-2-2. Modifications of third leg as in A. celer save that femur III lacks short thick setae and all coxae bear clumps of them. Palpus as in Figures 15, 31. Female (Pima Co., Arizona). Coloration as in male of A. celer. Total length 4.57 mm. Carapace 1.84 mm long, 1.42 mm wide, cephalic width 0.94 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.07, ALE 0.09, PME 0.09, PLE 0.09; anterior eye row 0.47 mm long, recui-ved; posterior eye row Spideu pAisriLY Anyphaemdak • Pintnick 229 0.63 mm long, prociirved; MOQ length 0.26 mm, front width 0.22 mm, back widtli 0.33 mm; eve interdi.stances (mm): AME- AME 6.0S, AME-ALE 0.04, PME-PME 0.15, PME-PLE 0.11, ALE-PLE 0.07. Sternum 1.01 mm long, 0.85 mm wide. Chelicerae 0.68 mm long with 4 promar- ginal teeth and 8 retromarginal denticle.s. Abdomen 2.74 nnn long, 1.85 mm wide. Epigastric furrow 0.86 mm from tracheal .spiracle, spiracle 0.94 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.51, 14; II 1.33, 17; III 0.94, 23; IV 1.48, 16. Ventral spination: tibiae I, II 2-2-2*, III 1-2-2, IV 2-2-2; metatarsi as in male. Epigynum as in Figure 41, internal geni- talia as in Figure 48. Natural Justory. Mature males and fe- males have been taken in July and August. Specimens have been taken at 7500 feet (2300 m) in yellow pine/ oak and douglas fir/ white fir forests. Distribution. Southeastern Arizona south to central Mexico (Map 1). Anyphaena arbida new species Map 1; Figures 17, 38 Types. Male holotype from Carr Can- yon, Huachuca Mountains, Cochise Co., Arizona, 26 August 1950 (M. A. Cazier), deposited in AMNH. Male paratype from Cochise Co., Arizona, deposited in MCZ. The specific name is an arbitrary combina- tion of letters. Diagnosis. AnypJiaena arhida is most closely related to A. catalina. Males of the former (Figs. 17, 38) lack the recurved retrolateral tegular apophysis of A. cata- lina; females of A. arbida are unknown. Male (Cochise Co., Arizona). Colora- tion as in AnypJiaena celer, except that posterior spinnerets are as in A. dixiana. Total length 6.95 mm. Carapace 3.28 mm long, 2.41 mm wide, cephalic width 1.22 mm, clypeus height 0.14 mm. Eyes: diameter (mm): AME 0.11, ALE 0.13, PME 0.13, PLE 0.15; anterior eye row 0.64 mm long, recurved; posterior eye row 0.89 mm long, procurved; MOQ lengtii 0.43 mm, front width 0.31 mm, back width 0.44 mm; eye interdistances (mm): AME- AME 0.09, AME-ALE 0.04, PME-PME 0.18, PME-PLE 0.14, ALE-PLE 0.09. Sternum 1.62 mm long, 1.33 mm wide. Chelicerae 1.30 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 3.71 mm long, 2.16 mm wide. Epigastric furrow 1.08 mm from tracheal spiracle, .spiracle 1.12 mm from base of spinnerets. Spinnerets surrounded by a clump of unusually long setae. Tibial lengths (mm) and indices: I 6.88, 5; II 3.35, 10; III 2.20, 16; IV 3.35, 10. Ventral spination: tibiae I 4-2-2*, II 3- 2-2*, III, IV 2-2-0; metatarsi I, II 2-2-0, III, IV 2-2-2. Third legs unmodified. Palpus as in Figures 17, 38. Female. Unknown. Natural history. Mature males have been collected in August. Habitat data is lack- ing- 1). Di.stribution. Cochise Co., Arizona (Map Anyphaena pectorosa Group Diagnosis. The pectorosa group is closely related to the pacifica group, but males may be distinguished by the spins on their coxae (Figs. 59-62). Females have the epigynum on a characteristic sclerotized plate (Figs. 74, 77, 79) and simple .sper- mathecae (Figs. 75, 78, 80). Description. Total length 4.5-6.5 mm. Carapace longer than wide, narrowed in front to less than half its maximum width in males, to slightly more than half its maximum width in females. Clypeus height more than 1.5 times the diameter of an an- terior median eye. Posterior median, pos- terior lateral and anterior lateral eyes sub- equal in size, almost twice the diameter of anterior medians. Procurved posterior eye row longer than slightly recur\xKl anterior row. Median ocular (juadrangle almost 230 Bulletin Museum of Comparative Zoolofi.ij, Vol. 146, No. 4 twice as wide in back as in front. Anterior median eyes separated l:)y sliglitly less than their diameter, sHghtly closer to anterior laterals than to each other. Posterior me- dians separated by slightly more than their diameter, slightly closer to posterior lat- erals. Anterior laterals separated by their radins from posterior laterals. Sternum longer than wide, with a low hirsute knob behind its middle in some males. Chelic- erae with 4 promarginal teeth and 7-9 retromarginal denticles. Abdomen longer than wide, tiacheal spiracle midway be- tween epigastric furrow and base of spin- nerets. Leg formula 1423. Metatarsi I and 11 with two pairs of ventral spines. Males with coxae II bearing round knobs, coxae III and IV bearing spurs. Palpus with an elongated median apophysis, en- larged conductor and inconspicuous embo- lus. Retrolateral tibial apophysis short. Epigynum on a sclerotized plate, without a hood. Two simple spermathecae. Variation. The species in this group show little intraspecific variation, individ- ual or geographical, in size, structure or coloration. Key to Species la. Coxae III of males with posterior spur bifid (Fiffs. 59, 61, 62); sternum of males with a low hirsute knob behind middle; sclerotized epigynal plate wider posteriorly than an- teriorly (Figs. 74, 79) 2 lb. Coxae III of males with posterior spur not bifid ( Fig. 60 ) ; sternum of males without a low hirsute knob behind middle; sclerotized epigynal plate wider anteriorly than pos- teriorly (Fig. 77) fratema 2a. Distal tip of palpal median apophysis bent sharply towards cymbium (Figs. 55, 58); sclerotized epigynal plate with pronounced posterolateral corners ( Fig. 74 ) 3 2b. Distal tip of palpal median apophysis not bent sharply towards cymbium ( Fig. 57 ) ; sclerotized epigynal plate without pro- nounced posterolateral corners (Fig. 79) alaclma 3a. Distal tip of palpal median apophysis meet- ing the recessed, dorsal branch of tlie apoph- ysis ( Fig. 55 ) ; sclerotized epigynal plate with pronounced posterolateral comers ( Fig. 74 ) pectorosa 3b. Distal tip of palpal median apophysis not meeting the recessed, dorsal branch of the apophysis (Fig. 58); females unknown __7flc?:a Anyphaena pectorosa L. Koch Map 2; Figures 51, 55, 59, 74, 75 Anyphaena pectorosa L. Koch, 1866, Arachn. Fam. Drass., 198, pi. 8, figs. 131, 132 { $). Male holotype from Baltimore, Maryland, in BMNH, examined. Bryant, 1931, Psyche, 38: 110, pi. 6, fig. 5, $ . Chickering, 1939, Pap. Michigan Acad. Sci., 24: 51, figs. 5-8, $,9. Comstock, 1940, Spider Book, rev. ed., p. 577, fig. 636, 9 . Kaston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 408, figs. 1453, 1477-1480, $, 9. Roewer, 1954, Katalog der Araneae, 2: 529. Bonnet, 1955, Bibliographia Araneorum, 2: 346. Aniiphaena calcarata Emerton, 1890, Trans. Con- necticut Acad. Sci., 8: 187, pi. 6, figs. 3-3d ( $, 9 ). Male holotype, female allotype from West Haven, Connecticut, in MCZ, examined. Emer- ton, 1902, Common Spiders, p. 12, figs. 42, 43, $, 9. Gaijenna calcarata, Banks, 1910, Bull. U.S. Nat. Mus., 72: 13. Gaijenna pectorosa, Comstock, 1912, Spider Book, p. 563 (in part), fig. 636, 9 (not fig. 637). Diagnosis. Anyphaena pectorosa is closest to A. alachua, but may readily be distin- guished from it by the highly curved me- dian apophysis of males (Fig. 55) and the pronounced posterolateral corners of the sclerotized epigvnal plate of females (Fig. 74). Male (Fairfax Co., Virginia). Total length 5.40 mm. Carapace 2.43 mm long, 1.98 mm wide, cephalic width 0.88 mm, clypeus height 0.11 mm, yellow with thin dark border and two dark paramedian longitudinal bands. Eyes: diameters (mm): Plate 5 Figures 51--54. Left palpi, ventral view. Figures 55-58. Left palpi, retrolateral view. Figures 59-62. ventral view. 51, 55, 59. Anyptiaena pectorosa L. Koch. 52, 56, 60. Anyphaena fraterna (Banks). Anyphaena alachua new species. 54, 58, 62. Anyphaena lacka new species. Male coxae, 53, 57, 61. Spider I'^aafily Anyphaenidak • Plaliuck 231 232 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 AME 0.06, ALE 0.11, PME 0.11, PLE 0.12; anterior e\e row 0.48 mm long, slightly re- cur^•ed; posterior eye row 0.65 mm long, procurved; MOQ length 0.28 mm, front width 0.20 mm, back width 0.35 mm; eye interdistances (mm): AME-AME 0.07, AME-ALE 0.04, PME-PME 0.14, PME- PLE 0.13, ALE-PLE 0.05. Sternum 1.35 mm long, 1.01 mm wide, pale yellow with translucent border, dark- ened extensions to coxae and a low hirsute knob behind middle. Chelicerae 0.73 mm long with 4 promarginal teeth and 7 retro- marginal denticles, pale yellow with boss outlined in gray. Labium and endites yel- low, darkest proximally. Endites slightly invaginated at middle. Abdomen 3.15 mm long, 1.67 mm wide, pale white with transverse rows of dark markings, venter pale. Epigastric furrow 1.01 mm from tracheal spiracle, spiracle 1.06 mm from base of spinnerets. Legs pale yellow with distal segments darkest. Tibial lengths (mm) and indices: I 3.10, 7; II 2.52, 9; III 1.82, 16; IV 2.56, 10. Ventral spination: tibiae I 2-2-1, II-IV 2- 2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2- 2-2. Coxae II, III and IV modified as in Figure 59. Palpus as in Figures 51, 55. Female (Fairfax Co., Virginia). Colora- tion as in male. Total length 5.44 mm. Carapace 2.41 mm long, 1.91 mm wide, cephalic width 0.97 mm, clypeus height 0.08 mm. Eyes: diameters (mm): AME 0.07, ALE 0.12, PME 0.11, PLE 0.12; anterior eye row 0.52 mm long, recurved; posterior eye row 0.71 mm long, procurved; MOQ length 0.33 mm, front width 0.20 mm, back width 0.37 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.04, PME-PME 0.15, PME-PLE 0.10, ALE-PLE 0.07. Sternum 1.31 mm long, 1.06 mm wide, without hirsute knob. Chelicerae 0.72 mm long with 4 promarginal teeth and 8 retro- marginal denticles. Abdomen 3.10 mm long, 1.76 mm wide. Epigastric furrow 0.70 mm from tracheal spiracle, spiracle 1.22 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 2.41, 11; II 2.05, 13; III 1.44, 19; IV 2.20, 12. Ventral spination: tibiae I, II 2-2-0, III, IV 1-2-1; metatarsi I, II 2-2-0, III, IV 2-2-2. Epigynum as in Figure 74, internal geni- talia as in Figure 75. Natural history. Mature males have been taken from mid-April through early Sep- tember, mature females from mid-April through mid-August. Specimens have been taken by sweeping foliage, in Malaise and pitfall ti^ips, and under rocks. Egg cases taken with females contained 65-95 eggs. Distribution. New England west to Michigan, south to western Florida and eastern Texas ( Map 2 ) . Anyphaena atachua new species Map 2; Figures 53, 57, 61, 79, 80 Types. Male holotype, female paratype from west of Gainesville, Alachua Co., Florida, 18 April 1938 (Willis J. Certsch), deposited in AMNH. Male and female paratypes from Alachua Co., Florida, de- posited in MCZ. The specific name is a noun in apposition and refers to the type locality. Diagnosis. Anyphaena alachua is closest to A. pectorosa but the median apophysis is not highly curved (Fig. 57) and the epigynal plate lacks pronounced postero- lateral corners (Fig. 79). Male (Alachua Co., Florida). Colora- tion as in Anyphaena pectorosa. Total length 4.90 mm. Carapace 2.41 mm long, 2.01 mm wide, cephalic width 0.79 mm, clypeus height 0.13 mm. Eyes: diameters (mm): AME 0.07, ALE 0.12, PME 0.12, PLE 0.13; anterior eye row 0.51 mm long, slightly recurved; posterior eye row 0.70 mm long, procurved; MOQ length 0.30 mm, front width 0.22 mm, back width 0.36 mm; eye interdistances (mm): AME-AME 0.07, AME-ALE 0.04, PME- PME 0.12, PME-PLE 0.11, ALE-PLE 0.06. Spider Family Anvi'haemdae • Plalnick 233 Sternum 1.26 mm long, 1.01 mm wide, with low hirsute knob Ix^hind middle. Chelicerae 0.76 mm long with 4 promar- ginal teeth r.nd 9 retromarginal denticle.s. Abdomen 2.48 mm long, 1.48 mm wide. Epigastric furrow 0.76 mm from tracheal spiracle, spiracle 0.8.3 mm from base of spinnerc>ts. Tibial lengtlis (nun) and indices: I 2.77, 10; II 2.27,^11; III 1.44, 22; IV 1.94. 14. \Vntral spination: tibiae I, II 2-2-0, III 1-2-2, IV 2-2-2; metatarsi I, II 2-2-0, III 2-0-2, IV 2-2-2. Coxae II, III and I\' modified as in Figure 61. Palpus as in Figures 53, 57. Female (Alachua Co., Florida). Colora- tion as in male of A. pectorosa. Total length 6.17 mm. Carapace 2.45 mm long, 1.80 mm wide, cephalic width 0.94 mm, clypeus height 0.12 mm. Eyes: diameters (mm): AME 0.08, ALE 0.13, PME 0.12, PLE 0.13; anterior eye row 0.57 mm long, slightly recurved; posterior eye row 0.73 mm long, procui-ved; MOQ IcMigth 0.30 mm, front width 0.22 mm, back width 0.40 mm; eye interdistances (mm): AME-AME 0.07, AME-ALE 0.04, PME- PME 0.15, PME-PLE 0.11, ALE-PLE 0.07. Sternum 1.35 mm long, 1.08 mm wide, without hirsute knob. Clielicerae 0.84 mm long with teeth as in male. Abdomen 3.53 mm long, 2.02 mm wide. Epigastric furrow 1.10 mm from tracheal spiracle, spiracle 1.21 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 2.30, 13; II 1.91, 14; III 1.31, 22; IV 2.09, 13. Ventral spination as in male save metatarsi III 2-2-2. Epigynum as in Figure 79, internal geni- talia as in Figure 80. Natural history. Mature males have been taken in late April and early May, mature females from late March through mid-May, by sweeping. Distribution. Known only from Florida (Map 2). Anyphaena lacka new species Map 2; Figures 54, 58, 62 Type. Male liolotxpe from Lake Corpus Christi State Park, southwest of Mathis, San Patricio Co., Texas, 28 Jvme 1962 (J. A. Beatty), deposited in MCZ. The specific name is an arbitrary combination of letters. Dia I ------v*- i£ Anyphoena fraterna .«\ -^> f* ■' 1 c "^ 1 ijiy. \ '( VM • ^ 1 r^"^- *' 1 1 )• ;' ''*~ • .-••\ • _ (• '~~^-r- ; • 1 V ;'• _ 1 ~'^ ";•:• z--- ..^ • • - i r^^ 1 Anyphaenc pacificQ / A -^w Tn P ^■-^ .__..__ Anyphoena colifornicQ - 1 \\v 1 1 "~ r ) Anyphoena locko s , 1 \\ i ,'---,' ; .' t>'- ^. ^ ■' ■• "> / \ \ 1 "\ : 1 Anyphoena aperta ' \ 1 -^ ^ .'- Map 2. Distributions of Anyphaena alachua, A. aperta, A. californica, A. fraterna, A. gertschi, A. lacka, A. pacif- ica and A. pectorosa. males of Anyphaena gertschi have rounded knobs on the coxae. Females lack the sclerotized epigynal plates found in the pectorosa group, but have a lightly sclero- tized atrivun-like area posteromedially (Figs. 66, 67, 72) and long, sometimes coiling, ducts (Figs. 68, 73, 76). Description. Total length 4-6 mm. Cara- pace longer than wide, narrowed in front by at least one-third of its maximum width, often by more than half. Clypeus height roughly equal to anterior median eye diam- eter. All eyes subequal in size. Procurved posterior eye row longer than slightly re- curved anterior eye row. Median ocular quadrangle longer than wide in front, wider in back than long. Anterior median eyes separated by less than their diameter, much closer to anterior laterals than to each other. Posterior medians separated by more than their diameter, much closer to posterior laterals. Anterior laterals sepa- rated by slightly more than their radius from posterior laterals. Sternum longer than wide, without a hirsute knob. Chelic- erae with 3 promarginal teeth and 6-9 retromarginal denticles. Abdomen longer than wide, tracheal spiracle midway be- tween epigastric furrow and base of spin- nerets. Leg formula 1423. Metatarsi I and II with two pairs of ventral spines. Males with legs unmodified. (A. pacifica and A. californica) or with coxae bearing round knobs and femora II and III bearing patches of short stiff setae ventrally (A. gertschi). Palpus with an elongated me- dian apophysis, enlarged conductor and inconspicuous embolus. Retrolateral tibial apophysis short. Epigyiuim not on a scle- rotized plate, without a hood, with a more or less pronoimced atrium-like lightly sclerotized area posteromedially. Internal genitalia with long ducts that coil in some species. Variation. Two species in this group, A. pacifica and A. californica, show a great deal of xariation in genitalic structure. In both species the shape oi the tip of the pal- 236 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 pal median apophysis and the coihng of the epigynal ducts are strikingly variable, and it was initially thought that many species were involved. Three sources of evidence, however, have indicated other- wise. First, many females are found in which the ducts on one side of the epigy- num coil differently from those on the other side. Secondly, when many speci- mens are taken together at one locality on a single da}", several variants are often found. Finally, the retrolateral tibial apophysis, which usually provides excellent diagnostic characters in anyphaenids, is stable within the species as they are de- fined here. Until such time as biological evidence on the breeding habits of these spiders can be obtained, it seems best to consider both A. pacifica and A. califorjuco as widespread, variable species. Key to Species la. Retrolateral tibial apophysis (RTA) without a dorsal process (Fig. 69). Median apoph- ysis with a deep invagination below tip giving the tip a chelate appearance (Fig. 65). Epigyninii with large wing-shaped paramedian flaps ( Fig. 72 ) gertschi lb. Retrolateral tibial apophysis (RTA) with a dorsal process (Figs. 70, 71). Median apophysis without a deep invagination be- low tip (Figs. 63, 64). Epigynum with- out large wing-shaped paramedian flaps (Figs. 66, 67) 2 2a. Dorsal process of RTA short, located dis- tally (Fig. 70). Median apophysis narrow- ing gradually towards tip (Fig. 63). In- ternal ducts with many coils (Fig. 68) — _ — pacifica 2b. Dorsal process of RTA long, located prox- inially (Fig. 71). Median apophysis nar- rowing abruptly towards tip (Fig. 64). In- ternal ducts without many coils (Fig. 73) — californica Anyphaena pacifica (Banks) Map 2; Figures 63, 66, 68, 70 Gaijcnna pacifica Banks, 1896, Trans. Amer. Ent. Soc, 23: 63. Female holot>'pe from Olympia, Washington, in MCZ, examined. Anyphaena pacifica, Simon, 1897, Hist. Natur. Araign., 2: 96. Bryant, 1931, Psyche, 38: 115, pi. 8, fig. 36, ?. Levi and Levi, 1951, Zoo- logica (New York), 36: 228, Tig. 25, $. Roe- wer, 1954, Katalog der Araneae, 2: 529. Bon- net, 1955, Bibliographia Araneorum, 2: 346. Anyphaena mundella Chamberlin, 1920, Pomona Coll. J. Ent. Zool., 12: 12, pi. 5, fig. 3 ( 9 , not $, = Aysha incursa) . Female holotype from Claremont, California, in MCZ, examined. Bryant, 1931, Psyche, 38: 120 (sub Aysha de- cepta [sic] ). Roewer, 1954, Katalog der Araneae 2: 534 (sub Aysha decepta [sic]). Bonnet, 1955, Bibliographia Araneorum, 2: 836 (sub Aysha decepta [sic]). NEW SYNONYMY. Anyphaena intermontana Chamberlin, 1920, Canad. Ent., 52: 200, fig. 22-6 ( $ ). Female holotype from Mill Creek, Salt Lake Co., Utali, in MCZ, examined. Bryant, 1931, Psyche, 38: 114 (sub Anyphaena californica [sic]). Roe- wer, 1954, Katalog der Araneae, 2: 528 (sub Anyphaena californica [sic]). Bonnet, 1955, Bibliographia Araneorum, 2: 343 (sub Any- phaena californica [sic]). NEW SYNONYMY. Gayenna saniuana Chamberlin and Gertsch, 1928, Proc. Biol. Soc. Wash., 41: 185. Male holotype from Verdure, San Juan Co., Utah, in AMNH, examined. Roewer, 1954, Katalog der Araneae, 2: 540. NEW SYNONYMY. Anyphaena saniuana, Bryant, 1931, Psyche, 38: 107. Bonnet, 1955, Bibliographia Araneorum, 2: 347. Anyphaena pomona Chamberlin and Ivie, 1941, Bull. Univ. Utah, Biol., 6: 23, pi. 2, fig. 16 ( 9 ). Female holotype from Mill Creek, Te- hama Co., California, in AMNH, examined. Roewer, 1954, Katalog der Araneae, 2: 529. NEW SYNONYMY. Gayenna jollensis Schenkel, 1950, Verb. Naturf. Ges. Basel, 61: 77, fig. 27 ( 9 ). Female holo- type from La Jolla, California, in Naturhistor- isches Museum, Basel, examined. Roewer, 1954, Katalog der Araneae, 2: 540. NEW SYN- ONYMY. Plate 6 Figures 63-65. Left palpi, ventral view. Figures 69-71. Left palpal tibiae, retrolateral view. Figures 66, 67, 72, 74, 77, 79. Epigyna, ventral view. Figures 68, 73, 75, 76, 78, 80. Internal genitalia, dorsal view. 63, 66, 68, 70. Anyptiaena pacifica {Banks). 64,67,71,73. Anyptiaena californica {Banks). 65,69,72,76. Anyphaena gertschi new species. 74, 75. Anyphaena pectorosa L. Koch. 77, 78. Anyphaena fraterna (Banks). 79, 80. Anyphaena alachua new species. Spider Family Anyphaenidae • Plalnick 237 68 ^V ?) 78 •^ £S^' 238 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 Diagnosis. Amjphaena pacifica is closest to A. californica, but males may be distin- guished by the short, distal, dorsal process of the retrolateral tibial apophysis (Fig. 70) and the gradually narrowing tip of the median apophysis (Fig. 63), while females have distinctive highly coiled internal ducts (Fig. 68). Variation in this species is discussed above. Male (El Dorado Co., California). Total length 5.18 mm. Carapace 2.34 mm long, 1.94 mm wide, cephalic width 0.86 mm, clypeus height 0.12 mm, pale orange with thin dark border and two dark paramedian longitudinal bands. Eyes: diameters (mm): AME 0.09, ALE 0.12, PME 0.10, PLE 0.11; anterior eye row 0.51 mm long, slightly procurved; posterior eye row 0.69 mm long, procurved; MOQ length 0.28 mm, front width 0.24 mm, back width 0.34 mm; eye interdistances (mm): AME-AME 0.07, AME-ALE 0.03, PME-PME 0.14, PME- PLE 0.10, ALE-PLE 0.05. Sternum 1.49 mm long, 1.04 mm wide, pale orange with darker border. Chelicerae 0.67 mm long with 3 promarginal teeth and 8 retromarginal denticles, dark orange- brown proximally, pale orange distally, with boss outlined in gray. Labium and endites orange, darkest proximally. En- dites slightly invaginated at middle. Abdomen 2.81 mm long, 1.69 mm wide, reddish-brown throughout. Epigastric fur- row 0.85 mm from tracheal spiracle, spira- cle 0.92 mm from base of spinnerets. Legs pale orange, unmodified. Tibial lengths (mm) and indices: I 2.11, 12; II 1.87, 13; III 1.44, 20; IV 2.07, 15. Ventral spination: tibiae I, II 2-2-0, III 1-2-2, IV 2-2-2; metatarsi I, II 2-2-0, III, IV 2-2-2. Palpus as in Figures 63, 70. Female (Mono Co., California). Color- ation as in male. Total length 5.39 mm. Carapace 2.34 mm long, 1.62 mm wide, cephalic width 0.94 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.10, ALE 0.12, PME 0.11, PLE 0.11; anterior eye row 0.51 mm long, slightly recurved; posterior eye row 0.73 mm long, procurved; MOQ length 0.29 mm, front width 0.25 mm, back width 0.36 mm; eye interdistances (mm): AME- AME 0.06, AME-ALE 0.03, PME-PME 0.15, PME-PLE 0.10, ALE-PLE 0.07. Sternum 1.44 mm long, 1.01 mm wide. Chelicerae 0.71 mm long with teeth as in male. Abdomen 3.02 mm long, 1.69 mm wide. Epigastric furrow 0.81 mm from tracheal spiracle, spiracle 0.86 mm from base of spinerets. Tibial lengths (mm) and indices: I 1.84, 15; II 1.71, 15; III 1.39, 19; IV 2.07, 13. Ventral spination as in male save tibiae III 1-1-2 and IV 1-2-2. Epigynum as in Figure 66, internal geni- talia as in Figure 68. Natural history. Mature males have been taken from late February through late July, mature females year round. Speci- mens have been taken in montane forests, in pitfall traps, under rocks and commonly in houses. Distribution. Western North America from British Columbia south to California, Ai-izona and New Mexico (Map 2). Anyphaena californica (Banks) Map 2; Figures 64, 67, 71, 73 Gaijenna californica Banks, 1904, Proc. California Acad. Sci., 3: 338, pi. 38, fig. 2(9). Female holotype from Palo Alto, California, in MCZ, examined. Amiphaena mens Chamberlin, 1920, Pomona Coll. J. Ent. Zoo!., 12: 11, pi. 5, fig. \ {$). Male holotype from Claremont, California, in MCZ, examined. Bryant, 1931, Psyche, 38: 113. Roewer, 1954, Katalog der Araneae, 2: 529. Bonnet, 1955, Bibliographia Araneorum, 2: 347. NEW SYNONYMY. Anyphaena californica, Bryant, 1931, Psyche, 38: 114. Roewer, 1954, Katalog der Araneae, 2: 528. Bonnet, 1955, Bibliographia Araneorum, 2: 343. Diagnosis. Amjphaena californica is most closely related to A. pacifica, but males have a long, proximal, dorsal process on the retrolateral tibial apophysis (Fig. 71) and an abruptly narrowed tip of the me- Spider Family Anyphaenidae • Phifnick 239 diaii apopliysis (Fig. 64), while the inter- nal dncts of the female are not highly coiled (Fig. 73). Variation in this species is discnssed above. Male (San Diego Co., California). Col- oration as in AnypJiaena pacifica except that the abdomen is pale white with trans- verse rows of dark markings. Total length 4.68 mm. Carapace 2.21 mm long, 1.78 mm wide, cephalic width 0.68 mm, clypeus height 0.07 mm. Eyes: diameters (mm): AME 0.07, ALE 6.09, PME 0.10, PLE 0.11; anterior eye row 0.43 mm long, recm-ved; posterior eye row 0.59 mm long, procurved; MOQ length 0.30 mm, front width 0.20 mm, back width 0.32 mm; eye interdistances (mm): AME- AME 0.06, AME-ALE 0.04, PME-PME 0.13, PME-PLE 0.11, ALE-PLE 0.07. Sternum 1.31 mm long, 0.90 mm wide. Chelicerae 0.60 mm long with 3 promar- ginal teeth and 8 retromarginal denticles. Abdomen 2.97 mm long, 1.34 mm wide. Epigastric furrow 0.79 mm from tracheal spiracle, spiracle 0.85 mm from base of spinnerets. Tibial lengths (mm) and indices: I 3.28, 5; II 3.20, 7; III 2.27, 8; IV 2.93, 7. Ventral spination: tibiae I 2-2-0, II 2-2-2, III 1- 1-2, IV 1-2-2; metatarsi I, II 2-2-0, III, I\' 2-2-2. Palpus as in Figures 64, 71. Female (Humboldt Co., CaHfornia). Coloration as in male. Total length 5.98 mm. Carapace 2.56 mm long, 1.91 mm wide, cephalic width 1.03 mm, clypeus height 0.08 mm. Eyes: diameters (mm): AME 0.09, ALE 0.11, PME 0.12, PLE 0.12; anterior eye row 0.52 mm long, recurved; posterior eye row 0.69 mm long, procurved; MOQ length 0.35 mm, front width 0.25 mm, back width 0.37 mm; eye interdistances (mm): AME- AME 0.07, AME-ALE 0.04, PME-PME 0.13, PME-PLE 0.10, ALE-PLE 0.06. Sternum 1.44 mm long, 1.08 mm wide. Chelicerae 0.86 mm long with 3 promar- ginal teeth and 9 retromarginal denticles. Abdomen 3.64 mm long, 2.43 mm wide. Epigastric furrow 1.15 mm from tracheal spiracle, spiracle 1.33 mm from base of spinnerets. Tibial lengths (mm) and indices: I 2.16, 12; II 1.87, 13; III 1.30, 19; IV 2.06, 14. Ventral spination as in male except tibiae II 1-2-0. Epigynum as in Figure 67, internal geni- talia as in Figure 73. Natural Jii.story. Mature males have been taken from early March through mid-July, mature females from mid-March through mid-November. Specimens have Ix^en taken in redwood forests, on citrus trees and in houses. Distribution. Oregon and California (Map 2). Anyphaena gertschi new species Map 2; Figures 65, 69, 72, 76 Types. Male holotype, female paratype from Bluff, San Juan Co., Utah, 11 May 1933 (Wilton Ivie), deposited in AMNH. Male and female paratypes from Emery Co., Utah, deposited in MCZ. The specific name is a patronym in honor of Willis J. Gertsch, who first recognized the species as new. Diagnosis. Anyphaena gertschi is a dis- tinctive species easily recognized by the chelate appearance of the tip of the me- dian apophysis of males (Fig. 65) and the large wing-shaped paramedian flaps on the female epigynum (Fig. 72). Male (Emery Co., Utah). Coloration as in Anyphaena pacifica except that cara- pace has paramedian bands only vaguely indicated and abdomen is pale yellow throughout. Total length 4.00 mm. Carapace 1.85 mm long, 1.42 mm wide, cephalic width 0.92 mm, clypeus height 0.14 mm. Eyes: diameters (mm): AME 0.09, ALE 6.09, PME 0.09, PLE 0.09; anterior eye row 0.45 mm long, slightly recurv'ed; pos- terior eye row 0.59 mm long, procurved; MOQ length 0.26 mm, front width 0.22 mm, back width 0.32 mm; e\e interdis- 240 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 tances (mm): AME-AME 0.04, AME- ALE 0.03, PME-PME 0.14, PME-PLE 0.08, ALE-PLE 0.04. Sternum 1.12 mm long, 0.85 mm wide. Chelicerae 0.65 mm long with 3 promar- ginal teeth and 6 retromarginal denticles. Abdomen 2.11 mm long, 1.31 mm wide. Epigasti'ic furrow 0.67 mm from tracheal spiracle, spiracle 0.76 mm from base of spinnerets. All coxae with round knobs ventrally. Femora II and III with patches of short, thick setae ventrally. Tibial lengths (mm) and indices: I 2.00, 9; II 1.69, 13; III 1.30, 17; IV 1.87, 12. Ventral spination: tibiae I 2-2-0, II 1-2-0, III, IV 1-2-2; metatarsi I, II 2-2-0, III, IV 2-2-2. Palpus as in Figures 65, 69. Female (San Diego Co., Cahfornia). Coloration as in male. Total length 5.04 mm. Carapace 2.25 mm long, 1.76 mm wide, cephalic width 0.95 mm, clypeus height 0.12 mm. Eyes: diam- eters (mm): AME 0.10, ALE 0.13, PME 0.10, PLE 0.13; anterior eye row 0.51 mm long, straight; posterior eye row 0.68 mm long, procurved; MOQ length 0.28 mm, front width 0.26 mm, back width 0.36 mm; eye interdistances (mm): AME-AME 0.06, AME-ALE 0.03, PME-PME 0.16, PME-PLE 0.08, ALE-PLE 0.05. Sternum 1.28 mm long, 0.90 mm wide. Chelicerae 0.70 mm long with teeth as in male. Abdomen 3.10 mm long, 2.02 mm wide. Epigastric furrow 0.77 mm from tracheal spiracle, spiracle 1.03 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 1.62, 14; II 1.49, 15; III 1.17, 21; IV 1.69, 14. Ventral spination as in male save tibiae III 1-1-0. Epigynum as in Figure 72, internal geni- talia as in Figure 76. Natural history. Mature males have been taken from late April through late June, mature females from mid-May through late September. Nothing is known of the habits of this species. Distribution. Southern Utah south to southern California and Arizona (Map 2). Anyphaena accentuata Group Diagnosis. Members of this group can be immediately differentiated from the other nearctic Anyphaena by the presence of only one pair of ventral spines on meta- tarsi I and II. Only one species occurs in America north of Mexico. Description. Total length 4-6 mm. Cara- pace longer than wide, narrowed in front to less than half its maximum width in males, to slightly more than half in females. Clypeus height roughly equal to anterior median eye diameter. Median eyes smaller than laterals. Procurved posterior eye row longer than recurved anterior row. Me- dian ocular quadrangle longer than wide in front, wider in back than long. Anterior median eyes separated by less than their diameter, closer to anterior laterals. Pos- terior medians separated by 1.5 times their diameter, closer to posterior laterals. An- terior laterals separated by their radius from posterior laterals. Sternum longer than wide, unmodified. Chelicerae with 3 promarginal teeth and 5-7 retrolateral den- ticles. Abdomen longer than wide, tracheal spiracle midway between epigastric furrow and base of spinnerets. Leg formula 1423, legs unmodified. Metatarsi I and II with one pair of ventral spines. Palpus with short median apophysis, short conductor and conspicuous embolus. Cymbial groove compressed to retrolateral side of cymbium. Epigynum with hood. Internal genitalia with anterior membranous dorsal cover. Variation. No significant variation was detected in Anyphaena aperta. Anyphaena accentuata (Walckenaer) Figure 134 Aranea accentuata Walckenaer, 1802, Faun. Paris, 2: 226. Type lost, presumed destroyed. Anyphaena accentuata, Roewer, 1954, Katalog der Araneae, 2: 522. Bonnet, 1955, Bibliographia Araneoriini, 2: 338. Spider Family Axypiiaenidak • Plafnick 241 A drawing of tlic palpus of tliis Enro- pc^an spider, type species ol tlie genus A/ij/- pJiaena, is included for pinposes of com- parison to A. aperta. Confusion exists between AnypJiaena accentuatii, A. ohscura (Sundex'all) and A. sabina L. Koch, and the female is therefore not illustrated and no description is gi\^en. The male illus- trated is from England. Anyphaena aperta (Banks) Map 2; Figures 135-137 Gaijenna aperta Banks, 1921, Pioc. California Acad. Sci., 11: 100, fig. ,3 ( 9 ). Female holo- t\'pe from OKinpia, Washington, in MCZ, ex- amined. Ani/pliacna aperta, Bryant, 1931, Psyche, 38: 114, pi. 8, fig. 35, 9 . Fox, 1938, Iowa State Coll. J. Sci., 12: 238, pi. 1, fig. 6, $. Roewer, 1954, Katalog der Araneae 2: 528. Bonnet, 1955, Bibliographia Araneornm, 2: 342. Didiinosis. In addition to the diagnostic character of the species group, Amjphaemi aperta can readily be distinguished from all other North American anyphaenids by the sharply pointed median apophysis of males (Fig. 135) and the membranous dor- sal cover of the internal genitalia of females (Fig. 137). Although the distribution indi- cates that this might be an inti-oduced species, no specimens or described species from the Palearctic or Oriental regions re- semble Anyphaena aperta. Male (Yamhill Co., Oregon). Total length 4.32 mm. Carapace 1.98 mm long, 1.63 mm wide; cephalic width 0.74 mm, clypeus height O.OS mm, light orange- brown, darker towards sides, with two dark paramedian longitudinal bands. Eyes: diameters (mm): AME 0.07, ALE 0.11, PME 0.09, PLE 0.11; anterior eye row 0.44 mm long, recurved; posterior eye row 0.62 mm long, procurved; MOQ length 0.26 mm, front width 0.20 mm, back width 0.32 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.03, PME-PME 0.14, PME-PLE 0.10, ALE-PLE 0.06. Sternum 1.04 mm long, 0.89 mm wide, pale orange \\'ith translucent border and darkened extensions to coxae. Chelicerae 0.55 mm long with 3 promarginal teeth and 5 retromarginal denticles, orange-brown with boss outlined in gray. Labium and endites pale orange, darkest proximally. Endites not invaginated. Abdomen 2.52 mm long, 1.51 mm wide, pale white with transx'crse rows of dark markings, venter pale with a clump of thick elongate setae posteriorly. Epigastric furrow 0.86 mm from tracheal spiracle, spiracle 0.74 mm from base of spinnerets. Legs pale yellow, unmodified. Tibial lengths (mm) and indices: I 1.87, 12; II 1.70, 13; III 1.27, 18; IV 1.73, 14. Ventral spination: tibiae I, II 2-2-2, III 1-2-2, IV 2-2-2; metatarsi I, II 2-0-0, III 2-0-2, IV 2-2-2. Palpus as in Figure 135. Female (Curry Co., Oregon). Colora- tion as in male. Total length 5.83 mm. Carapace 2.65 mm long, 2.05 mm wide, cephalic width 1.17 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.10, ALE 0.12, PME 0.12, PLE 0.13; anterior eye row 0.61 mm long, slightly recurved; posterior eye row 0.87 mm long, procurved; MOQ length 0.35 mm, front width 0.30 mm, back width 0.44 mm; eye interdistances (mm): AME-AME 0.09, AME-ALE 0.05, PME- PME 0.19, PME-PLE 0.14, ALE-PLE 0.07. Sternum 1.46 mm long, 1.04 mm wide. Chelicerae 0.80 mm long with 3 promar- ginal teeth and 7 retromarginal denticles. Abdomen 4.00 mm long, 2.60 mm wide, without thick setae ventrally. Epigastric furrow 0.81 mm from tracheal spiracle, spiracle 1.03 mm from base of spinnerets. Tibial lengths (mm) and indices: I 1.87, 16; II 1.77, 16; III 1.31, 22; IV 1.87, 17. Ventral spination as in male except tibiae I 2-2-0 and IV 1-2-2. Epigynum as in Figure 136, internal genitalia as in Figure 137. Natural history. Mature males have been taken from late Marcli through early Sep- tember, mature females from ],\tc March 242 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 through early November. Specimens have been taken from redwoods and red cedars. Distribution. Pacific coast from British Coknnbia south to southern Cahfornia (Map 2). Wulfila O. P.-Cambridge WuIfiJa O. P.-Cambridge, 1895, Biologia Central! Americana, Aran., 1: 158. Type species Wulfila pallidtis O. P.-Cambridge, 1895, designated by Simon, 1897, Hist. Natur. Araign., 2: 103. Cragits O. P.-Cambridge, 1896, Biologia Centrali Americana, Aran., 1: 215. Type species by mono- typy Cragiis pallidus O. P.-Cambridge, 1896. NEW SYNONYMY. Anyphaenella Bryant, 1931, Psyche, 38: 115. Type species by original designation Clubiona salta- hunda Hentz, 1847. NEW SYNONYMY. Diagnosis. Wulfila may be easily recog- nized by their long, thin, pale white legs. Leg I in particular is greatly elongated, with its tibial index usually 5 or less. Pal- pal structure indicates that this genus is closely related to Amjphaena. There are probably more than fifty species in this genus; most occur in Central America and the West Indies. Description. Total length 2.5-4.5 mm. Carapace longer than wide, narrowed in front to from one-half to two-thirds its maximum width. Clypeus height greater than anterior median eye diameter. Pos- terior median, posterior lateral and anterior lateral eyes subequal in size, somewhat larger than anterior medians. Procurved posterior eye row longer than straight an- terior row. Median ocular quadrangle twice as wide in back as in front. Anterior median eyes separated by less than their diameter, by roughly their diameter from anterior laterals. Posterior medians sepa- rated by almost twice their diameter, by their diameter from posterior laterals. An- terior laterals separated by roughly their diameter from posterior laterals. Sternum longer than wide, unmodified. Chelicerae with 3-6 promarginal teeth, often on ca- rina, and 5-10 retromarginal denticles. Abdomen longer than wide, tracheal spira- cle midway between epigastric furrow and base of spinnerets. Leg formula 1423, legs long, thin, pale white. Leg I greatly elon- gated. Metatarsi I and II with two pairs of ventral spines. Coxae of males often with spurs and knobs; leg III spination often reduced. Palpus with an elongated median apophysis, enlarged conductor and conspicuous embolus. Retrolateral tibial apophysis greatly expanded except in W. wunda. Epigyna and internal genitalia small and diverse. Variation. None of the species in this genus show any significant individual or geographic intraspecific variation in struc- ture, size or coloration. Key to Species la. Carapace and abdomen with dark mark- ings saltabunda lb. Carapace and abdomen without dark markings 2 2a. Males 3 2b. Females . 7 3a. At least one pair of coxae modified with spurs or knobs 4 3b. All coxae unmodified alba 4a. Coxae I and/or II modified with spurs or knobs 5 4b. Coxae III and/or IV modified with spurs or knobs - 6 5a. Retrolateral tibial apophysis more than half the tibial length (Fig. 93) — bryantae 5b. Retrolateral tibial apophysis less than half the tibial length ( Fig. 95 ) ._ wunda 6a. Retrolateral tibial apophysis greatly ex- panded at tip (Fig. 86) ..._. tantilh 6b. Retrolateral tibial apophysis not greatly expanded at tip (Fig. 88) immaculella 7a. Epigynum with long ducts (Figs. 91, 97, 98 ) 8 7b. Epigynum without long ducts (Figs. 90, 96) : 10 8a. Epigynum with a heart-shaped atrium (Fig. 97) wunda 8b. Epigynum without a heart-shaped atrium 9 9a. Epigynal ducts terminating far anterior of epigynal openings ( Fig. 91 ) tantilla 9b. Epigynal ducts temiinating near epigynal openings ( Fig. 98 ) immaculella 10a. Epigynum with anterolateral flaps, with- out a medial ridge (Fig. 90) alba 10b. Epigynum without anterolateral flaps, with a medial ridge (Fig. 96) bryantae Spideh Family Anyphaenidaf, • Platnick 243 ^; V) i ) !-•- \U^ •^ •, rf ? • ^ _- •^-^•.^% •\ \/^ Wulfila saltabundQ^ 1 --—— ^^"^ ^^-j ^ \ Ifila tantilla 1 V i r i 1 1 ■^ Wulfila immaculella | VV \ U '^ — u-^/- ... r Wulfllo albo \-- - 1 ' ■ 1 — -'i V .' N . ^"^ /^ v-4 /■•f \ \ Wulfllo bryantae "\ Map 3. Distributions of Teudis calcar, Wulfila alba, W. bryantae, W. immaculella, W. saltabunda, W. tantilla and IV. wunda. Wulfila pallidas O. P.-Cambridge Figure 144 Wulfila palUdus O. P.-Cambridge, 1895, Biologia Central! Americana, Aran., 1: 159, pi. 19, fig. 11 ( 9 ). Female holotype from Teapa, Ta- basco, Mexico, in BMNH, examined. Bonnet, 1959, Bibliographia Araneorum, 2: 4832. Wulfila pallida, Simon, 1897, Hist. Natur. Araign., 2: 94. Roewer, 1954, Katalog der Araneae, 2: 554. Vulfila pallida, Simon, 1897, Hist. Natur. Araign., 2: 103. Thi.s Mexican .species, though belonging to a distinct species group, closely resem- bles the North American Wulfila in body form, leg length and coloration. It is the type .species of Wulfila. Wulfila saltabunda (Hentz), new combination Map 3; Figures 81, 82, 89, 99 Cluhiona saltabunda Hentz, 1847, J. Boston Soc. Natur. Hist., 5: 453, pi. 23, fig. 23 ( 9 ). Fe- male holotype from Alabama in Boston Soc. Natur. Hist. (Boston Mu.seum of Science), de- stroyed by beetles. Amjphaena saltabunda, Fmerton, 1890, Trans. Connecticut Acad. Sci., 8: 187, figs. 4-4d, $, $ . Emerton, 1902, Common Spiders, p. 14, figs. 46, 47, 5 , 9 . Gayenna saltabunda, Comstock, 1912, Spider Book, p. 563, figs. 638, 639, $, 9 . Anyphaenella saltabunda, Bryant, 1931, Psyche, 38: 116, pi. 7, figs. 18, 22, $, 9. Comstock, 1940, Spider Book, rev. ed., p. 576, figs. 638, 639, $, 9. Ka.ston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 406, figs. 1465- 1470, $, 9. Roewer, 1954, Katalog der Araneae, 2: 530. Bonnet, 1955, Bibliographia Araneorum, 2: 349. Diarior eye row 0.49 mm long, slightly rcx-urved; posterior eye row 0.62 mm long, procurved; MOQ length 0.26 mm, front width 0.20 mm, back width 0.30 mm; eye interdistances (mm): AME- AME 0.07,^ AME-ALE 0.07, PME-PME 0.15, PME-PLE 0.13, ALE-PME 0.06. Sternum 0.97 mm long, 0.55 mm wide. Chelicerae 0.73 mm long with 3 promar- ginal teeth on a carina and 7 retromarginal denticles. Abdomen 1.80 mm long, 1.12 mm wide. Epigastric fvnrow 0.56 mm from tracheal spiracle, spiracle 0.68 mm from base of spinnerets. Coxae I with a small knob, coxae II with two spurs. Tibial lengths (mm) and in- dices: I 2.76, 4; II 1.85, 7; III 0.92, 15; IV 1.89, 7. \'entral spination: tibiae I, II 2- 2-0, III 1-2-0, IV 1-1-0; metatarsi I, II 2- 2-0, III, IV 2-1-2. Palpus as in Figures 92, 93. Female (Hidalgo Co., Texas). Colora- tion as in male of Wulfila alba. Total length 3.78 mm. Carapace 1.44 mm long, 0.99 mm wide, cephalic width 0.74 mm, clypeus height 0.07 mm. Eves: diameters (mm): AME 0.06, ALE 6.07, PME 0.06, PLE 0.07; anterior eye row 0.42 mm long, slightly recurved; posterior eye row 0.59 mm long, procurved; MOQ length 0.23 mm, front width 0.17 mm, back width 0.27 mm; eye interdistances (mm): AME-AME 0.06, AME-ALE 0.05, PME- PME 0.14, PME-PLE 0.13, ALE-PLE 0.06. Sternum 0.74 mm long, 0.64 mm wide. Chelicerae 0.62 mm long with 5 promar- ginal teeth and 5 retromarginal denticles. Abdomen 2.59 mm long, 2.16 nun wide. Epigastric furrow 0.90 mm from tracheal spiracle, spiracle 0.88 mm from base of spinnerets. 250 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 Legs unmodified. Tibial lengths (mm) and indices: I 2.24, 5; II 1.37, 9; III 0.72, 19; IV 1.35, 10. Ventral spination as in male except tibiae III, IV 2-2-0 and meta- tarsi III, IV 2-0-2. Epigynum as in Figure 96, internal genitalia as in Figure 102. Natural history. Mature males have been taken from late April through early June, mature females from early April through early December. Nothing is known of the habits of this species. Distribution. Southern Texas and Ta- maulipas (Map 3). Wulfila wunda new species Map 3; Figures 94, 95, 97, 104 Wulfila immaculata, Bryant (not Banks), 1936, Psyche, 43: 98, fig. 1, $. Male allotype from Brichell Hammock, Florida Keys, in MCZ, examined. Not Wulfila immaculata Banks, 1914, Bull. Amer. Mus. Natur. Hist., 33: 640, pi. 43, fig. 7, 9 . Female holotype from Vinales, Pinar del Rio, Cuba, in AMNH, examined. Types. Male holotype, female paratype from Tavernier, Monroe Co., Florida, 16 February 1951 (A. M. Nadler), deposited in AMNH. Male and female paratypes from Dade Co., Florida, deposited in MCZ. The specific name is an arbitrary combina- tion of letters. Diagnosis. Wulfila wunda is a distinc- tive species the genitalia of which are quite different from those of the other Wulfila in America north of Mexico: the retrolateral tibial apophysis is very short ( Fig. 95 ) and the epigynum has an atrium (Fig. 97). Male (Dade Co., Florida). Coloration as in Wulfila alba. Total length 3.42 mm. Carapace 1.55 mm long, 1.08 mm wide, cephalic width 0.68 mm, clypeus height 0.06 mm. Eyes: diameters (mm): AME 0.05, ALE 0.07, PME 0.08, PLE 0.08; anterior eye row 0.48 mm long, straight; posterior eye row 0.59 mm long, procurved; MOQ length 0.20 mm, front width 0.14 mm, back width 0.29 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.09, PME-PME 0.14, PME-PLE 0.12, ALE-PLE 0.04. Sternum 1.06 mm long, 0.70 mm wide. Chelicerae 0.85 mm long with 4 promar- ginal teeth and 6 retromarginal denticles. Abdomen 1.91 mm long, 1.01 mm wide. Epigastric furrow 0.70 mm from ti'acheal spiracle, spiracle 0.76 mm from base of spinnerets. Coxae II with a small knob. Tibial lengths (mm) and indices: I 4.10, 3; II 1.87, 8; III 1.01, 15; IV 2.05, 7. Ventral spination: tibiae I, II 2-2-0, III 0-1-0, IV 0-2-0; metatarsi I, II 2-2-0, III 0-2-0, IV 2-1-2. Palpus as in Figures 94, 95. Female (Dade Co., Florida). Colora- tion as in male of Wulfila alba. Total length 3.74 mm. Carapace 1.55 mm long, 1.15 mm wide, cephalic width 0.72 mm, clypeus height 0.07 mm. Eyes: diameters (mm): AME 0.04, ALE 0.06, PME 0.07, PLE 0.07; anterior eye row 0.49 mm long, straight; posterior eye row 0.59 mm long, procurved; MOQ length 0.20 mm, front width 0.15 mm, back width 0.30 mm; eye interdistances ( mm ) : AME- AME 0.06, AME-ALE 0.10, PME-PME 0.16, PME-PLE 0.12, ALE-PME 0.04. Sternum 0.90 mm long, 0.67 mm wide. Chelicerae 0.65 mm long with 5 promar- ginal teeth and 9 retromarginal denticles. Abdomen 2.16 mm long, 1.15 mm wide. Epigastric furrow 0.74 mm from tracheal spiracle, spiracle 0.83 mm from base of spinnerets. Legs unmodified. Tibial lengths (mm) and indices: I 3.13, 4; II 1.44, 10; III 0.76, 20; IV 1.58, 9. Ventral spination as in male except tibiae III 1-2-0 and IV 0-1-0 and metatarsi III 1-2-0 and IV 1-2-2. Epigynum as in Figure 97, internal geni- talia as in Figure 104. Natural history. Mature males have been taken from mid-February through mid- May, mature females apparently year- round. Nothing is known of the habits of this .species. Spider Family Anyphaenidae • Platnick 251 Distrihuiion. Southern Florida, Culxi, and Mona Island ( Map 3). Aysha Keyserling Aysha Keyserling, 1891, Spinn. Ainer. ( Brasil. Spiiin.), 3: 83, 129. Type species Aysha pros- pera Keyserling, 1891, designated by Simon, 1897, Hist. Natm-. Araig., 2: 104. Diagnosis. Aysha is easily recognized by the greatly adxanced placement of the tracheal spiracle, located just behind the epigastric furrow. The genitalic structure is quite different from that of Amjphacna and Wulfila and the genus undoubtedly represents a different evolutionary line. There are probably more than thirty spe- cies in this genus; they occur commonly in both North and South America. Description. Total length 4-9 mm. Cara- pace longer than wide, narrowed in front to more than half its maximum width. Clypeus height roughly equal to anterior median eye diameter. All eyes subequal in size. Procurved posterior eye row longer than recurved anterior row. Median ocular quadrangle longer than wide in front, wider in back than long. Anterior median eyes separated by slightly less than their diameter, slightly closer to anterior laterals. Posterior medians separated by up to twice their diameter, closer to posterior laterals. Anterior laterals separated by their radius from posterior laterals. Sternum longer than wide, unmodified. Chelicerae with 3- 4 promarginal teeth and 7-9 retromarginal denticles. Abdomen longer than wide, tracheal spiracle much closer to epigastric furrow than to base of spinnerets. Leg formula 1423, legs unmodified. Metatarsi I and II with one pair of \'entral spines. Pal- pus with greatly enlarged base of embolus, long curving embolus and short conductor. Ventral tibial apophysis sometimes present in addition to retrolateral tibial apophysis. Epigynum with anterior median opening and two sidepieces. Internal genitalia with long, sometimes coiling, ducts. Variation. Only Aysha gracilis shows significant variation, and that is in size and not strnetine or coloration. The size of both the whole animal and ol the genitalia vary geographically. The largest .specimens ( males with cymbium length averaging 1.3 mm) occur in Virginia and surrounding states, with smaller individuals occurring in the north (New England and Michigan males with cymbium length averaging 1.1 mm) and in the south (Texas males with cymbium length averaging 0.9 mm). Key to Species la. Males 2 lb. Females -..- 7 2a. Palpus without a ventral tibial apophysis (VTA) (Figs. Ill, 119) -.._ 3 2b. Palpus with a ventral tibial apophysis (VTA), sometimes small, transparent, eas- ily overlooked (Figs. 113, 115, 117, 121) 4 3a. Embolus restricted to distal half of palpal bulb (Fig. 118) arunda 3b. Embolus not restricted to distal half of pal- pal bulb (Fig. 110) velox 4a. VTA erect, sclerotized, relatively large (Figs. 113, 115) 5 4b. VTA recumbent, transparent, relativelv small (Figs. 117, 121) '. 6 5a. Distal retrolateral tip of tegulum with a flap covering embolus (Fig. 112) decepta 5b. Distal retrolateral tip of tegulum with a sharp point underlying embolus (Fig. 114) — incursa 6a. Base of embolus recurved, with a sharp spike (Fig. 120) camhridgei 6b. Base of embolus not recurved, forming a smooth arc (Fig. 116) fitacilis 7a. Internal genitalia with simple uncoiled ducts (Figs. 124, 127, 141, 143) 9 7b. Internal genitalia coiled or with accessory ducts (Figs. 125, 142) 8 8a. Internal genitalia highly coiled ( Fig. 125 ) '...... vclox 8b. Internal genitalia not coiled but with loop- ing accessory ducts (Fig. 142) arunda 9a. Median epig\'nal opening near anterior rim (Figs. 123, 126, 138) 10 91). Median epigynal opening near middle of epigynum ( Fig. 140 ) -.. gracilis 10a. Median epig\nal opening much wider than epigynal sidepieces (Fig. 138) cand)ridgc'i lOb. Median epigynal opening not wider than epigynal sidepieces (Figs. 123, 126) 11 11a. Base of epig>nal sidepieces near epigastric furrow (Fig. 126); internal genitalia with angular ducts (Fig. 127) incursa 252 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 . ^ Aysha gracilis I / ; 1 / HM ~T— 7 I Ayshc velox r? i? ♦^^'^r cQ^* i i " 1 \ r' ) ( -A Aysha arunda .„ V Map 4. Distributions of Aysha arunda. A. cambridgei, A. decepta, A. gracilis, A. incursa and A. velox. 111). Base of epigynal sidepieces far from epigastric furrow (Fig. 123); internal genitalia with rounded ducts (Fig. 124) decepta Aysha prospers Keyserling Figure 145 Ay.sha prospera Keyserling, 1891, Spinnen Amer- ikas (Brasil. Spinn.), 3: 129, pi. 4, fig. 88 ( 9 ). Female holotype from Rio Grande, Brasil, in BMNH, examined. Roewer, 1954, Katalog der Araneae, 2: 533. Bonnet, 1955, Biblio- graphia Araneorum, 2: 838. This South American species, type spe- cies of Aysha, is a member of a large, dis- tinct species group. Somatic characters clearly ally it with the North American forms included in the genus. Aysha gracilis (Hentz) Map 4; Figures 116, 117, 140, 143 Chihiona gracilis Hentz, 1847, J. Boston Soc. Natur. Hist., 5: 452, pi. 23, fig. 9(5). Type specimens from North Carolina and Alabama in Boston Soc. Natur. Hist. (Boston Museum of Science), destroyed by beetles. Anijphaena gracilis, L. Koch, 1836, Arach. Fam. brassidae, p. 195, pi. 8, fig. 130, 9 . Comstock, 1912, Spider Book, p. 561, fig. 633, $ (not fig. 632). Anijphaena rubra Emerton, 1890, Trans. Connecti- cut Acad. Sci., 8: 186, pi. 6, fig. 1(9). Male allotype (?) from Franklin Park, Boston, Mas- sachusetts, in MCZ, examined. Emerton, 1909, Trans. Connecticut Acad. Sci., 14: 220, pi. 9, fig. 8-8c, $ . Aysha gracilis, Bryant, 1931, Psyche, 38: 119, pi. 7, fig. 13, pi. 8, fig. 26, $, 9. Chickering, 1939, Pap. Michigan Acad. Sci., 24: 53, figs. 9-11, $, 9. Comstock, 1940, Spider Book, rev. ed., p. 575, fig. 633, $ (not fig. 632). Kaston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 405, figs. 1452, 1459-1464, $, 9 . Roewer, 1954, Katalog der Araneae, 2: 534. Bonnet, 1955, Bibliographia Araneorum, 2: 837. Diagnosis. Aysha gracilis is closest to A. cambridgei but lacks the sharp spike on the proximal edge of the base of embolus (Fig. 116) of that .species. Females have Spider Family Anyphaenidae • Platnick 253 Plate 9 Figures 110, 112, 114, 116. Left palpi, ventral view. Figures 111, 113, 115, 117 110, 111. Aysha velox (Becker). 112, 113. Aysha decepta (Banks) 116, 117. Aysha gracilis (Hentz). Left palpi, retrolateral view/. 114, 115. Aysha incursa (Chamberlin). the median epigynal opening near the mid- long, 2.02 mm wide, cephalic width 1.17 die of the epigynum (Fig. 140). Variation mm, clypeus heigiit 0.09 mm, light orange- in thi.s species is discussed above. brown, darkest anteriorly, with thin dark Male (Middlesex Co., Massachusetts), border and two dark paramedian longitudi- Total length 5.73 mm. Carapace 2.56 mm nal bands. Eyes: diameters (mm): AME 254 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 0.09, ALE 0.11, PME 0.09, PLE 0.11; an- terior eye row 0.60 mm long, slightly re- cnrved; posterior eye row 0.80 mm long, procurved; MOQ length 0.32 mm, front width 0.26 mm, back width 0.38 mm; eye interdistances (mm): AME-AME 0.09, AME-ALE 0.07, PME-PME 0.19, PME- PLE 0.15, ALE-PLE 0.06. Sternum 1.44 mm long, 1.08 mm wide, light orange-brown with translucent border and darkened extensions to coxae. Chelic- erae 1.12 mm long with 4 promarginal teeth and 8 retromarginal denticles, dark orange-brown proximally, dark brown dis- tally. Labium and endites light orange- brown, darkest proximally. Endites sharply invaginated at middle. Abdomen 3.20 mm long, 1.73 mm wide, pale grayish-brown with transverse rows of dark markings, venter pale. Epigastric furrow 0.40 mm from tracheal spiracle, spiracle 1.73 mm from base of spinnerets. Legs light orange-brown with distal seg- ments darkest. Tibial lengths (mm) and indices: I 2.64, 10; II 1.87, 15; III 1.19, 26; IV 2.09, 15. Vential spination: tibiae I, II 2-2-2, III 1-2-2; IV 2-2-2; metatarsi I, II 2-0-0, III 2-1-2, IV 2-2-2. Palpus as in Figure 116, 117. Female (Washington Co., Arkansas). Coloration as in male. Total length 8.42 mm. Carapace 2.75 mm long, 2.11 mm wide, cephalic width 1.47 mm, clypeus height 0.10 mm. Eyes: diameters (mm): AME 0.14, ALE 0.14, PME 0.13, PLE 0.14; anterior eye row 0.43 mm long, recurved; posterior eye row 1.04 mm long, procurved; MOQ length 0.43 mm, front width 0.36 mm, back width 0.49 mm; eye interdistances (mm): AME- AME 0.09, AME-ALE 0.08, PME-PME 0.22, PME-PLE 0.18, ALE-PLE 0.06. Sternum 1.84 mm long, 1.31 mm wide. Chelicerae 1.57 mm long with teeth as in male. Abdomen 5.76 mm long, 3.53 mm wide. Epigastric furrow 0.68 mm from tracheal spiracle, spiracle 3.24 mm from base of spinnerets. Tibial lengths (mm) and indices: I 2.56, 16; II 1.94, 20; III 1.26, 30; IV 2.30, 17. Ventral spination: tibiae I 2-2-0, II 1-2-1, III 1-1-2, IV 1-2-2; metatarsi I, II 2-0-0, III 2-0-2, IV 2-2-2. Epigynum as in Figure 140, internal genitalia as in Figure 143. Natural history. Mature males and fe- males have been taken year-round. Speci- mens have been taken by sweeping, in pitcher plants, on loblolly pine, in fall web- worm nests and frequently in houses. Distribution. New England west to Wis- consin and Iowa, south to Florida and east- ern Texas ( Map 4 ) . Aysha cam bridge! Bryant Map 4; Figures 120, 121, 138, 141 Aysha cambiidgei Bryant, 1931, Psyche, 38: 119, pi. 7, fig. 15 { $ ). Male holotype from Guanajuato, Mexico, in MCZ, examined. Roe- wer, 1954, Katalog der Araneae, 2: 532. Bon- net, 1955, Bibliographia Araneorum, 2: 836. Diagnosis. Aysha cambridgei is closely related to A. gracilis but has a distinctive spike on the proximal edge of the base of the embolus (Fig. 120) and the median epigynal opening near the anterior rim of the epigynum ( Fig. 138 ) . Male (Jeff Davis Co., Texas). Colora- tion as in Aysha gracilis except that the ab- domen is pale white with two dark para- median longitudinal bands. Total length 5.87 mm. Carapace 2.41 mm long, 1.91 mm wide, cephalic width 0.97 mm, clypeus height 0.11 mm. Eyes: diameters (mm): AME 0.11, ALE 0.12, PME 0.11, PLE 0.11; anterior eye row 0.57 mm long, recurved; posterior eye row 0.75 mm long, procurved; MOQ length 0.33 mm, front width 0.28 mm, back width 0.38 mm; eye interdistances (mm): AME- AME 0.06, AME-ALE 0.05, PME-PME 0.16, PME-PLE 0.11, ALE-PLE 0.05. Sternum 1.42 mm long, 1.01 mm wide. Chelicerae 0.98 mm long with 4 promar- ginal teeth and 7 retromarginal denticles. Abdomen 3.49 mm long, 1.58 mm wide. Epigastric furrow 0.68 mm from tracheal Spideu Family Anyphaenidae • Platnick 255 122 126 127 125 Plate 10 Figures 118, 120. Left palpi, ventral view. Figures 119, 121. Left palpi, retrolateral view. Figures 122, 123, 126. Epigyna, ventral view. Figures 124, 125, 127. Internal genitalia, dorsal view. 118, 119. Aysha arunda new species. 120, 121. Aysha cambridgei Bryant. 122, 125. Aysha velox (Becker). 123, 124. Aysha decepta (Banks). 126, 127. Aysha incursa (Chamberlin). .spiracle, spiracle 1.55 mm from base ot Ventral spination: tibiae I, II 2-2-2, III spinnerets. 1-2-2, IV 2-2-2; metatarsi I, II 2-0-0, III, Tibial lengths (mm) and indices: I 3.06, IV 2-2-2. 8; II 1.87, 13; III 1.28, 21; IV 2.16, 12. Palpus as in Figures 120, 121. 256 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 Female (Henderson Co., Texas). Color- ation as in male. Total length 8.50 mm. Carapace 3.35 mm long, 2.52 mm wide, cephalic width 1.69 mm, clypeus height 0.12 mm. Eyes: diameters (mm): AME 0.14, ALE 0.16, PME 0.14, PLE 0.14; anterior eye row 0.84 mm long, recm'ved; posterior eye row 1.11 mm long, procurved; MOQ length 0.42 mm, front width 0.37 mm, back width 0.50 mm; eye interdistances (mm): AME- AME 0.10, AME-ALE 0.07, PME-PME 0.22, PME-PLE 0.20, ALE-PLE 0.05. Sternum 1.91 mm long, 1.22 mm wide. '&' Chelicerae 1.69 mm long with teeth as in male. Abdomen 5.04 mm long, 2.88 mm wide. Epigastric furrow 0.61 mm from tracheal spiracle, spiracle 3.17 mm from base of spinnerets. Tibial lengths (mm) and indices: 12.88, 12; II 2.07, 17; III 1.40, 26; IV 2.57, 15. Ventral spination as in male except tibiae I, II 2-2-0 and III 2-2-2. Epigynum as in Figure 138, internal genitalia as in Figure 141. Natural history. Mature males have been taken from mid-June through early August, mature females from late May through early August. Specimens have been taken on trees and shrubs. Distribution. South central states from Alabama to western Texas, south to central Mexico (Map 4). Aysha decepta (Banks) Map 4; Figures 112, 113, 123, 124 Amjphaena decepta Banks, 1899, Proc. Ent. Soc. Washington, 4: 190. Female holotype from Brazos Co., Texas, in MCZ, examined. Aysha mimita F. O. P.-Canibridge, 1900, Biologia Centrali Americana, Aran., 2: 99, pi. 7, figs. 18-19 { $, ? ). Male holotype, female allotype from Guatemala, in BMNH, examined. Bryant, 1931, Psyche, 38: 120, pi. 7, fig. 17, $. Roe- wer, 1954, Katalog der Araneae, 2: 533. Bon- net, 1955, Bibliographia Araneorum, 2: 838. NEW SYNONYMY. Aijsha decepta, Bryant, 1931, Psyche, 38: 120, pi. 7, fig. 16, pi. 8, fig. 27, $, 9. Roewer, 1954, Katalog der Araneae, 2: 534. Bonnet, 1955, Bibliographia Araneorum, 2: 836. Diagnosis. Aysha decepta is very closely related to A. incursa but has a characteris- tic flap (on the retrolateral tip of the tegu- lum) that covers the embolus (Fig. 112), while the base of the epigynal sidepieces is a considerable distance from the epigas- tric furrow ( Fig. 123 ) . Both morphological and zoogeographical data (Map 4) indi- cate that these two species are each other's nearest relatives. Male (Hidalgo Co., Texas). Coloration as in Aysha camhridgei. Total length 4.82 mm. Carapace 2.25 mm long, 1.76 mm wide, cephalic width 1.06 mm, clypeus height 0.10 mm. Eyes: diameters (mm): AME 0.08, ALE 0.10, PME 0.11, PLE 0.11; anterior eye row 0.58 mm long, straight; posterior eye row 0.75 mm long, procurved; MOQ length 0.23 mm, front width 0.24 mm, back width 0.39 mm; eye interdistances ( mm ) : AME- AME 0.08, AME-ALE 0.06, PME-PME 0.18, PME-PLE 0.12, ALE-PLE 0.05. Sternum 1.37 mm long, 0.85 mm wide. Chelicerae 0.97 mm long with 4 promar- ginal teeth and 7 retromarginal denticles. Abdomen 2.74 mm long, 1.39 mm wide. Epigastric furrow 0.38 mm from tracheal spiracle, spiracle 1.28 mm from base of spinnerets. Tibial lengths (mm) and indices: I 2.54, 9; II 1.67, 14; III 1.01, 25; IV 1.89, 16. Venti-al spination: tibiae I 2-2-0, II 1-2-0, III 1-2-2, IV 2-2-2; metatarsi I, II 2-0-0, III, IV 2-2-2. Palpus as in Figures 112, 113. Female (E. Baton Rouge Parish, Louisi- ana). Coloration as in male of Aysha cam- hridgei. Total length 5.76 mm. Carapace 2.45 mm long, 1.80 mm wide, cephalic width 1.17 mm, clypeus height 0.09 mm. Eyes: diameters (mm): AME 0.10, ALE 0.13, PME 0.12, PLE 0.12; anterior eye row 0.60 mm long, recurved; posterior eye row 0.76 mm long, procurved; MOQ length 0.30 mm, front width 0.26 mm, back width Spider Family Anyphaenidae • Plahiick 257 0.3(S mm; eye interdistances (mm): AME- AME 0.07, AME-ALE 0.05, PME-PME 0.15, PME-PLE 0.11, ALE-PLE 0.04. Sternum 1.35 mm long, 0.95 mm widc\ Chelicerae 0.89 mm long with 4 promar- ginal teeth and 8 retromarginal dentieles. Abdomen 3.51 mm long, 2.20 mm wide. Epigastric furrow 0.41 mm from tracheal spiracle, spiracle 1.78 mm from base of .spinnerets. Tibial lengths (mm) and indices: I 1.84, 14; II 1.40, 19; III 0.93, 29; IV 1.82, 17. Ventral spination as in male except tibiae III 1-1-2 and metatarsi III 2-1-2. Epigynum as in Figure 123, internal genitalia as in Figure 124. Natural history. Mature males and fe- males have been taken every month except January and February. Specimens are commonly found in great quantities in wasp nests and occasionally in houses. Distribution. Northern Florida west to eastern Texas, south to Costa Rica ( Map 4). Aysha incursa (Chamberlin) Map 4; Figures 114, 115, 126, 127 Anypliacna incursa Chamberlin, 1919, Pomona Coll. J. Ent. Zool., 12: 12, pi. 5, fig. 2(9). Female holotype from Claremont, California, in MCZ, examined. Bryant, 1931, Psyche, .38: 120 (sub Aysha dccepta [sic]). Roewer, 1954, Katalog der Araneae, 2 : 534 ( sub Aysha de- cepta [sic]). Bonnet, 1955, Bibliographia Araneorum, 2: 836 (sub Aysha deccpta [sic]). Anyphaena johnstoni Chamberlin, 1924, Proc. California Acad. Sci., 12: 662, figs. 105, 106 (5, 9 ). Female holotype, male allotype from San Pedro Nolasco Island, Gulf of California, in California Academy of Sciences. Paratype male from San Marcos Island, Gulf of California, in MCZ, examined. Bryant, 1931, Psyche, 38: 120 (sub Aysha decepta [sic]). Bonnet, 1955, Bibliographia Araneorum, 2: 836 (sub Aysha dccepta jsic] ). Anyphaena ni^.rifwns Chamberlin and Woodburw 1929, Proc. Biol. Soc. Washington, 42: 137, pi. 1, fig. 4 ( 9 ). Female holotype from St. George, Utah, in AMNH, e.\amined. NEW SYNONYMY. Aysha nigrifrons, Bryant, 1931, Psyche, 38: 121. Roewer, 1954, Katalog der Araneae, 2: 534. Bonnet, 1955, Bibliographia Araneorum, 2: 838. Diapwsis. Aysha incursa is very closely related to A. decepta but has a distinctive sharp point on the retrolateral tip of the tegulum (Fig. 114), while the base of the epigynal sidepieces is near the epigastric furr(')w (Fig. 126). Male (Tulare Co., California). Colora- tion as in Aysha camhrid^ei. Total length 6.08 mm. Carapace 3.02 mm long, 2.18 mm wide, cephalic width 1.22 mm, clypeus height 0.12 mm. Eyes: diameters (mm): AME 0.11, ALE 0.12, PME 0.11, PLE 0.12; anterior eye row 0.66 mm long, recurved; posterior eye row 0.84 mm long, procurved; MOQ length 0.33 mm, front width 0.31 mm, back width 0.42 mm; eye interdistances (mm): AME- AME 0.10, AME-ALE 0.07, PME-PME 0.21, PME-PLE 0.17, ALE-PLE 0.05. Sternum 1.67 mm long, 1.08 mm wide. Chelicerae 1.22 mm long with 3 promar- ginal teeth and 8 retromarginal denticles. Abdomen 3.38 mm long, 1.80 mm wide. Epigastric furrow 0.50 mm from tracheal spiracle, spiracle 1.75 mm from base of spinnerets. Tibial lengths (mm) and indices: I 3.15, 9; II 2.16, 14; III 1.40, 26; IV 2.34, 16. Ventral spination: tibiae I 2-2-0, II, III, IV 2-2-2; metatarsi I, II 2-0-0, III, IV 2- 2-2. Palpus as in Figures 114, 115. Female (Santa Barbara Co., California). Coloration as in male of Aysha camhridgei. Total length 5.72 mm." Carapace 2.09 mm long, 1.66 mm wide; cephalic width 1.04 mm, clypeus height 0.05 mm. Eyes: diameters (mm): AME 0.08, ALE 0.09, PME 0.10, PLE 0.10; anterior eye row 0.50 mm long, recurved; posterior eye row 0.67 mm long, procurved; MOQ length 0.25 mm, front width 0.23 mm, back width 0.33 mm; eye interdistances (mm): AME- AME 0.07, AME-ALE 0.05, PME-PME 0.14, PME-PLE 0.11, ALE-PLE 0.06. Sternum 1.30 mm long, 0.85 mm wid(\ Chelicerae 0.70 mm long with 3 promar- ofinal teeth and 8 retromariiiinal dcMiticles. 258 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 Abdomen 4.00 mm long, 2.38 mm wide. Epigastric furrow 0.70 mm from tracheal spiracle, spiracle 1.91 mm from base of spinnerets. Tibial lengths (mm) and indices: I 1.57, 14; II 1.26, 17; III 0.86, 29; IV 1.64, 15. Ventral spination as in male except tibiae II, III 1-1-0 and IV 1-1-2 and metatarsi III 2-0-2. Epigynum as in Figure 126, internal genitalia as in Figure 127. Natural history. Mature males have been taken from late April through early Sep- tember, mature females year-round. Speci- mens have been taken on poplars, in fields, and in houses. Distribution. California west to Utah, south to southern Mexico (Map 4). Aysha velox (Becker) Map 4; Figures 110, 111, 122, 125 Anijphaena vcIox Becker, 1879, Ann. Ent. Soc. Belgique, 22: 83, pi. 2, figs. 5-7 ( 5 ). Female holotype from Pascagoula, Mississippi, should be in the Institute Royal des Sciences Naturelles de Belgique but could not be located there by Mr. J. Kekenbosch in 1971; lost, presumed de- stroyed. Banks, 1904, Proc. Acad. Nat. Sci. Philadelphia, 56: 123, pi. 8, fig. 19, $. Amjphaena floridana Banks, 1896, Trans. Amer. Ent. Soc, 23: 63. Female holotype from Lake Worth, Florida, in MCZ, examined. Aysha orlandensis Tullgren, 1901, Bih. Svenska Akad., 27: 19, fig. 4 ( ? ). Female holotype from Orlando, Florida, in Uppsala Univ. Zool. Mus., examined. Bryant, 1931, Psyche, 38: 119 (sub Aysha gracilis [sic]). Roewer, 1954, Katalog der Araneae, 2: 534 (sub Aysha graci- lis [sic]). Bonnet, 1955, Bibliographia Araneo- rum, 2: 837 (sub Aysha gracilis [sic]). NEW SYNONYMY. Aysha velox, Banks, 1909, Estacion central agrono- mica de Cuba, Second Report, p. 158. Bryant, 1931, Psyche, 38: 119, pi. 7, fig. 14, pi. 8, fig. 34, $ , 5 . Roewer, 1954 Katalog der Araneae 2: 534. Bonnet, 1955, Bibliographia Araneo- rum, 2: 839. Chiracanthium falculum Chamberlin, 1925, Bull. Mus. Comp. Zool., 67: 220. Male holotype from Sebastian, Florida, in MCZ, examined. Diagnosis. Aysha velox is a distinctive species easily recognized by its short retro- lateral tibial apophysis and its lack of a ventral tibial apophysis (Fig. Ill) and the embolus' not being restricted to the distal half of the palpal bulb (Fig. 110). The coiled internal ducts of females (Fig. 125) are diagnostic. Male (Alachua Co., Florida). Colora- tion as in Aysha gracilis except that the abdomen lacks dark markings. Total length 7.31 mm. Carapace 3.45 mm long, 2.52 mm wide, cephalic width 1,51 mm, clypeus height 0.13 mm. Eyes: diameters (mm): AME 0.15, ALE 0.14, PME 0.13, PLE 0.15; anterior eye row 0.80 mm long, recurved; posterior eye row 1.01 mm long, procurved; MOQ length 0.42 mm, front width 0.38 mm, back width 0.47 mm; eye interdistances (mm): AME- AME 0.09, AME-ALE 0.09, PME-PME 0.22, PME-PLE 0.18, ALE-PLE 0.05. Sternum 1.92 mm long, 1.28 mm wide. Chelicerae 1.58 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 4.14 mm long, 1.76 mm wide. Epigastric furrow 0.31 mm from tracheal spiracle, spiracle 2.46 mm from base of spinjierets. Tibial lengtlis (mm) and indices: I 3.92, 8; II 2.86, 12; III 1.69, 21; IV 2.52, 14. Ventral spination: tibiae I-IV 2-2-2; meta- tarsi I, II 2-0-0, III, IV 2-2-2. Palpus as in Figure 110, 111. Female (Alachua Co., Florida). Colora- tion as in male. Total length 8.42 mm. Carapace 3.96 mm long, 2.88 mm wide; cephalic width 1.87 mm, clypeus height 0.14 mm. Eyes: diameters (mm): AME 0.15, ALE 0.14, PME 0.14, PLE 0.14; anterior eye row 1.02 mm long, recurved; posterior eye row 1.31 mm long, procurved; MOQ length 0.48 mm, front width 0.45 mm, back width 0.57 mm; eye interdistances ( mm ) : AME- AME 0.14, AME-ALE 0.14, PME-PME 0.28, PME-PLE 0.27, ALE-PLE 0.09. Sternum 2.16 mm long, 1.62 mm wide. Chelicerae 1.87 mm long with teeth as in male. Abdomen 4.50 mm long, 2.41 mm wide. Spider Family Axyphaexidae • Phi f nick 259 Epigastric furrow 0.36 mm from traclical spiracle, spiracle 2.48 mm from base of .spinnerets. Tibial lengths (mm) and indices: I 3.46, 11; II 2.68,^14; III 1.66, 24; IV 2.79, 14. Ventral spination as in male. Epigynum as in Figure 122, internal genitalia as in Figure 125. Natural histonj. Mature males and fe- males have been taken year-round. Speci- mens have been taken on Casuarina sp., Citrus sp., Paurotis sp., Calatuandra sp., Pinus sp., Ncluniho sp., and in houses. Distribution. North Carolina west to Ar- kansas, south to east Texas and Florida, Cuba, Haiti, the Dominican Republic and Bermuda ( Map 4 ) , Aysha arunda new species Map 4; Figures 118, 119, 139, 142 Types. Male holotype, female paratype from Edinburg, Hidalgo Co., Texas, May 1934 (Mulaik), deposited in AMNH. Male and female paratypes from Hidalgo Co., Texas, deposited in MCZ. The specific name is an arbitrary combination of letters. Diagnosis. Aysha arunda is a distinctive species easily recognized by the restriction of the embolus to the distal half of the palpal bulb (Fig. 118) and the triangular .shape of the epigynum (Fig. 139). Male (Hidalgo Co., Texas). Coloration as in Aysha cambridgei. Total length 6.23 mm. Carapace 3.04 mm long, 2.02 mm wide, cephalic width 1.49 mm, clypeus height 0.10 mm. Eyes: diameters (mm): AME 0.13, ALE 0.13, PME 0.14, PLE 0.14; anterior eye row 0.74 mm long, recurved; posterior eye row 0.95 mm long, procurved; MOQ length 0.44 mm, front width 0.34 mm, back width 0.43 mm; eye interdistances (mm): AME- AME 0.07, AME-ALE 0.06, PME-PME 0.16, PME-PLE 0.19, ALE-PLE 0.07. Sternum 1.73 mm long, 1.24 mm wide. Chelicerae 1.62 mm long with 4 promar- ginal teeth and 8 retromarginal denticles. Abdomen 3.62 mm long, 1.67 mm wide. Epigastric Imrow 0.31 mm from tracheal spiracle, .spiracle 1.75 mm from base of spinnerets. Tibial lengths (mm) and indices: I 4.03, 7; II 2.72, 12; III 1.53, 20; IV 2.58, 15. Ventral spination: tibiae I-IV 2-2-2; meta- tarsi I, II 2-0-0, III, IV 2-2-2. Palpus as in Figures 118, 119. Female (Hidalgo Co., Texas). Colora- tion as in male of Aysha cambridgei. Total length 6.59 mm. Carapace 2.99 mm long, 2.23 mm wide, cephalic width 1.33 mm, clypeus height 0.09 mm. Iwes: diameters (mm): AME 0.11, ALE 6.13, PME 0.13, PLE 0.13; anterior eye row 0.70 mm long, recurved; posterior eye row 0.91 mm long, procurved; MOQ length 0.40 mm, front width 0.32 mm, back width 0.43 mm; eye interdistances (mm): AME- AME 0.10, AME-ALE 0.07, PME-PME 0.18, PME-PLE 0.17, ALE-PLE 0.06. Sternum 1.62 mm long, 1.13 mm wide. Chelicerae 1.37 mm long with 4 promar- ginal teeth and 9 retromarginal denticles. Abdomen 3.76 mm long, 2.12 mm wide. Epigastric furrow 0.40 mm from tracheal spiracle, spiracle 2.23 mm from base of spinnerets. Tibial lengths (mm) and indices: I 2.65, 12; II 2.00,^6; III 1.22, 27; IV 2.25, 16. Ventral spination as in male. Epigynum as in Figure 139, internal genitalia as in Figure 142. Natural history. Mature males have been taken from early May through late Septem- ber, mature females from early April through late September. Nothing is known of the habits of this species. Distribution. Southern Texas (Map 4). Oxysoma Nicolet Oxy.soma NMcolet, 1849, in Gay: Hist. Chili, 10 (3): 511. Type species Oxysonui punctatum Nicolet, 1849, designated by Simon, 1897, Hist. Natur. Araign., 2: 100. Gaycimina Ceitsch, 1935, Anier. Mus. Novitates, No. 805: 21. Type species by iii()i.()t\p\' Gaijcn- ninii hritrlwri Certsch, 1935. Diagnosis. O.xysoma can be (juickly distintiuished from all other North Ameri- 260 Bulletin Museum of Comparative Zoology, Vol. 146, No. 4 can anyphaenids by the presence of only two teeth on the cheKceral retromaigin. In addition, the coloration pattern shown in Fignre 109 is typical for the genus through- out its range. Predominantly South Ameri- can, only one species occurs north of Mexico. Oxysoma is more closely related to Aijsha than to Anyphaena or Wulfila. Description. Total length 5-7 mm. Cara- pace longer than wide, narrowed in front to about half its maximum width. Clypeus height more than twice the anterior median eye diameter. Posterior median, posterior lateral and anterior lateral eyes subequal in size, much larger than anterior medians. Procurved posterior eye row longer than recurved anterior row. Median ocular quadrangle more than twice as wide in back as in front. Anterior median eyes separated by their diameter, closer to anterior laterals than to each other. Pos- terior medians separated by almost three times their diameter, closer to posterior laterals. Anterior laterals separated by their diameter from posterior laterals. Sternum longer than wide, unmodified. Chelicerae with 3 promarginal and 2 re- tromarginal teeth. Abdomen longer than wide, tracheal spiracle roughly midway between epigastric furrow and base of spinnerets. Leg formula 1423, legs un- modified. Metatarsi I and II with one pair of ventral spines. Palpus bulbous, with elongated conductor and conspicuous em- bolus. Retrolateral tibial apophysis lacking. Epigynum on a sclerotized plate. Internal genitalia with two large spermathecae and accessory ducts. Variation. The two males of Oxysoma cubana known from Arizona are slightly larger than the eastern specimens. One has a broken conductor, the other matches the eastern specimens in genitalic details. Oxysoma punctatum Nicolet Oxifsoma punctatum Nicolet, 1849, in Gay: Hist. Chili, 10(3): 513, pi. 4, fig. 13 (9). Female holotype from Chile, possibly in Museum Na- tional d'Histoire Naturelle, Paris, unavailable. Roewer, 1954, Katalog der Araneae, 2: 544. Bonnet, 1958, Bibliographia Araneorum, 2: 3269. Types of this species, type species of Oxysoma, were unfortunately unavailable for examination. Oxysoma cubana Banks Map 5; Figures 105-109 Oxysoma cubana Banks, 1909, Estacion central agronomica de Cuba, Second Report, II ( 2 ) : 157, pi. 10, fig. 7 {$). Male holotype from Havana, Habana, Cuba, in MCZ, examined. Bryant, 1940, Bull. Mus. Comp. Zool., 86: 435, pi. 16, figs. 218, 222, pi. 17, fig. 234, $,9. Kaston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 405. Gaijennina hritcheri Gertsch, 1935, Amer. Mus. Novitates, No. 805: 21, figs. 35, 36 ( ? ). Fe- male holotype from Woods Hole, Massachu- setts, in AMNH, examined. Kaston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 405. Roewer, 1954, Katalog der Araneae, 2: 540. Bonnet, 1957, Bibliographia Araneorum, 2: 1981. Oxysoma cubanum, Roewer, 1954, Katalog der Araneae, 2: 543. Bonnet, 1958, Bibliographia Araneorum, 2: 3268. Diagnosis. The characters of the genus distinguish this species from all other nearctic anyphaenids. The bulbous palp (Fig. 107) and characteristic epigynum (Fig. 106), as well as the color pattern ( Fig. 109 ) , are diagnostic. Variation in this species is discussed above. Male (Suffolk Co., New York). Total length 5.22 mm. Carapace 2.68 mm long, 2.14 mm wide, cephalic width 1.08 mm, clypeus height 0.23 mm, pale yellow with a median dark band and two submarginal longitudinal rows of dark spots. Eyes: diameters (mm): AME 0.05, ALE 0.11, PME 0.09, PLE 0.08; anterior eye row 0.49 mm long, slightly recurved; posterior eye row 0.76 mm long, procurved; MOQ length 0.26 mm, front width 0.19 mm, back width 0.44 mm; eye interdistances (mm): AME-AME 0.06, AME-ALE 0.05, PME- PME 0.26, PME-PLE 0.16, ALE-PLE 0.12. Sternum 1.42 mm long, 0.95 mm wide, pale yellow with translucent border. Che- licerae 0.65 mm long, pale yellow with 3 Spider Family Anyphaenidae • Platnick 261 ft-,--.' vl'^ \ / V i^^^^^^^^ Map 5. Distributions of Oxysoma cubana and Teudis mordax. promarginal and 2 retromarginal teeth. Labium and endites pale yellow. Endites not in^'aginated at middle. Abdomen 2.97 mm long, 1.39 mm wide, pale white with a median longitudinal dark band, venter pale. Epigastric furrow 0.58 mm from tracheal spiracle, spiracle 1.13 mm from base of spinnerets. Legs pale yellow with scattered dark markings, unmodified. Tibial lengths (mm) and indices: I 2.09, 17; II 1.78, 20; III 1.35, 26; IV 2.07, 14. Ventral spination: tibiae I, II 2-2-2, III 1-2-2, IV 2-2-2; metatarsi I, II 2-0-0, III 2-0-2, IV 2-2-2. Palpus as in Figures 105, 107. Female (Barnstable Co., Massachusetts). Coloration as in male. Total length 5.90 mm. Carapace 2.66 mm long, 1.91 mm wide, cephalic width 1.01 mm, clypeus height 0.14 mm. Eyes: diameters (mm): AME 0.06, ALE 0.12, PME 0.09, PLE 0.10; anterior eye row 0.46 mm long, recurved; posterior eye row 0.74 mm long, procurved; MOQ length 0.34 mm, front width 0.19 mm, back width 0.41 mm; eye interdistances (mm): AME- AME 0.05, AME-ALE 0.04, PME-PME 0.25, PME-PLE 0.13, ALE-PLE 0.11. Sternum 1.42 mm long, 0.86 mm wide. Chelicerae 0.86 mm long with teeth as in male. Abdomen 3.71 mm long, 1.71 mm wide. Epigastric furrow 1.19 mm from tracheal spiracle, spiracle 1.13 mm from base of spinnerets. Tibial lengths (mm) and indices: I 1.55, 23; II 1.46, 23; III 1.10, 21; IV 1.42, 18. Ventral spination as in male except tibiae III 0-2-2 and metatarsi IV 0-0-0. Epigynum as in Figure 106, internal genitalia as in Figure 108. Natural history. Mature males have been taken from late May through late August, mature females from late March through late August. One specimen was taken in a pitfall trap, but the habits of this wide- spread but rare species are unknown. Distrihution. Southeastern Arizona to Michigan, Massachusetts, Florida, and Cuba (Map 5). Teudis O. P. -Cambridge Teudis O. P.-Canibridge, 1896, Biologia Centrali Americana, Aran., 1 : 198. Type species Teudis gentilis O. P.-Cambridge, 1896 (= Teudis geminus Petrunkevitch, 1911), designated by F.O. P.-Canibridge, 1900, ihid., 2: 100. Dianibium 3 2(1) Palpal tibia with a distal bulge (Fig. 13); paracymbium pointed at tip (Fig. 13); median apophysis with two long spines (Fig. 14); Europe keyserUngi - Palpal tibia with sides parallel ( Fig. 5); paracymbium rounded at tip (Fig. 5 ) ; median apophysis with short spines (Fig. 6); North America and Europe __.. atrica 3(1) Tcgulum of palpus without projection (other than ctmductor) (Figs. 28, 29); cosmopolitan x-notata - Tegulum of palpu:^ with a projection or sculpturing (other than conductor) (Figs. 42, 56, 68) 4 4(3) Tegulum in "horizontal" position in CN'iiibium, its long axis transverse to that of c>nibium (Fig. 119); projection with teeth "vertical" and surrounding conductor (Fig. 119); paracymbium a hook, barely modified (Figs. 118-120); Japan sia - Tegulum in more or less "vertical" po- sition in cymbium, its long a.xis parallel to cymbium (Fig. 100); projection not vertical 5 5(4) Tegulum projection in ventral view in the shape of a human ear (Fig. 100); paracymbium square (Fig. 101) thorelli - Tegulum with simple projection; para- cymbium not a square (Figs. 68, 75) - 6 6(5) In lateral view tegulum projection al- most as long or longer than tegulum width (Figs. 68, 110) 7 - In lateral view tegulum projection shorter than width of tegulum ( Figs. 75, 86) ._ 8 7(6) Tegulum projection pointed (Fig. 68); paracymbiiun with a dorsally directed point (Fig. 68); Cdliiornia.. ..carpenteri - Tegulum projection truncate ( Fig. 109); paracymbium a ventrally directed lobe (Fig. 110); Europe stroemi 8(6) Palpus with a j,clerite (? terminal apophysis) more or less parallel to em- bolus (in ventral view) in distal part of palpus (Figs. 41, 55); paracymbium complex with a notch (Figs. 47-49, 59) 9 - Palpus with no sclerite parallel to em- bolus (in ventral view) (Figs. 19, 74, 85 ) ; paracymbium without notch (Figs. 20, 75, 86) _-._ 10 9(8) A sclerite (? tenninal apophysis) longer than embolus in \entral \iew ( Fig. 41); paracymbium with a distal notch (Figs. 47-49); eastern Asia, North America - - — - diapar - Terminal apophysis shorter tlian em- bolus in ventral view (Fig. 55); para- cymbium with a ventral notch ( Fig. 59); Europe .montana 10(8) Base of conductor sitting in a depres- sion surrounded by a rim, or base of conductor surrounded by wrinkles (Figs. 75, 85); Eurasia _„ 11 - Base of conductor not surrounded by wrinkles or a rim (Fig. 19); Canary Isl. - - .rninitna 11(10) Base of conductor in a depression sur- rounded by a rim (Fig. 85) kochi 272 BuUctin Museum of Comparative Zoology, Vol. 146, No. 5 — Base of conductor surrounded only by wrinkles of the tegulum (Figs. 74, 75) caspica Key to Females of Zygiella Species 1 Posterior rini of epigynum with a semi- circular lobe (Fig. 71) caspica — Posterior rim otherwise 2 2(1) Epigynum with a scape (Figs. 89, 93, 97, 104, 112) 3 — Epigynum without a scape (Figs. 3, 10, 16, 22, 34, 77, 82) 7 3(2) Openings ventral underneath heart- shaped scape (Fig. 112); Japan sia — Openings posterior 4 4(3) Scape constricted at base (Figs. 89, 93 ) 5 — Scape not constricted at base (Figs. 97, 104) 6 5(4) Scape heart-shaped, slightly longer than wide (Fig. 89) kochi — Scape more than twice as long as wide (Fig. 93); Palestine inconveniens 6(4) Scape a broad lobe with a posterior median extension (Fig. 97) thorelli — Scape much longer than wide with parallel sides (Fig. 104) stroemi 7(2) No depression, openings or sculpturing visible in ventral view of epigynum, at most a posterior rim (Figs. 16, 22); posterior view with two separate open- ings (Figs. 18, 24) 8 — In ventral view a depression, openings or sculpturing visible (Figs. 3, 10, 34, 77, 82); no distinct pair of openings in posterior view 9 8(7) Total length more than 4 mm; epigynum heavily sclerotized (Figs. 22, 24); probably cosmopolitan x-notata — Total length less than 3.5 mm; epigy- num lightly sclerotized (Figs. 16, 18); Canary Isl. minima 9(7) Semicircular openings bordered on ven- ter of epigynum (Fig. 82); Burma nielanocrania — Openings not so; venter of epigynum with a median depression or bulge (Figs. 3, 10, 34, 77) .....10 10(9) A median, posterior, indistinctly bor- dered, dark depression in ventral view of epigynum (Fig. 77); fourth coxae drawn out posteriorly into a spine ( Fig. 80); Malaysia calijptrata — Posterior depression or bulge distinctly bordered (Figs. 3, 10, 34); fourth coxae without a spine; Holarctic region 11 11(10) Median area of epigynum a depression in ventral view much wider than long (Fig. 63); California .carpenteri - Median area at most one and one half times as wide as long (Figs. 3, 10, 34) 12 12(11) Median area of epigynum a bulging lobe framed anteriorly only by a lip (Fig. 10); Europe keyserlingi - Median area depressed, framed an- teriorly and laterally (Figs. 3, 34, 52). .13 13(12) Median depressed area extending pos- teriorly in ventral view (Fig. 3); in posterior view sides of epigynum lightly sclerotized and smaller than median depression (Fig. 4); Europe, North America atrica - Median depressed area not projecting beyond sclerotized area of epigynum in ventral view (Figs. 34, 38, 52); in pos- terior view sides of epigynum heavily sclerotized and sclerotized areas larger in area than median depression (Figs. 35, 39, 54) 14 14(13) Median area with a constriction as seen in botli ventral and posterior views (Figs. 34, 35, 38, 39); Eastern Asia, North America dispar - Median area without constriction as seen in both ventral and posterior views (Figs. 52, 54); Europe montana Zygiella atrica (C. L. Koch) Plate 1; Figures 1-8 EucJuiria atrica C. L. Koch, 1843, Die Arachniden, 12: 103, figs. 1030, 1031, 9, 5. Specimens came from Germany and France and are pre- siunably in the museum of the Humboldt Uni- versitiit, Berlin. Zilla atrica, - Wiehle, 1931, Tierwelt Deutschlands, 23: 33, figs. 38-40, 9, $. Zy fiiclla atrica, - Bonnet, 1959, Bibliographia Araneorum, 2: 4998. Gertsch, 1964, Anier. Mus. Novitates, No. 2188: 16, figs. 18-20, $, ?. Diagnosis. This species can be confused only with Z. keyserlingi. The male differs from other Zygiella by the long palpal tibia and, unlike that of Z. keyserlingi, the tibia has its sides parallel (Figure 5) with setae equally distributed. The epigynum has a wide median lobe extending posteriorly in ventral view; the lobe is depressed in the middle and the lateral sclerotized areas are relatively small (Figures 3, 4). Natural history. This species is common- Orb-weaver Genus Zygiella • Levi 273 Figures 18. Zygiella atrica (C. L. Koch). 1. Female. 2 4. Epigynum. 2. Posterodorsal view, cleared. 3. Ventral. 4. Posterior. 5-7. Left male palpus. 5. Lateral view. 6. Ventral. 7. Expanded. 8. Eye region and chelicerae of female. Figures 9 14. Z. keyserlingi (Ausserer). 9-11. Epigynum. 9. Dorsal, cleared. 10. Ventral. 11. Posterior. 12. Female. 13-14. Male palpus. 13. Lateral. 14. Ventral. Abbreviations. C, conductor; DH, distal hematodocha; E, embolus; M, median apophysis; P, paracymbium; R, radix; S, subtegulum; T, tegulum. Size lines. 0.1 mm, except Figures 1, 8, 12, 1 mm. est on ocean coasts, but is found in Europe in coastal areas of tlie ocean and of Lake also in other locations (Wiehle, 1931), on Erie. On the peninsula of Nahant, Massa- shrubs, junipers, etc. In America the spe- chusetts, it is very common under and be- cies is certainly introduced and is foiuid tween boulders placed to prevent the road 274 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 from being washed away by high tides. As it can be found abundantly among these boulders year after year, the species can presumably tolerate the occasional high waves and salt spray from the ocean. Adult males and females have been found in this location in October. The web (Wiehle, 1931) has more radii (43-50) than that of Z. x-notata and many other orb-weavers; most radii are in the lower half of the web. The free sector is narrow and the hub has a fine mesh. There are many frame threads, many close to the spiral region. The retreat is not as well built as that of Z. x-notata. Emerton's picture of the web ( 1902, The Common Spiders of the United States) shows only a few radii. Distribution. Europe; in America from Nova Scotia to Long Island, New York; Port Credit, Ontario, and British Columbia coast. For map see Gertsch ( 1964 ) . Zygiella keyserlingi (Ausserer) Figures 9-14 Zilla keyserlingi Ausserer, 1871, Verhandl. zool. hot. Ges. Wien, 21: 830, pi. 5, fig. 11, ?. Fe- male holotype from Dalmatia in the Keyserling collection of the British Museum, Natural His- tory, not examined. Wiehle, 1931, Tierwelt Deutschlands, 23: 35, figs. 41, 42, $, $. Zygiella keyserlingi, - Roewer, 1942, Katalog der Araneae, 1: 884. Bonnet, 1959, Bibliographia Araneorum, 2: 5002. Description. Female from unknown lo- cality in Europe. Carapace light brown, cephalic region not much darker. Legs not banded. Dorsum of abdomen with characteristic pattern (Figure 12) and venter with a white line on each side. Diameter of posterior median eyes 0.9 di- ameter of anterior medians, anterior laterals 0.8, posterior laterals 0.7 diameter of an- terior median eyes. Anterior median eyes one diameter apart, one from laterals. Pos- terior median eyes one diameter apart, 1.5 from laterals. There are three teeth on the anterior margin of the chelicerae, three on the posterior, with denticles between the margins. Total length 8.0 mm. Carapace 2.7 mm long, 2.3 mm wide. First femur, 3.1 mm; patella and tibia, 4.0 mm; metatar- sus, 3.1 mm; tarsus, 1.0 mm. Second pa- tella and tibia, 2.9 mm; third, 1.7 mm; fourth, 2.6 mm. Male from unknown locality. Coloration like that of female. Diameter of posterior median eyes and of anterior lateral eyes 0.7 diameter of anterior median eyes; that of posterior lateral eyes 0.6 diameter of an- terior medians. Anterior median eyes slightly less than their diameter apart, and slightly less than their diameter from lat- erals. Posterior median eyes slightly less than one diameter apart, 1.5 diameters from laterals. Total length 6 mm. Cara- pace 2.9 mm long, 2.0 mm wide. First femur, 4.1 mm. Second patella and tibia, 3.6 mm; third, 2.0 mm; fourth, 2.9 mm. Additional female and male specimens were available from Krivosije, Dalmatia. Diagnosis. The long palpal tibia is found also in Z. atrica; however, Z. keyserlingi has the tibia distally swollen with the swollen area having more setae than the basal part ( Fig. 13 ) . The epigynum has a central bulging area bordered anteriorly only by a transverse lip (Figures 10, 11). The species is less pigmented than Z. atrica. Distribution. Portugal, Italy, Hungary to Greece (Bonnet, 1959). Zygiella minima (Schmidt) Figures 15-20 Zygiella x-notata minima Schmidt, 1968, Zool. Beitr., 14: 414, fig. 11, $. Female, male syn- types in poor physical condition from Esperanza Forest, Tenerife, Canary Islands, owned by the author G. Schmidt, but made a\'ailable to me. Description. Female. Coloration diffi- cult to determine. Eyes seem about sub- equal in size. The anterior median eyes slightly less than their diameter apart, their radius from laterals. Posterior median eyes their radius apart, about 0.8 diameter from laterals. Total length 3 mm. Cara- Orb-weaver Genus Zygiella • Levi 275 Figures 15-20. Zygiella minima (Schm\6\). 15-18. Epigynum. 15. Ventral, cleared. 16. Ventral. 17. Posterior, cleared. 18. Posterior. 19-20. Left male palpus. 19. Ventral. 20. Lateral. Figures 21 31. Z. x-notata (Clerck). 21-25. Epigynum. 21. Ventral, cleared. 22. Ventral. 23. Posterior, cleared. 24. Posterior. 25. Dorsal, cleared. 26. Female. 27. Female abdomen, ventral. 28-31. Male palpus. 28. Ventral. 29. Lateral. 30,31. Expanded. 30. Subventral view of bulb. 31. Dorsal view of bulb. Abbreviations. C, conductor; DH, distal hematodocha; E, embolus; I, stipes; M, median apophysis; P, paracym- bium; R, radix; T, tegulum; Y, cymbium. Size lines. 0.1 mm except Figures 26, 27, 1 mm. 276 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 pace 1.5 mm long. First femur, 2.0 mm; patella and tibia, 2.4 mm; metatarsus, 1.8 mm; tarsus, 0.7 mm. Second patella and tibia, 1.7 mm; third, 1.0 mm; fourth, 1.6 mm. Male. In slightly better physical condi- tion than female. Eyes subequal in size. Anterior median eyes their diameter apart, 0.8 diameter from laterals. Posterior me- dian eyes slightly less than their diameter apart, their diameter from laterals. Total length 2.5 mm. Carapace 1.2 mm long, 0.9 mm wide. First femur, 1.6 mm; patella and tibia, 2.1 mm; metatarsus, 1.6 mm; tarsus, 0.7 mm. Second patella and tibia, 1.5 mm; third, 0.8 mm; fourth, 1.2 mm. Diagnosis. Zygiella minima differs from Z. x-notata in that the female has the epig- ynum lightly sclerotized and with differ- ently sized openings in posterior view (Figures 16, 18); the male has a small tooth-shaped projection on the face of the tegulum (Figures 19, 20) absent in Z. x-notata. Distribution. Canary Islands. Zygiella x-notata (Clerck) Figures 21-31, 57-58 Araneiis x-notatus Clerck, 1758, Aranei Svecici, 46, pi. 2, fig. 5. A Clerck specimen bearing this name as labeled by Thorell is in the Swed- ish Museum of Natural History, Stockholm; not examined. Zilla bosenhergi Keyserling, 1878, Verhandl. zool. bot. Ges. Wien, 28; 575, pi. 14, fig. 4, 5, 2 , $. Female and male syntypes from Uruguay in the nmseum of the University of Hamburg and the British Museum (Natural History), exam- ined. NEW SYNONYMY. Zilla caUjornica Banks, 1896, J. New York Ent. Soc, 4: 90. Female holotype from Palo Alto, California, in the Museum of Comparative Zool- ogy, examined. Gertsch (in letter, 1957) in- dicated that Stanford University Museum had specimens marked types. This spider collection has since been sent to the Los Angeles County Museum and was destroyed (C. L. Hogue, per- sonal communication). Larinia maulliana Mello-Leitao, 1951, Rev. Chi- lena Hist. Natur., 51-53: 331, figs. 5, 6, $. Male holotype from Maullin, Chile, in the Museu Nacional, Rio de Janeiro, examined. NEW SYNONYMY. Zygiella x-notata, - Bonnet, 1959, Bibliographia Araneorum, 2: 5007. Gertsch, 1964, Amer. Mus. Novitates, No. 2188, 12, figs. 2, 15-17, 2,5, map. Diagnosis. The epigynum, unlike that of Z. minima, is heavily sclerotized. It has two diagnostic openings seen in posterior view (Figure 24). The palpus is simple with the tegulum's long axis parallel to that of the cymbium (Figures 28-30); the lack of terminal apophysis (Figures 30, 31) separates males from those of other species, the lack of a tegulum projection from males of Z. rninima. Natural history. Numerous references to habits and webs can be found in Bonnet (1959). The web, which has a vacant sector, has been used in cthological studies. It is illustrated in Wiehle (1931), figure 37, and J. Comstock, 1940, The Spider Book, figure 470. The species is very common in southern Chile. In the city park of Osorno, Chile, I found suspended from a web on a telephone pole a dried, shrivelled, 6-cm long lizard on which a Z. x-notata had apparently fed (15 March 1965). Figures 32-50. Zygiella dispar (Kulczynski). 32-39. Epigynum. 32, 33. Posterior view, cleared. 34, 36, 38. Ventral. 35, 37, 39. Posterior. 32. (Michigan). 33-35. (Mendocino Co., California). 36, 37. British Columbia. 38, 39. (Virginia). 40-50. Left male palpus. 40. Expanded. 41. Ventral. 42. Lateral. 43-46. Embolus and apophysis. 43. (Alaska). 44. (California). 45. (Manitoba). 46. (Maine). 47-49. Paracymbium. 47. (Alaska). 48. (California). 49. (Manitoba). 50. (Maine). Figures 51-56. Zygiella montana (C. L. Koch). 51-54. Epigynum. 51. Anterodorsal view, cleared. 52. Ventral. 53. Posterior, cleared. 54. Posterior. 55, 56. Male palpus. 55. Ventral. 56. Lateral. Abbreviations. A, terminal apophysis; C, conductor; E, embolus; H, hematodocha; I, stipes; M, median apophysis; T, tegulum. Scale lirtes. 0.1 mm. Orb-weaver Genus Zygiella • Levi 277 SSSrSfe'Wiv.i 278 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 Adult males are found from July until September on the Pacific coast of North America. Distribution. Europe, but probably cos- mopolitan, carried around the world by man. It is introduced in America and found along the Atlantic coast from Maine to Virginia, the Pacific coast from southern British Columbia to southern California. Gertsch ( 1964 ) maps the North American distribution. It is very common in Chile and is found in Uruguay and Argentina. Zygiella dispar (Kulczyhski) Figures 32-50 Zilla dispar Kulczynski, 1885, Denkschrift. Akad. Wissenschaften Krakow, 11: 24, pi. 9, fig. 7, 5, $ . Male type from Kanitchatka, Siberia, in Polish Academy of Sciences, Warsaw, in poor physical condition, examined. Ztjgic'IIa montana, - numerous authors of American records only. Zygiella dispar, - Gertsch, 1964, Amer. Mus. Novi- "tates. No. 2188: 7, figs. 7-10, 9, $. Zygiella nearctica Gertsch, 1964, Amer. Mus. Novitates, No. 2188: 4, figs. 3-6, 9, $. Male holotype from Seba, Alberta, in the American Museum of Natural History, not examined. NEW SYNONYMY. Note. Gertsch (1964) used the name dispar for the population along the Pacific coast from Alaska to south-central Cali- fornia; other specimens he called nearctica. Gertsch separated Z. nearctica from Z. dis- par by the following characters: the male palpus has the apical [= ? subterminal] apophysis less developed, and has "differ- ences of the various apophysis"; the female epigynum has the "fovea" visible from be- low. The last character is a matter of po- sition of the epigynum during examination. Gertsch's figure 7 (dispar) is much more characteristic of all specimens of the spe- cies in ventral view than is figure 4 (nearc- tica), which is the view from slightly pos- terior. The subterminal apophysis differs among individuals ( Figures 43-46 ) , as do the paracymbium (Figures 47-50) and, to a lesser extent, the median apophysis (not illustrated). Similarities of the internal female genitalia also indicate that we have only one species, not two. California specimens of the species are the largest; a male from Alaska was the smallest speci- men examined. Gertsch is probably correct in stating that Z. dispar is distinct from Z. montana of Europe. Perhaps intermediates will be found in the vast area between Europe and Siberia from which no collections have been examined, but I would not expect this. Diagnosis. Females of Z. dispar differ from those of the related Z. montana in that the median depression of the epigy- num has a consti^iction (Figures 35, 39) in posterior view. The palpus is similar to that of Z. monta)Ui but differs in that most sclerotized parts of the palpus have a different shape and are positioned slightly differently (Figures 41, 42). Natural history. The species is found on trees and rocks (Emerton, 1902, The Com- mon Spiders, p. 185). Parts of the web have been illustrated by Emerton (1884, pi. 40, fig. 2). Emerton collected the species in the Adirondack Mountains, New York State, and the White Mountains, New Hampshire. Distribution. Kamtchatka, Siberia, and North America along the Pacific coast, across Canada, the northern states, south in the western states, and in the Appala- chian mountains in the east ( Gertsch, 1964, fig. 1, a map ) . Zygiella montana (C. L. Koch) Figures 51-56, 59-61 Zilla montana G. L. Koch, 1839, Die Arachniden, 6: 146, pi. 536, 537, 9, $. Syntypes probably from Nassfelder Alpen in Salzburg, Austria, in the Museum of the Humboldt University, Ber- lin, not examined. Wiehle, 1931, in Dahl, Tier- welt Deutschlands, 23: 38, figs. 46-48, 9, $. Zygiella montana, - Roewer, 1942, Katalog der Araneae, 1: 886. Bonnet, 1959, Bibliographia Araneorum, 2: 5003. Description. Female from Seefeld, Tirol, Austria. Coloration as in other species. Orb-weaver Genus Zygiella • Levi 279 Figures 57-58. Zygiella x-notata (Clerck). 57. Eye region and chelicerae. 58. Left chelicera from inside. Figures 59-61. Z. montana (C. L. Koch). 59. Paracymbium dorsolateral view. 60 61. Left male palpus, ex- panded. 60. Subventral. 61. Dorsal, cymbium removed. Figures 62 69. Z. carpenter! Archer. 62-64. Epigynum. 62. Posterior, cleared. 63. Ventral. 64. Posterior. 65. Female. 66. Female abdomen, ventral 67-69. Male palpus. 67. Ventral. 68. Lateral. 69. Paracymbium. Abbreviations. A, terminal apophysis; C, conductor; E, embolus; M, median apophysis; P, paracymbium; R, radix; T, tegulum. Scale lines. 0.1 mm, except Figures 57, 65, 66, 1.0 mm. 280 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 Secondary eyes 0.8 diameter of anterior medians. Anterior median eyes 0.7 diam- eter apart, 0.6 diameter from laterals. Pos- terior median eyes one diameter apart, 1.2 from laterals. Total length 8.0 mm. Cara- pace 2.9 mm long, 2.2 mm high. First femur, 3.0 mm; patella and tibia, 3.7 mm; metatarsus, 2.9 mm; tarsus, 1.2 mm. Second patella and tibia, 3.1 mm; third, 1.9 mm; fourth, 2.6 mm. Male from Seefeld, Tirol, Austria. Sec- ondary eyes 0.6 diameter of anterior me- dians. Anterior median eyes 0.6 diameter apart, 0.5 diameter from laterals. Posterior median eyes their diameter apart, 1.6 from laterals. Total length 6.5 mm. Carapace 3.0 mm long, 2.4 mm wide. First femur, 3.2 mm; patella and tibia, 5.0 mm; metatar- sus, 4.3 mm; tarsus, 1.0 mm. Second pa- tella and tibia, 3.9 mm; third, 2.3 mm; fourth, 3.2 mm. Diagnods. This European species can easily be confused with Z. dispar but the epigynum lacks the constriction of the me- dian depression in posterior view (Figure 54; the palpus has many sclerites, all slightly different in shape (Figures 55, 56). Natural history. According to Wiehle (1931) this is a mountain species found in the Alps above 1000 m elevation, most commonly between 1300-1800 m. The species is found on buildings, rocks, bark and branches of trees and shrubs. The web is similar to that of Z. x-notata with 19-35 radii. The vacant sector is especially wide and the hub has a rough structure. Both sexes are mature from June until September, and may take several years to mature. Distribution. European mountains. Zygiella carpenter! Archer Figures 62-69 Zygiella carpenteri Archer, 1951, Anier. Mus. Novitates, No. 1487: 18, fig. 34, 9. Female holotype from Del Monte Forest, Pacific Grove, Monterey Co., California, in the American Mu- seum of Natural History, examined. Gertsch, 1964, Amer. Mus. Novitates, No. 2188: 9, figs. 1, 11-14, 9, 5, map. Diagnosis. The wide depression of the epigynum (Figure 63), the long, pointed projection of the palpal tegulum (Figure 68) and the shape of the paracymbium (Figures 68, 69) separate the species from Z. dispar. Distribution. Sierra mountains of Ore- gon and Washington. There are also a few records from near Spokane, Washing- ton, and the coast of California. There is a distribution map in Gertsch (1964). Zygiella caspica (Simon) Figures 70-75 Zilla caspica Simon, 1889, Verh. zool. bot. Ges. Wien, 39: 382. Two female, one male syntypes from Transylvania in the Museum National d'Histoire Naturelle, Paris, examined. Zi/fiiella caspica, - Roewer, 1942, Katalog der Araneae, 1: 883. Bonnet, 1959, Bibliographia Araneorum, 2: 5002. Description. Female. Color like that of other species. Legs not banded, yellowish brown. Dorsal pattern as is characteristic in Zygiella (Figure 70), venter with very Figures 70-75. Zygiella caspica {S\mon). 70. Female. 71,72. Epigynum. 71. Ventral. 72. Posterior. 73-75. Left male palpus. 73. Mesal. 74. Ventral. 75. Lateral. Figures 76-80. Z. calyptrata (Workman). 76-78. Epigynum. 76. Dorsal, cleared. 77. Ventral. 78. Posterior. 79. Female. 80. Fourth coxae, ventral. Figures 81 84. Z. melanocrania (Thorell). 81-83. Epigynum. 81. Ventral, cleared. 82. Ventral. 83. Posterior. 84. Female. Figures 85-86. Z. kocfii (Thorell), male palpus. 85. Ventral. 86. Lateral. Scale lines. 0.1 mm except Figures 70, 79, 84, 1.0 mm. Orb-weaver Genus Zygiella • Levi 281 .:fc-|^.^ vv 282 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 little black pigment. The posterior median eyes are slightly smaller than anterior me- dians, laterals 0.8 diameter of anterior me- dian eyes. The anterior median eyes are their radius apart, their radius from lat- erals. Posterior median eyes their diameter apart, and slightly more than one diameter from laterals. Total length 6.5 mm. Cara- pace 2.4 mm long, 1.9 mm wide. First fe- mur, 2.8 mm; patella and tibia, 3.6 mm; metatarsus, 2.6 mm; tarsus, 0.9 mm. Second patella and tibia, 2.8 mm; third, 1.7 mm; fourth, 2.3 mm. Male. Coloration as in female. The eyes are slightly larger and closer together. Total length 5.0 mm. Carapace 2.3 mm long, 1.7 mm wide. First femur, 2.9 mm; patella and tibia, 4.3 mm; metatarsus, 2.9 mm; tarsus, 1.2 mm. Second patella and tibia, 2.9 mm; third, 1.7 mm; fourth, 2.3 mm. Diagnosis. While the short semicircular scape of the epigynum (Figures 71, 72) is distinct, the palpus is similar to that of Z. kochi, but differs in the shape of the tegu- lum at the base of the conductor and the terminal apophysis (Figures 73-75). Distribution. Trans-Carpathian region. Zygiella calyptrata (Workman) Figures 76-80 Epeira calyptrata Workman, 1894, Malaysian Spi- ders, p. 21, plate 21. One female lectotype here designated and two female paralectotypes from Singapore in the National Museum of Ireland, Dublin, examined. Epeira (Zilla) calyptrata, - Thorell, 1895, Descr. Catalogue of the Spiders of Burma, p. 188. Zygiella calyptrata, - Roewer, 1942, Katalog der Araneae, 1 : 886. Araneus cahjptratus, - Bonnet, 1955, Bibliogra- phia Araneorum, 2: 450. Description. Female lectotype. Carapace brown; head region very much darker, glossy. Sternum, legs brown. Dorsum of abdomen white with black marks (Figure 79). Sides brownish black. Venter gray. Anterior median eyes much larger than others. Diameter of posterior median eyes 0.8 diameter of anterior medians; laterals 0.6 diameter of anterior median eyes. An- terior median eyes their diameter apart, slightly more than their diameter from lat- erals. Posterior median eyes slightly less than their radius apart, 2.5 diameters from laterals. The fourth coxa has a posterior distal spine (Figure 80). Total length 4 mm. Carapace 1.7 mm long, 1.3 mm wide. First femur, 1.5 mm; patella and tibia, 2.0 mm; metatarsus, 1.5 mm; tarsus, 0.6 mm. Second patella and tibia, 1.7 mm; third, 1.0 mm; fourth, 1.5 mm. Diagnosis. Unlike females of Z. melano- crania, those of Z. calyptrata have the me- dian area of the epigynum dark with an indistinct border ( Figure 77 ) . It is doubt- ful that this species belongs to Zygiella and is related to the other Zygiella species. Distribution. Malaysia, Burma. (Of ThorelFs specimens labeled Epeira calyp- trata in the British Museum, Natural His- tory, one is this species, the other specimen is a related species.) Zygiella melanocrania (Thorell) Figures 81- 84 Epeira melanocrania Thorell, 1887, Ann. Mus. Civica Storia Xatur. Genova, (2)5: 209. Fe- male holotvpe from Teinzo, Burma, in the Museo Civico di Storia Naturale, Genova, exam- ined. Zygiella melanocrania, - Roewer, 1942, Katalog der Araneae, 1 : 886. Aranetis melanocranius, - Bonnet, 1955, Bibliog- raphia Araneorum, 2: 543. Description. Carapace shiny brown. Head region dark brown. Chelicerae brown, darker than head region. Sternum yellow-brown. Legs brown, first two darker than last two, with faint indications of darker rings. Dorsum of abdomen with characteristic black and white Zygiella pat- tern (Figure 84). Venter with white pig- ment spots only. Secondary eyes 0.8 diam- eter from anterior median eyes. Anterior median eyes are a diameter apart, slightly more than one diameter from laterals. Pos- terior median eyes are their radius apart, 2.5 diameters from laterals. The laterals are separated by their radius from each Orb-weaver Genus Zygiella • Levi 283 other. The heiglit of tlie clypcus is about equal to the radius of the anterior median eyes. The chehcerae of one specimen have four teeth on the anterior margin of tlie fang furrow, but the posterior margin has four on one cheHcera, three on the other; there are denticles in the furrow. The ab- domen is oval and hairy. Total length 5.5 mm. Carapace 2.6 mm long, 2.0 mm wide. First femur, 2.7 mm; patella and tibia, 3.3 mm; metatarsus, 2.1 mm; tarsus, 0.9 mm. Second patella and tibia, 2.7 mm; third, 1.6 mm; fourth, 2.2 mm. Diai:,nosis. The epigynum (Figures 82, 83), with two semicircular openings on the ventral side, separates this species from all other known Zygiella. Distribution. This species is known only from the type specimen. The specimen il- lustrated by Dyal ( 1935, Bull. Dept. Zool, Panjab Univ. 1: 183, pi. 16, fig. 125) is probably not this species. Zygiella kochi (Thorell) Figures 85-91 Zilla kochii Thorell, 1870, Remarks on Synonyms of European Spiders, p. 33. Synt>'pes from Nice and Monaco presumably in the Stockholm Nat- ural History Museum. Rosenberg, 1901, Zool- ogica, 13: 43, pi. 3. fig. 32, 9, $. Wiehle, 1929, Z. Morphol. Okol. Tiere, 15: 262-308. Wiehle, 1931, in Dahl, Tierwelt Deutschlands, 23: 41, figs. 52, 53, 9, $. Zygiella koclii, - Simon, 1929, Arachnides de France, 6(3): 663, 754, figs. 1021, 1025, 9, $. Roewer, 1942, Katalog der Araneae, 1: 884. Ronnet, 1959, Ribliographia Araneoiinn, 2: 5002. Description. Female from France. Cara- pace brown, with darker lines going from eye region to thoracic depression ( Figure 87). Sternum brown. Legs very indis- tinctly banded. Dorsum of abdomen with usual pattern ( Figure 87 ) . Venter with a black spot framed by white on each side. Secondary eyes 0.7 diameter of anterior medians. Anterior median eyes 0.7 diam- eter apart, one diameter from laterals. Pos- terior median eyes one diameter apart, a little less than two diameters from laterals. Total length 7.5 mm. Carapace 3.5 mm long, 2.5 mm wide. First femur, 3.2 mm; patella and tibia, 4.3 mm; metatarsus, 3.0 mm; tarsus, 1.3 mm. Second patella and tibia, 3.2 mm; third, 2.0 mm; fourth, 2.9 mm. The entrance into the seminal recep- tacles is through pockets and folds rather than through distinct ducts (Figures 88, 91). Description of male from unknown lo- cality. Posterior median eye diameter about the radius of anterior median eyes; anterior lateral eyes 0.7 diameter of anterior median eyes; posterior lateral eye diameter about the radius of anterior median eyes. An- terior median eyes their radius apart and about their radius from laterals. Posterior median eyes their diameter apart, 1.5 di- ameters from laterals. There are no modi- fications on appendages. Total length 7 mm. Carapace 3.1 mm long, 2.3 mm wide. First femur, 3.2 mm; patella and tibia, 4.7 mm; metatarsus, 3.1 mm; tarsus, 1.3 mm. Second patella and tibia, 3.6 mm; third, 2.0 mm; fourth, 2.6 mm. Diagnosis. The heart-shaped scape of the epigynum (Figure 89) separates this species readily from others. The scape has a central depression. The rim on the tegu- lum surrounding the base of the conductor, and the shape of the subterminal apophysis of the palpus separate males from Z. cas- pica (Figures 85, 86). Natural histonj. The species is found on triuiks of trees, cork bark and chestnut in Corsica; its retreat is in cracks in bark (Wiehle, 1929, 1931). The web is simihir to that of Z. x-notata; of fifteen webs four did not have the vacant sector but had complete orbs (Wiehle, 1929). Distribution. Central and southern Europe, Mediterranean region and North Africa (Bonnet, 1959). Zygiella inconveniens (O. P. -Cambridge) Figures 92-94 Epeira inconveniens (O.P.-Cambridge), 1872, Proc. Zool. Soc. London, p. 298. Female holo- 284 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 type and juvenile lectotype from Beirut, Leba- non. Zijgiella inconveniens, - Roewer, 1942, Katalog der Araneae, 1: 883. Araneiis inconveniens, - Bonnet, 1955, Bibliog- raphia Araneorum, 2: 522. Description. Coloration characteristic for the genus ( Figure 92 ) . The secondary eyes are about 0.8 diameter of anterior medians. Anterior median eyes are 0.7 diameter apart, their radius from laterals. The posterior median eyes are slightly less than one diameter apart, slightly more than one from laterals. Total length 5.5 mm. Carapace 2.5 mm long, 1.9 mm wide. First femur, 2.3 mm; patella and tibia, 3.2 mm; metatarsus, 2.4 mm; tarsus, 0.9 mm. Second patella and tibia, 2.5 mm; third, 1.5 mm; fourth, 2.2 mm. Diagnosis. Females differ from Z. kochi in the longer, narrower scape of the epigy- num ( Figure 93 ) . Distribution. Only known from Beirut, Lebanon. Zygiella thorelli (Ausserer) Figures 95-101 Zilla thorelli Ausserer, 1871, Verhandl. zool. bot. Ges. Wien, 21: 830, pi. 5, fig. 10, ?. Female from Prater (amusement park), Vienna, Aus- tria, probably in the Naturhistorisches Museum, Wien, not examined. Wiehle, 1931, in Dahl, Tierwelt Deutschlands, 23: 39, figs. 49-51, ?, $. Zygiella thorelli, - Simon, 1929, Arachnides de France, 6(3): 663, 664, 755, figs. 1019, 1024, 9 S. Roewer, 1942, Katalog der Araneae, 1: 884. Bonnet, 1959, Bibliographia Araneorum, 2: 5006. Description. Female from France. Cara- pace brown, black lines from each pos- terior lateral eye to thoracic region, fusing there with a lateral branch; black line around margin of the thoracic region ( Fig- ure 95). Chelicerae dark brown. Sternum dark brown with light brown median longi- tudinal narrow band. Legs brown with narrow dark bands. Dorsum with charac- teristic pattern (Figure 95) containing black and with white pigment spots. Ven- ter black between genital furrow and spin- nerets, with a white line on each side. Pos- terior median eyes 0.6 diameter of anterior median eyes. Anterior lateral eyes 0.7 di- ameter of anterior medians, posterior lat- eral eyes 0.5 diameter of anterior median eyes. Anterior median eyes 0.7 diameter apart, one diameter from laterals. Posterior median eyes slightly less than their diam- eter apart, 1.7 from laterals. On the anterior margin of tlie fang furrow, the chelicerae have three large teeth; on the posterior margin, four teeth and, farthest from fang, a denticle. Total length 10 mm. Carapace 4.5 mm long, 3.2 mm wide. First femur, 4.9 mm; patella and tibia, 6.7 mm; metatarsus, 5.0 mm; tarsus, 1.7 mm. Second patella and tibia, 5.0 mm; third, 2.9 mm; fourth, 4.0 mm. Male from Kochem on the Mosel, Ger- many. Coloration like that of female. Sec- ondary eye diameter 0.6 diameter of an- terior median eyes. Anterior median eyes slightly less than their radius apart, their diameter from laterals. Posterior median eyes 0.7 diameter apart, 1.5 diameters from laterals. The chelicerae have three teeth on the anterior margin; three smaller teetli on the posterior. Total length 7.5 mm. Carapace 3.9 mm long, 2.8 mm wide. First femur, 4.8 mm; patella and tibia, 7.0 mm; metatarsus, 6.5 mm; tarsus, 1.8 mm. Second patella and tibia, 5.0 mm; third, 2.8 mm; fourth, 3.5 mm. Diagnosis. This species has a longer, narrower carapace than is seen in other species of Zygiella. Females are distinct in the shape of the epigynal scape, a lobe with a distal extension ( Figure 97 ) . Males are characterized by the sculptured, hu- man-ear-shaped tegulum (Figure 100). No other known species is close to Z. thorelli. Natural history. This central European species prefers warm locations such as walls of ruins and cliffs. It has also been found on wooden buildings. The sexes are mature in August and September (Wiehle, 1931). The web, a typical Zygiella web, Ohb-weaver Genus Zygiella • Levi 285 Figures 87 91. Zygiella kochi {ThoreW). 87. Female. 88 91. Epigynum. 88. Dorsal, cleared. 89. Ventral. 90. Posterior. 91. Posterior, cleared. Figures 92-94. Z. inconveniens (O. P. -Cambridge). 92. Female. 93, 94. Epigynum. 93. Ventral. 94. Posterior. Figures 95 101. Z. thorelli {Ausserer). 95. Female. 96 99. Epigynum. 96. Dorsal, cleared. 97. Ventral. 98. Posterior. 99. Posterior, cleared. 100, 101. Left male palpus. 100. Ventral. 101. Lateral. Scale lines. 0.1 mm except Figures 87, 92, 95, 1.0 mm. 286 Bitlletin Museum of Comparative Zoology, Vol. 146, No. 5 is pictured in Lendl ( 1891, Potpiiz Tennesz. kozl., Budapest, 13: 31, figure 8). Distribution. France, southern Ger- many, Czechoslovakia, Pohmd to Italy and Roumania (Bonnet, 1959). Zygiella stroemi (Thorell) Figures 102-110 Zilla stroemi Thorell, 1870, Remarks on Synonyms of European Spiders, p. 235. New name for Zilla montana, Westring (not C. L. Koch) from Sweden. Wiehle, 1931, in Dahl, Tierwelt Deutschlands, 23: 36, figs. 43^5, 9, $. Zygiella x-notata, - Roewer, 1942, Katalog der Araneae, 1: 884 (not x-notata Clerck). Zygiella stroemi, - Locket and Millidge, 1953, British Spiders, 2: 163, figs. 108b, 109c, 9, $. Bonnet, 1959, Bibliographia Araneorum, 2: 5005. Description. Female from Plitvice, Cro- atia, Jugoslavia. Coloration similar to that of other species (Figure 103). Diameter of posterior median eyes 0.8 diameter of anterior medians. Anterior lateral eyes 0.9 diameter of anterior medians and posterior lateral eyes 0.8 diameter of anterior me- dians. Anterior median eyes slightly less than their radius apart, the same distance from laterals. Posterior median eyes their diameter apart, slightly more than their diameter from laterals. Total length 4.5 mm. Carapace 1.9 mm long, 1.5 mm wide. First femur, 2.2 mm; patella and tibia, 2.7 mm; metatarsus, 2.0 mm; tarsus, 0.9 mm. Second patella and tibia, 1.9 mm; third, 1.3 mm; fourth, 1.9 mm. Male from Plitvice, Croatia, Jugoslavia. Diameter of secondary eyes 0.7 diameter of anterior medians. Anterior medians 0.3 diameter apart, the same distance from lat- erals. Posterior median eyes slightly less than their diameter apart, slightly more than their diameter from laterals. Total length 3.4 mm. Carapace 1.7 mm long, 1.6 mm wide. First femur, 2.2 mm; patella and tibia, 3.0 mm; metatarsus, 2.6 mm; tar- sus, 1.0 mm. Second patella and tibia, 2.2 mm; third, 1.1 mm; fourth, 1.5 mm. Diagnosis. The flat, long scape with al- most parallel sides (Figure 104) separates females from all other Zygiella. Males are \ distinguished by the truncate projection of the tegulum of the palpus ( Figures 108- 110). Natural historij. The web is on the trunks of pines (Wiehle, 1931; Locket and Millidge, 1953); the retreat is under bark, Wiehle ( 1931 ) reports that the species ma- tures from May until June, and several specimens may be found near each other. Distribution. Most of Europe to Siberia and Turkestan (Bonnet, 1959). Zygiella sia (Strand) Figures 111-120 Aranea (Zilla) sia Strand, 1906, in Bosenberg and Strand, Abhandl. Senckenberg. Ges., 30 (1-2): 237, pi. 4, fig. 24, $. Adult female, male, and 5 juvenile syntypes from Japan in the Senckenberg Museum, Frankfurt, examined. Zygiella sia, - Roewer, 1942, Katalog der Araneae, 1: 884. Araneus sia, - Bonnet, 1955, Bibliographia Araneorum, 2: 598. Yaginuma, 1960. Spiders of Japan in Colour, Osaka, p. 54, figs. 1, 3, plate 19, fig. 115, ?, $. Zilla sia, - Saito, 1959, The Spider Book Illus- trated in Colours, Tokyo, p. 109, fig. 23, pi. 17, fig. 129 a, b, pi. 18, fig. 129 d, ?, web. Description. Female syntype. Carapace brown, head region darker brown, darker area coming to a point posteriorly in tho- racic depression. Some white hairs on sides. Sternum dark brown. Legs indistinctly to distinctly banded. Abdomen with the characteristic pattern. Venter with a white longitudinal line on each side. Posterior median eyes 0.6 diameter of anterior me- dians, anterior lateral eyes 0.6, posterior laterals 0.5 diameter. Anterior median eyes 0.8 diameter apart, 1.5 diameters from lat- erals. Posterior median eyes 0.7 diameter apart, 3.0 diameters from laterals. Lateral eyes slightly separated. There are three teeth on the anterior margin of chelicerae. Total length 7 mm. Carapace 2.7 mm long, 2.2 mm wide. First femur, 2.9 mrn; patella and tibia, 4.0 mm; metatarsus, 2.7 mm; tarsus, 0.7 mm. Second patella and tibia, 3.4 mm; third, 1.9 mm; fourth, 2.7 mm. Orb-weaver Genus Zygiella • Levi 287 Figures 102 110. Zygiella stroemi (ThoreW). 102. Eye region and chelicerae. 103. Female. 104 106. Eplgy- num. 104. Ventral. 105. Anterodorsal, cleared. 106. Posterior. 107-110. Left male palpus. 107. Expanded. 108. Mesal. 109. Ventral. 110. Lateral. Abbreviations. A, terminal apophysis; C, conductor; E, embolus; \, stipes; M, median apophysis; R, radix; SA, subterminal apophysis; T, tegulum. Scale lines. 0.1 mm except Figures 102, 103, 1.0 mm. Male syntype from Japan. Coloration as in female, but abdominal pattern more dis- tinct. Diameter of secondary eyes about equal to radius of anterior medians. Pos- terior lateral eyes slightly smaller than other secondary eyes. Anterior median eyes 0.7 diameter apart, one diameter from laterals. Posterior median eyes 0.6 diam- eter apart, three diameters from laterals. There are three teeth anteriorly on chelic- eral fang margin and three posteriorly. Total length 6 mm. Carapace 2.6 mm long, 2.1 mm wide. First femur, 3.0 mm; patella and tibia, 4.1 mm; metatarsus, 2.9 288 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 Figures 111-120. Zygiella sia (Strand). 111-115. Epigynum. 111. Ventral, cleared. 112. Ventral. 113. Ven- tral, scape torn off. 114. Posterior, cleared. 115. Posterior. 116. Female. 117-120. Left male palpus. 117, 118. Expanded. 119. Ventral. 120. Lateral. Abbreviations. A, terminal apophysis; C, conductor; E, embolus; H, hematodocha; M, median apophysis; P, para- cymbium; R, radix; T, tegulum; TA, projection of tegulum. Scale lines. 0.1 mm except Figure 116, 1.0 mm. Ohb-weaver Genus Zygiella • Levi 289 mm; tarsus, 1.0 mm. Second patella and tibia, 3.3 mm; third, 1.8 mm; fourth, 2.6 mm. Note on size. Several other specimens of this species were examined. They were much larger. And with the size increase there was a proportionate increase in the distance of the laterals from the median eyes. A female from Shiga Prefecture was 12.5 mm total length. The specimen had a carapace 4.7 mm long and 4.1 mm wide, about 1.7 times the size of the female syn- type. The legs were of proportionate length, 1.7 times that of the syntype. The comparative eye sizes stayed the same but anterior median eyes were about twice their diameter from laterals (a distance in- crease of about 1.3 times) and the posterior medians slightly less than five times from laterals (a distance increase of 1.6 times almost proportionate to growth ) . The eyes thus grew relatively less. Male specimens from Naga Prefecture were also larger: total length 7.5 and 10.5 mm; carapace 3.8 and 5.1 mm long, 2.7 and 3.9 mm wide. These measurements are 1.4 times and 2.0 times the corresponding mea- surements of the syntype; the appendage articles were, however, 1.7 times and 2.2 times the length of the carapace of the type. Growth of males' legs thus did not seem proportional. However, in the two different-sized males from Naga Prefec- ture, carapace and leg sizes were in pro- portion. The Naga males had the diameter of the secondary eyes 0.7 diameter of the medians (the syntypes about 0.5 diameter). The syntype had the anterior median eyes one diameter from laterals, the smaller Naga specimen one and one-half, the larger one slightly less than two. The posterior me- dian eyes were three diameters from lat- erals in the syntype, about four in the smaller Naga specimen (1.3 times the dis- tance), about five times in the larger one ( 1.7 times the distance in tlie syntype). The eye distances increase less than size; there appears to be only little increase in eye sizes. Presumably the specimens had matured in different instars. But tliesc proportional differences seem surprising considering the similarity in proportion and size of the epigyna and male palpi. The male specimen whose carapace was twice as long as the carapace of the syn- type, also had the palpal tibia 2.5 times as long as that of the syntype (a propor- tional increase with leg length), but the critical palpal cymbium was only 1.4 times longer than that of the syntype. The larger specimen thus had relatively a much longer palpal tibia. No differences were noted in the position and proportion of the sclerites held within the cymbium. Diagnosis. The heart-shaped scape cov- ering the ventral openings of the epigynum (Figure 112) separates the female from all other Zygiella. The scape has a transverse light mark. The male palpus (Figures 119- 120) is superficially very different from other species: it has a huge basal hema- todocha, a minute tegulum bearing a toothed projection, and the median apoph- ysis has a projecting hook (Figures 117- 118). As in Z. atrica the palpal tibia is slightly elongated, but of a different shape. There is some doubt in placing this species in Zygiella, because of the wider spacing of the eyes and the cap on the pal- pal embolus in the expanded palpus (Fig- ure 117), not otherwise seen in the genus. The course of the duct into and through the tegulum remains uncertain, despite its having been illustrated in Figures 117, 118. Distribution. Japan. Fox ( 1938, J. Wash- ington Acad. Sci. 28: 367) reported speci- mens from Szechwan Prov. China, but the specimens of the U.S. National Museum could not be found. REFERENCES CITED Archer, A. 1951a. Studies in the orbweaving spiders ( Argiopidae), 1. Amer. Mus. Novi- tates, No. 1487: l-,52. . 1951b. Studies in the orbweaving spi- 290 Bulletin Museum of Comparative Zoology, Vol. 146, No. 5 ders ( Argiopidae), 2. Amer. Mus. Novitates, No. 1502: 1-34. Bonnet, P. 1959. Bibliographia Araneorum, Vol. 2. Imprimerie Douladoure, Toulouse, vol. 2, 4231-5058. Emerton, J. H. 1884. New England spiders of the family Epeiridae. Trans. Connecticut Acad. Arts Sci., 6: 295-342. Gertsch, W. J. 1964. The spider genus Zy- giella in North America (Araneae, Argiop- idae). Amer. Mus. Novitates, No. 2188: 1-21. Roewer, C. F. 1942. Katalog der Araneae. Verlag Natura, Hamburg, vol. 1. Simon, E. 1895. Histoire Naturelle des Araig- nees. Roret, Paris: Libraire Encyclopedique vol. 1. Wiehle, H. 1931. Araneidae. In Dahl, F. Die Tierwelt Deutschlands, G. Fischer Verlag, Jena 23: 1-136. Valid names are printed in italics, illustrations. alpina, Zilla 270 ancora, Epeira 270 atrica, Eucharia 272 atrica, Zilla 272 atrica, Zijgiella 269*, 272, 273* aureola, Zilla 270 bosenbergi, Zilla 276 californica, Zilla 276 calophylla, Aranea 271 calyptrata, Epeira 282 calyptrata, Zygiella 281*, 282 calyptratus, Araneus 282 carpenteri, Zygiella 279*, 280 caspica, Zilla 280 caspica, Zygiella 280, 281* crucinotata, Zilla 271 decolorata, Zilla 271 dispar, Zilla 278 dispar, Zygiella 277*, 278 gigans, Zilla 271 guttata, Zilla 271 guyanensis, ZiUa 271 inconveniens, Araneus 284 inconveniens, Epeira 283 inconveniens, Zygiella 283, 285* keyserlingi, Zilla 274 keyserlingi, Zygiella 273*, 274 kochii, Zilla 283 INDEX Page numbers refer to main references, starred page numbers to kochi, Zygiella 281*, 283, 285* maulliana, Larinia 276 melanocephala, Linyphia 271 melanocrania, Epeira 282 melanocrania, Zygiella 281*, 282 melanocranius, Araneus 282 minima, Zygiella 274, 275* montana, Zilla 278 montana, Zygiella 211*, 278, 279" montana, Zygiella 278 nawazi, Zilla 271 nearctica, Zygiella 278 punctata, Zilla 271 rogenhoferi, Zilla 271 sia, Aranea 286 sia, Araneus 286 sia, Zilla 286 sia, Zygiella 286, 288* stroemi, Zilla 286 stroemi, Zygiella 286, 287* thorelli, Zilla 284 thorelli, Zygiella 284, 285* x-notata, Zygiella 275*, 276, 279* x-notata, Zygiella 286 x-notatus, Araneus 276 Zygia 268 Zygiella 268 us ISSN 0027-4100 Sulletin OF THE Museum of Compardtive Zoology The Orb-weaver Genera Ataniella and Nuctenea (Araneae: Araneidae) HERBERT W. LEVI HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS, U.S. A VOLUME 146, NUMBER 6 21 NOVEMBER 1974 PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Bkeviora 1952- BULLETIN 1863- Memoirs 1864-1938 JoHNsoNiA, Department of MoUusks, 1941- OccAsiONAL Papers on Mollusks, 1945- SPECIAL PUBLICATIONS. 1. Whittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredini- dae (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms. 284 pp. 4. Eaton, R. J. E., 1974. A Flora of Concord. 250 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mammalian Hibernation. Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15. Proceedings of the New England Zoological Club 1899-1948. (Complete sets only.) Publications of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publications Office, Museum of Comparative Zoology, Hai^vard University, Cambridge, Massa- chusetts, 02138, U.S.A. © The President and Fellows of Harvard College 1974. THE ORB-WEAVER GENERA ARANIELLA AND NUCTENEA (Araneae: Araneidae) HERBERT W. LEVI Abstract. The species included in Araniella and Nuctenea have traditionally been included in Ara- neiis, but males differ in lacking the embolus cap. The lack of embolus cap can be related to differ- ences in mating behavior. Those orb-weavers witii cap (Amnetis) can mate only once with each palpus; Nuctenea males lacking a cap can mate several times. Four species of Araniella are known, one of diem Holarctic, the others Palearctic. Some Euro- pean populations are of interest as there are indi- cations that the species hybridize. Of the si.x species known to belong to Nuctenea, three are Holarctic, and three Palearctic. One of the Holarctic species may be a recent introduc- tion to North America; another may be cosmopoli- tan. Other species belonging to these two genera may be hidden among the two thousand si^ecies placed in Araneus and mostly poorly described. INTRODUCTION Araniella and Nuctenea species have traditionally been placed in Araneus. They include our commonest orb-weavers. Nev- ertheless the species are not well known, and in looking through the collections available, I found that many specimens were misidentified. The species of both genera are mainly Palearctic with some Holarctic species. I would like to thank the following for providing specimens for this study: D. Bixler; W. J. Gertsch and J. A. L. Cooke of the American Museum of Natural History, Cornell and Utah University collections; J. E. Carico; M. Grasshoff of the Sencken- berg Museum; C. Dondale and R. Leech, Canadian National collections, Ottawa; W. Hackman; G. H. Locket; M. Martelli of the Zoological Museum of the University of Florence; W. W. Moss of the Academy of Natural Sciences, Philadelphia; W. Peck, Exline-Peck collection; R. X. Schick of the California Academy of Sciences; J. Proszynski and W. Star^ga of the Polish Academy of Sciences; B. Vogel; H. K. Wal- lace and H. V. Weems, Florida State Col- lection of Arthropods. Information was provided by T. Kronestedt of the Natural History Museum, Stockholm; M. Moritz of the Zoological Museum of the Humboldt University, Berlin; and F. Wanless and D. Newman of the British Musevmi, Natural History. Lorna R. Levi and Lm Mackay edited the paper. This investigation and its publication were supported in part by National Science Foundation grant number GB-3616L One of the striking and puzzling features of these common orb-weavers is the enor- mous individual variation in genitalic structiu-e (Figs. 8-15, 17, 18, 67-76'), while there is little variation in the size and shape of the whole animal. This variation is found in Holarctic Araniella (Usplicata and A. cucurhitina of Europe as well as in Holarctic Nuctenea patagiata and Holarctic N. cornuta. Not infrequently specimens of these common species are sent to the museum by collectors who believe them to Bull. Mas. Comp. Zool., 146(6) : 291-316, November, 1974 291 292 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 be a new species. In their unusual varia- tion, Araniella and Nuctenea contrast with the small Aroneus species (Levi, 1973). Even though the species are fairly wide- spread, the differences between small Araneus species are far less than is found among individuals of a single collection of N. cornuto. The larger-sized Araneus species (A. nordmanni, A. saevus) are in- termediate in this respect ( Levi, 1971 ) . Large variation was not found in any of the widespread, common thcridiids such as Achaearanea tepidarioriim, although it does occur in the Tidarren species and Enoplognatha ovata. Variation in the genitalic structure of individuals was found in the zone of overlap between Araneus gemma and Araneus gemmoides; all evidence indicates hybridization (Levi, 1971 ) . Araniella species may also hybridize in Europe. Perhaps the individual varia- tion of Araniella cucurhitina of Europe is due to separation of northern and southern populations during the Pleistocene and the later hybridization occurred after the reced- ing of the ice. In none of the species of Araniella or Nuctenea is the variation geographic. But I have not studied various populations in detail. Shortly after Pet- iimkevitch ( 1925 ) wrote on the remarkable variation of genitalia of Agelena naevia, Seyler (1940), following up a hint from Gertsch (1934), correctly found that what Petrunkevitch called one species was in fact several. But I doubt that populations of Araniella displicata or Nuctenea patagiata and N. cornuta consist of sibling species. Of considerable interest is the relation- ship of Araniella species in Europe. Mr. Locket made me aware of this. While A. inconspicua and A. alpica appear distinct on the Continent, intermediates are found in Great Britain. While in many American araneid and theridiid species specimens from the Gulf Coast and Florida are noticeably smaller than those from other parts of North Amer- ica, Alaskan specimens of N. cornuta are smaller than those from southern Canada and the United States. To judge by the labels, N. cornuta is less dependent on houses in Alaska and probably competes with the native Araneus species. In north- eastern America all three Nuctenea species are usually found on buildings, but this is not true throughout their ranges. Nu- ctenea patagiata may be found under bark in woods. It is most unfortunate that at times names have to be changed as a result of revisionary studies. Araniella Chamberlin and Ivie AmnicUa Chamberlin and Ivie, 1942, Bull. Univ. Utah, biol. ser., 7(1): 76. Type species Epeira displicata Hentz, by original designation. The name is of feminine gender. Note. Chamberhn and Ivie (1942) do not give reasons for separating E. displicatus from Araneus other than that the species is close to those of Neoscona. I agree with this opinion. Diagnosis. There are no good superficial characters that separate female Araniella from the small species of Araneus. Females have a glabrous carapace and an oval abdomen, widest in the middle, lacking setae and lacking a folium pattern, but usually with paired black spots (Plate 1, Figs. 1, 16). The epigynum has a short, wide, wrinkled scape (Figs. 8, 25, 34, 40). The scape is not always clearly set off from the base of the epigynum. Unlike Araneus species, however, Araniella has, besides a single pair of seminal receptacles, a pair of sclerotized sacs (Figs. 7, 27, 33, 39) between the external entrance from outside to the connecting duct and the seminal receptacles. The entrance of the duct to the sclerotized sacs on each side is a slit. The ducts are hard to see in very sclerotized epigyna (A. cucurhitina). Unlike other araneids I have examined, Araniella has three macrosetae (Figs. 31, 42) on the patella of the male palpus. Species of most genera have only two or one. However, the palpal femur has a basal ventral tooth facing a tooth on the Orb-weaver Auaniella and Nuctenea • LcxA 293 endite as in Araneus. The palpus resembles that of Neoscona (Berman and Levi, 1971, fig. 31) in having the sclerites nearly fused, a small flap-like terminal apophysis (A in Figs. 20-22), and in laekiug a distal hematodoeha (Fig. 21). This eontrasts with the huge terminal apophysis and distal hematodoeha in Araneus and Nu- ctenea (Figs. 58, 61). AranicUa speeies also differ from Neoscona and Araneus in the hook-shaped, selerotized, median apoph- ysis ( M ) , dorsally directed toward the eymbium (Figs. 20, 22), and in the un- usually complex, large conductor (C). The embolus (E) lacks a cap. An embolus cap is always present in virgin males of Araneus species; the cap breaks off in mating. Araniella males have a hook on the distal margin of the first coxa, and the second femur has a matching depression. The legs of males are longer than those of females, and bear many macrosetae. Most Araniella are green or yellow to reddish in color when alive, similar to small species of Araneus. The color, except the black spots, washes out in alcohol. All species are about the same size and pro- portions (Figs. 1, 16). Natural history. All species build a small web between leaves; the web may be liorizontal. The spiders are active at day- time. The egg sac has a loose woollen ap- pearance (Plate 1).^ Distribution. The known species are all Palearctic, except A. displicata which is Holarctic in distribution. Misplaced species. Araniella geayi Ca- ' I liad anticipated that Araniella males, which resemble males of Nuctenea species in lacking a cap on the embolus, would likewise be able to mate repeatedly. Therefore, it was with consid- erable interest that, after submitting my own manuscript, I received R. Blanke ( 1973, Neue Ergebnisse zum Sexualverhalten von Araneus cu- ctirbitinus, Forma et Functio, 6: 279-290). iilanke did indeed observe differences between the behavior of A. cucurbitinus and that of Araneus species: The female approaches the male, the male taps the tarsi of females, and the male mates several times. Plate 1. Araniella displicata (Hentz). Above, female with egg sac. Below, male. Both laboratory photo- graphs, from Massachusetts. poriacco, 1954, Comm. Pontifica Acad. Sci., 16: 104 from Guyana has as a holot^'pe a juvenile specimen belonging to the genus Araneus deposited in the Zoological Mu- seum of the University of Florence, ex- amined. Key to Species of Araniella 1 Male, conductor of palpus with a distal lobe directed at and close to palpal pa- tella (Figs. 37, 42); female with epigy- 294 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Map 1. North American distribution of Araniella displicata (Hentz). niim scape having proximal part widest, much wider than tip (Figs. 34, 40) — _ 2 - Male palpal conductor without distal lobe toward patella (Figs. 18, 30); female epigynum with scape having parallel sides or proximal constriction (Figs. 10, 25 ) - 3 2(1) Median apophysis with fin toward the embolus ( Fig. 42 ) ; female with base of epigynum showing on each side of scape ( Fig. 40 ) ; Europe alpica - Median apophysis without fin (Fig. 37 ) ; female epigynum base not showing in ventral view (Fig. 34); Eurasia — _ inconspicua 3(1) Conductor with a set-off piece which holds terminal apophysis and embolus (Figs. 22, 30, 31); scape of epigynum narrower or of equal width to part of base visible on each side of it ( Figs. 23, 25); Eurasia, perhaps Arctic North Amer- ica cucu rbitina - Conductor without set-off piece holding terminal apophysis and embolus ( Figs. 17, 18); scape of epigynum much wider than part of base showing on each side of it (Figs. 8-14); North America and Eurasia displicata Araniella displicata (Hentz) Plate 1 ; Figures 1-21 ; Map 1 Epeira displicata Hentz, 1847, J. Boston Soc. Natur. Hist., 5: 476, pi. 31, fig. 17. Types from Alabama, May, Oct., destroyed. Emerton, 1884, Trans. Connecticut Acad. Sci., 6: 313, pi. 34, fig. 4, pi. 36, fig. 20, 342. Kevserling, 1893, Spinnen Amerikas, 4: 219, pi. 10, fig. 162, 9. Emerton, 1902, Common Spiders, p. 172, fig. 405. Kaston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 258, fig. 806, ? . Epeira decipiens Fitch, 1856, Trans. New York Agric. Soc, 15: 451. Male specimen from New York, lost. Epeira sexpunctata Keyserling, 1884, Verhandl. Zool. Bot. Gesell. Wien, 34: 530, pi. 13, fig. 28, 9 . Female type from North America in the Museum of Comparative Zoology, examined. Keyserling, 1892, Spinnen Amerikas, 4: 200, pi. 9, fig. 148. Orb-weaver Araniella and Nuctenea • Levi 295 ^r^W''%'^ Figures 1-19. Araniella displicata (Hentz). 1-15. Female. 1. Dorsal view. 2. Carapace. 3. Eyes and chelic- erae. 4. Internal genitalia with epigynum cleared. 5. Epigynum having scape torn off. 6-7. Internal genitalia. 6. Anterolateral. 7. Posterior. 8-15. Epigynum, 8, 10, 12, 14. Ventral view. 9, 11, 13, 15. Posterior view. 8, 9, 10, 11, 14, 15. (California.) 12, 13. (Oregon.) 16 19. Male. 16. Dorsal view. 17, 18. Left male palpus, mesal view. 19. Palpus, ventral view. 17. (Minnesota.) 18, 19. (Montana.) Scale lines. 0.1 mm; Figs. 1-3, 16, 1 mm. 296 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Epeira alba Keyserling, 1884, Verhandl. Zool. Bot. Gesell. Wien, 34: 530, pi. 13, fig. 20, 9. Fe- male t>pe from Kentucky in the Museum of Comparative Zoology, examined. Epeira cuctirhitina, - McCook, 1893, American Spiders, 3: 150, pi. 3, figs. 1-3, pi. 4, fig. 6, ?, $ . Not A. cuctirhitina (Clerck). Aranetis croaticus Kulczynski, 1905, Bull. Acad. Sci. Cracovie, 233, pi. 7, figs. 22, 30, 9 . Female holotype from Croatia in the Polish Academy of Sciences, Warsaw, examined. NEW SY- NONYMY. Aranea displicata, - Comstock, 1912, Spider Book, p. 494, figs. 524, 525, ? , web. Wiehle, 1931 in Dahl, Tierwelt Deutschlands, 23: 109, figs. 167- 170, 9, $. Comstock, 1940, Spider Book, 2nd ed., p. 508, figs. 524, 525, 9, web. Roewer, 1942, Katalog der Araneae, 1: 798. Araniella displicata, - Chamberlin and Ivie, 1942, Bull. Univ. Utah, biol. sen, 32(13): 76. Araniella displicata octopunctata Chamberlin and Ivie, 1942, Bull. Univ. Utah, biol. ser., 32(13): 76. Female holotype from Emigration Canyon, Wasatch Mts., Utah, in the American Museum of Natural History, paratype, examined. Araneus displicatns, - Locket and Millidge, 1953, British Spiders, 2: 149, figs. 96b, 97c, 99c, lOOc-e, 9, $. Araneus cucurbitimts displicatns, - Bonnet, 1955, Bibliographia Araneonnu, 2: 478. Note. It is of interest that this species described from Alabama is very rare at the present time in the Gulf states if present at all. Only a juvenile from Bankhead National Forest, Alabama, might be this species. Others collected by Archer in Alabama were Armieus gadus Levi that had been misidentified. Description. Female from California. Carapace yellowish with eyes on black spots. Legs yellow. Dorsum of abdomen yellowish with three pairs of circular black spots on the posterior part (Fig. 1). Carapace smooth, almost without setae. There is no thoracic depression. Diameter of posterior median eyes 1.2 diameters of anterior medians; anterior laterals 0.8 di- ameters of anterior medians; posterior laterals subequal in size to anterior medians. Anterior median eyes L5 diameters apart, 3.2 from laterals. Posterior median eyes one diameter apart, three from laterals. The height of the clypeus is about 1.5 di- ameters of the anterior median eyes. The chelicerae, which are not very strong, have four teeth on the anterior margin, three on the posterior. The legs are relatively heavy, with many macrosetae. The abdo- men is suboval, widest in the middle. Total length 5.3 mm. Carapace 2.5 mm long, 1.9 mm wide. First femur, 1.9 mm; patella and tibia, 2.4 mm; metatarsus, 1.4 mm; tarsus, 0.6 mm. Second patella and tibia, 2.2 mm; third, 1.3 mm; fourth, 2.2 mm. Male from California. Coloration as in female, except legs tend to be banded or distal ends of leg articles darker. Carapace smooth with head region relatively high and a shallow thoracic depression, having two lines crossing each other at right angles (Fig. 16). Eyes are subequal in size. An- terior median eyes 1.7 diameters apart, two from laterals. Posterior median eyes their diameter apart, 2.5 from laterals. The height of the clypeus is 1.5 diameters of the anterior median eyes. Total length 4.0 mm. Carapace 1.8 mm long, 1.7 mm wide. First femur, 1.8 mm; patella and tibia, 2.3 mm; metatarsus, 1.3 mm; tarsus, 0.6 mm. Second patella and tibia, 2.0 mm; third, 1.2 mm; fourth, 1.9 mm. Variation. Total length of females 4.8 to 7.2 mm. Carapace 2.0 to 2.7 mm long, 1.7 to 2.0 mm wide. Total length of males 4.0-5.0 mm. Carapace 2.0-2.4 mm long, 1.7-2.2 mm wide. The coloration is much more variable than the size; it is often greenish, reddish, brownish or yellowish on the abdomen. This pigment washes out, however, and in alcohol the abdomen is generally white. Diagnosis. This is probably the only species of Araniella occurring in North America, although A. cuctirhitina may occur in northern Canada and Alaska. Araniella displicata females can be sepa- rated by the wider and longer scape, which often hides the base of the epigynum; the widest part of the scape is generally toward its middle (Figs. 8-14). In pos- terior view the base is shorter, wider ( Figs. 9, 11, 13, 15) than that of A. cucurbitina. Orb-weaver Aranieu.a and Nuctenfa • Levi 297 Figures 20-21. Araniella displicata (Hentz). 20. Left male palpus, ventral view without cymbium, cleared. 21. Palpus expanded. Figures 22-31. A. cucurbitina (Clerck). 22. Male palpus, expanded. 23 26. Epigynum. 23, 25. Ventral. 24, 26. Posterior. 23, 24. (Taunus, Germany.) 25, 26. (Scotland.) 27-29. Epigynum cleared. 27. Ventral. 28. Lateral. 29. Posterior. 30, 31. Palpus, mesal. 30. (Poland.) 31. (Germany.) Abbreviations. A, terminal apophysis; C, conductor; E, embolus; M, median apophysis; R, radix; T, tegulijm. Scale lines. 0.1 mm. 298 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Males can readily be separated from A. cucurbitina and other species by the shape of the conductor, which holds the tip of the embolus and terminal apophysis ( Figs. 17-21). That of A. displicata has several large teeth around its margin while a special lobe on the conductor of A. cucur- bitina holds the tip of embolus and terminal apophysis (Figs. 22, 30, 31). The main sclerotized part of the conductor of A. cucurbitina has just one tip. Habits. Males are mature from late spring until late summer, females until fall. The spider is found by sweeping meadows and low bushes. This species makes a relatively small orb web, often among the leaves of bushes or underneath a single large leaf. Distribution. Europe, North America, from the Arctic to North Carolina, probably Alabama, and Arizona in the south; but apparently absent from the south-central states, southern Iowa, Nebraska to the Gulf (Map 1). Araniella cucurbitina (Clerck) Figures 22-31 Araneiis cucurbitimis Clerck, 1757, Aranei Svecici, p. 44, pi. 2, fig. 4, ? . Cleick's specimens in tlie Natural History Museum of Stockholm, origi- nally pinned and labeled by Thorell, not exam- ined. Locket and Millidge, 1953, British Spi- ders, 2: 144, figs. 96a, 97b, 98a, 99a, 9, $. Bonnet, 1955, Bibliographia Araneorum 2: 472 ( in part ) . Epeim proxima Kulczynski, 1885, Pamiet. Akad. Umiejet. Krakow, li: 19, pi. 9, fig. 11. Male holotype from Kamchatka in tlie Polish Acad- emy of Sciences, examined. NEW SYNONYMY. Araneus cucurbitina opisthographa Kulczynski, 1905, Bull. Acad. Sci. Cracovie, p. 232, pi. 7, figs. 2, 20, 23, 26, 9, $. Syntypes from nu- merous localities in Poland in the Polish Acad- emy of Sciences, Warsaw, examined. Aranea cucurbitina, - Wiehle, 1931, in Dahl, Tier- welt Deutschlands, 23: 106, figs. 161, 164, 9, $. Roewer, 1942, Katalog der Araneae, 1: 785. Aranea proxima, - Roewer, 1942, Katalog der Araneae, 1: 790. Araneus proximus, - Bonnet, 1955, Bibliographia Araneorum, 2; 571. Note. I could not find any consistent differences between specimens labeled opisthographa and others. Figure 30 was prepared from a syntype of A. opistho- grapha. Diagnosis. The short, narrow scape with parallel sides which exposes most of the base of the epigynum in ventral view (Figs. 23-26) readily separates A. cucur- bitina from other Araniella females. The more complex conductor with a distinct, separate lobe holding the tip of embolus and terminal apophysis (Figs. 22, 30, 31) distinguishes males (from other Araniella species ) . Natural history. Very common in trees and bushes in Europe (see Wiehle, 1931). Distribution. Common and widespread in Eurasia from Great Britain to Kamchatka (Wiehle, 1931; Locket and Millidge, 1953). The species is believed not to occur in North America. The single record is prob- ably the result of comparing specimens and misplacing them, or perhaps the species may occur in poorly collected Alaska and northern Canada. The female was from Fort Smith, Northwest Territory, 20. VI. 1967 (R. Leech), in white poplar. Araniella inconspicua (Simon) Figures 32-37 Epeira inconspicua Simon, 1874, Arachnides de France, 1: 84. Female type in the Musemn National d'Histoire Naturelle, Paris, not exam- ined. Aranea inconspicua, - Wiehle, 1931, in Dahl, Tier- welt Deutschlands, 23: 112; figs. 174-176, 9, $. Roewer, 1942, Katalog der Araneae, 1: 787. Arai^eus inconspicuus, - Locket and Millidge, 1953, British Spiders, 2: 146; figs. 97a, 98c, 99d, 100b, 9 , $ . Bonnet, 1955, Bibliographia Araneorum, 2: 521. Diagnosis. The abdomen usually lacks the black spots. The female has a triangular wrinkled scape that completely hides the base in ventral view (Fig. 34); in pos- terior view, the median groove is shorter (Figs. 35, 36) tlian that of A. alpica. The palpus lacks the distally directed fin of Orb-weaver Araniella and Nuctenea • Levi 299 .^, .fiftr^i'ji-^'^- Figures 32-37. Araniella inconspicua (Simon). 32-36. Epigynum. 32. Ventral, cleared. 33. Posterior, cleared. 34. Ventral. 35, 36. Posterior. 32-35. (France.) 36. (England.) 37. Left male palpus, mesal. Figures 38-42. A. alpica (L. Koch). 38-41. Epigynum. 38. Ventral, cleared. 39. Posterior, cleared. 40. Ven- tral. 41. Posterior. 42. Male palpus, mesal. Scale lines. 0.1 mm. the median apophysis (Fig. 37) present in A. alpica. While Continental specimens are readily separated from A. alpica, this is not true for those of the British Isles. Perhaps as a result of a recent introduc- tion of A. alpica they hybridize. Natural history. Found in trees and bushes at low elevations (Wiehle, 1931). Distribution. Europe from Great Brit- ain, northern Spain to Macedonia ( Wiehle, 1931). There are also references to A. inconspicua occurring in eastern Asia (Bonnet, 1955). Araniella alpica (L. Koch) Figures 38-42 Epeira alpica L. Koch, 1869, Z. Ferdinandeum Tirol, (3)14: 173. Specinien.s from Tyrol and other locaHtios in the Kocli collection of the Briti.sh Mu.seum (Natural History); presiunahly types but not examined. Aranca alf)ica, - Wiehle, 1931, in Dahl, Tier- welt Deutschlands, 23: 110, figs. 171-173, 9, $. Roewer, 1942, Katalog der Araneae, 1: 781. 300 Bulletin Museum of Comparative Zoology, Vol, 146, No. 6 Araneus alpiciis, - Locket and Millidge, 1953, Brit- ish Spiders, 2: 149, figs. 96c, 98d, 99b, 100a, 5 , $ . Bonnet, 1955, Bibliographia Araneomni, 2: 428. Araniella alpica, - Archer, 1951, Natur. Hist. Misc., 84: 3, fig. 4, $. Diagnosis. In ventral view the base of the epigynum shows as two bulges pos- terior and lateral to the scape (Fig. 40). In posterior view the central area is much longer (Fig. 41) than that of A. iricon- spicua. The median apophysis has a dis- tally directed fin ( Fig. 42 ) and the terminal apophysis is wider than that of other species. The abdomen has at most four black spots (see note under A. inconspicua diagnosis ) . Natural history. This species is found in European mountains above 300 m to krummholz ( 1800 m in the alps ) and is limited to fir and spruce. Males are ma- ture until August (Wiehle, 1931). Distribution. Great Britain ( Locket and MilHdge, 1953), Scandinavia, Central Eu- rope (Wiehle, 1931), Balkans (Bonnet, 1955). Nuctenea Simon Niictenea Simon, 1864, Histoire Naturelle des Araignees, p. 261. New subgenus of Epeira with the type species Epeira umbratica desig- nated by Bonnet, 1950, Bibhographia Araneo- rum, 2: 3118. The name is of feminine gender. Cyphepeira Archer, 1951, Natur. Hist. Misc., Chi- cago, 84: 4. New subgenus of Epeira with the type species by original designation Epeira ( Cy- phepeira) silvicultrix C. L. Koch. The name is of feminine gender. Note. The species included here had been placed by Wiehle ( 1931 ) in groups 4 and 5 of Aranea, by Locket and Millidge (1953) in groups 3 and 5 of Araneus. Archer (1951), following F. P.-Cambridge ( 1903 ) , considered the group a distinct genus, Epeira, with Araneus cornutus (Clerck) the type of the genus. But this type designation is an error as Epeira is an objective synonym of Ara^iea and a sub- jective synonym of Araneus, having Epeira diadernata as type designated by Latreille, 1810 (Levi, 1971). In considering the group included here as distinct, I am following older authors and also Archer. In 1959 Yaginuma and Archer included the species in Cyphepeira, as did Proszynski and Star^ga (1971). Wiehle (1927), discussing orb-web build- ing, included Epeira umbratica, E. sclo- petaria, E. cornuta and E. patagiata in a group making an unibraticus-type web. Gerhardt (1926) separated the group be- cause of different mating behavior: males can mate three or four times with each palpus. Araneus species can mate only once with each palpus. Also, the female assumes a different mating position, ap- proaching and hanging opposite the male, with cephalothorax lowered and abdomen raised, and pulls in the male on threads. The male will court a female that does not have an orb-web, and in mating the male's body is not as close to the female's as in Araneus. Males do not refill their palpi with sperm immediately after mating, as males of Araneus have been observed to do (observations on N. umbratica, N. cornuta, N. sclopetaria) . Description. All species are gray to brown, none brightly colored (Plate 2). The abdomen is dorsoventrally flattened, oval in outline, widest in the middle, with a folium on the dorsum (Figs. 97-109). The cardiac mark is usually dark. The venter of the abdomen is black with a pair of comma-shaped or bracket-shaped white marks (Figs. 98, 99, 102, 104). The genitalia are heavily sclerotized. The opening of the epigynum is hard to find and the connecting duct difficult to make out, even in epigyna digested with 10 percent NaOH. The openings of the N. umbratica epigynum are anterior on the base (Figs. 45, 46), those of other species, posterior in a groove (Figs. 71, 72, 81, 82, 85, 86). The palpus has a simple conductor ( C in Figs. 58-62), unlike that of Araniella (Figs. Orb-weaver Araniella and Nuctenea • Levi 301 20, 22) and even simpler than that of opening on (>ach side of the base. The Araneiis. A complex terminal apophysis palpus of Metazijgia has a terminal apoph- shiclds the embolus from above (A in Figs, ysis with a large proximal part and is very 58-62) and is connected to the bulb by different from that of Nwctenea. The palpal distal hematodocha. The distal hema- femur lacks the basal ventral tooth present todocha may be sclerotized, and reveals in Araneiis and Araniella but has a cor- its origin by the presence of folds and rc\sponding tooth on the side of the endite. grooves. Despite sclerotization, parts of Natural history. Nuctenea .species, at it expand (Figs. 58, 61, 62). The embolus least those found in North America, may lacks a cap and is a relatively simple be mature the year around; adult males structure. The tiny structure visible on can be found at all seasons. In contrast, the opening of the embolus (Figs. 119, 122, in all Araneus species observed in the 123, 125) is found only in mated males and temperate region, mature males can be is presumably dried up sperm fluid. The found only during a short period of the median apophysis is on the mesal side and year. Males, as well as females, can mate may project (except in N. umhratica) . In numerous times; .species belonging to most species it is biforked (Figs. 61, 62, Araneus, to judge by the work of U. 110-117, 126-129). It is a simple projec- Gerhardt (Levi, in preparation), can mate tion in N. silvicultrix (Figs. 55, 60) and a only twice, once with each palpus, perhaps lancet-shaped, appressed sclerite in N. three times if the mating was imsuccessful. umhratica (M in Fig. 58). Males have a No doubt these differences in habits are a liook on the distal margin of the first result of the cap, which is found on the coxa and a corresponding depression on embolus of Araneus, and is absent in Nu- the second femur; also the second tibia ctenea. may be swollen and bear macrosetae. The species all build in the evening and Diagnosis. In the Americas the species are nocturnal. The webs of all the species of Nuctenea can be confused only with have few radii (fewer than 20) with few those of Metazijgia. However, the carapace viscous threads, widely separated; those of Nuctenea is setose (Figs. 105-109), that of adult N. umhratica are separated by 10 of Metazijgia lacks setae. Unlike most mm or more. Thus the web, especially in species of Araneus, Nuctenea species have wind, gives the impression of being a flimsy the abdomen dorsoventrally flattened, oval structure. The center is small with rough in outline, and widest in the middle with a threads and few scaffolding threads. Al- dorsal folium. The venter is black between though usually made nearly vertical, the genital furrow and spinnerets, enclosing a web may be horizontal (N. cormita, Plate white, comma-shaped mark on each side 2). All species make a retreat or sit near (Figs. 97-109). The cardiac mark on the the web during the day, in the web at abdomen, if present, is dark, not light as night. it often is in Araneus. The only Araneus In the northeastern United States the species that have the abdomen oval are species arc commonly found on houses, marked differently. More important, Nu- Comstock (1912, 1940) refers to these ctenea genitalia are much more sclerotized, spiders as the House Araneas, J. H. Emer- and the embolus lacks a cap. Externally ton calls them House Epeiras. But, as the epigynum base seems more complex Comstock points out, they are often found (Figs. 43-46, 50-54, 71-76, 81-92) than in suitable habitats far away from houses, in Araneus and Metazijgia. The Metazijgia In America the species have a wide epigynum lacks the scape present in Nuc- distribution, N. cornuta from the Arctic tenea; in its place there is a ventrally to the tropics. Only N. sclopetaria ap- extended, laterally flattened lobe with the pears introduced; the other two species, 302 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 though they are commonly found on build- ings and in trash (Maps 2-4), as inti'o- duced species often are, have a continuous Holarctic distribution. Misplaced species. Epeira carolinalis Archer, 1951, Amer. Mus. Novitates, No. 1487: 40, fig. 57, ?, may belong to the genus Metazygia. The male is not known. Key to Species of Nuctenea 1 Male with median apophysis of palpus spHt into two branches (Figs. 110- 117); female with scape of epigynum originating at anterior of base (Figs. 71-91 ) 3 - Male with median apophysis of palpus not split into two branches (Figs. 47, 48, 55, 56, 58, 60); female with scape originating in center of base or lacking scape (Figs. 43, 51); Eurasia, North Africa 2 2(1) Terminal apophysis of male palpus a mas- sive shield (Figs. 48, 49, 58); median apophysis spindle-shaped (Figs. 47, 58); epigynum with triangular scape pointing posteriorly (Fig. 43); Europe and North Africa umbratica - Tenninal apophysis of palpus with teeth (Figs. 55, 60); median apophysis a knob (Figs. 55, 60); epig>'num with cone- shaped base drawn out anteriorly ( Figs. 51, 53); northern. Central Europe to Siberia silvicultrix 3(1) Males 4 - Females 7 4(3) Terminal apophysis a prong with a nar- row neck (Figs. 61, 110); embolus with tip hidden by a lamella (Figs. 61, 110, 118, 119), probably cosmopolitan _. cornuta - Terminal apophysis a sclerotized lobe without neck (Figs. 112-117); embolus cylindrical (Figs. 112, 114, 116) 5 5(4) Embolus with a distal, set-off finger (Fig. 116); Central Europe to Iran ixobola - Embolus cylindrical, pointed at end but without set-off finger (Figs. 112, 114, 120-125) 6 6(5) Median apophysis massive with the two prongs about of equal width in ventral view (Fig. 113); conductor leaning to- ward median apophysis (Fig. 113); Holarctic _ patagiata - Median apophysis more slender with "up- per" prong narrower than "lower" one (Fig. 115); conductor bending away from median apophysis (Fig. 115); Holarctic sclopetaria 7(3) Epigynal base posteriorly corrugated (Fig. 84); scape widest near tip witli a narrow neck (Figs. 78-81); Holarctic patagiata - Epigvnal base not corrugated (Figs. 74, 76, 88, 92) 8 8(7) Epigynum covered by a lamella on each side (Figs. 65-76); base with a ven- tral, anterolateral fold and swollen pos- teriorly (Figs. 63, 65, 67, 69, 73, 75); probably cosmopolitan cornuta - Epigynal lamellae not visible in ventral view or hidden behind base ( Figs. 85— 92); base without anterolateral fold and posterior swelling 9 9(8) Anterior part of scape framed by a lip of the base (Fig. 87); Holarctic sclopetaria - Anterior part of scape not framed by lips from base, base with anterolateral pockets (Fig. 91); Central Europe to Iran _... ixobola Nuctenea umbratica (Clerck) Figures 43-49, 58, 59, 93, 99, 105 Araiieus timbraticus Clerck, 1757, Aranei Svecici, p. 31, pi. 1, fig. 7, $ . Clerck's specimens from Sweden in the Museum of Natural History, Stockholm, labeled by Thorell; not examined. Locket and Millidge, 1953, British Spiders, 2: 139, fig. 92, ? , $. Bonnet, 1955, Bibliographia Araneonmi, 2: 621. Aranea sexpunctata Linnaeus, 1758, Systema Nat- urae, 10 ed., p. 622. Type specimens believed lost. Wiehle, 1931, in Dahl, Tienvelt Deutsch- lands, 23: 93, figs. 138-141, 9, $. Roewer, 1942, Katalog der Araneae, 1: 791. Epeira (Nuctenea) umbratica, - Simon, 1864, Histoire Naturelles des Araignees, p. 261. Epeira umbratica, - Nielsen, 1932, The Biology of Spiders, Copenhagen, vol. 2, figs. 299-304, web, egg sac, retreat. Cliinestcla umbratica, - Archer, 1951, Natur. Hist. Misc., Chicago, 84. Proszynski and Stargga, 1971, Katalog Fauny Polski, 16: 85. Description. Female from England. Cara- pace dark brown. Legs dark brown, with light bands. Dorsum of abdomen with usual dark brown pattern. Venter black with two lateral white marks. Diameter of posterior median eyes 0.9 diameters of anterior medians, laterals 0.8 diameters of anterior medians. Anterior median eyes their diameter apart, posterior medians slightly more than their diameter apart. Height of clypeus equals about the diam- eter of the anterior median eyes. The abdo- Orb-weaver Araniella and Nuctenea • Levi 303 Figures 43-49. Nuctenea umbratica (Clerck). 43 46. Epigynum. 43. Ventral. 44. Posterior. 45. Ventral, cleared. 46. Posterior, cleared. 47-49. Left male palpus. 47. Mesal. 48. Ventral. 49. Apical. Figures 50-57. N. silvicultrix (C. L. Koch). 50-54. Epigynum. 50. Anterior. 51. Ventral. 52. Posterior. 53. Lateral 54. Posterior, cleared 55 57. Male palpus. 55. Mesal. 56. Ventral. 57. Tibial macrosetae. Scale lines. 0.1 mm. men is much flattened. Total length 12 Male from England. Coloration as in mm. Carapace 4.7 mm long, 4.1 mm wide, female, with abdominal pattern more dis- First femur, 3.9 mm; patella and tibia, 5.(S tinct. Diameter of posterior median eyes mm; metatarsus, 3.6 mm; tarsus, 1.8 mm. 0.8 diameters of anteriors. Laterals very Second patella and tibia, 5.3 mm; third, slightly smaller than posterior median eyes. 3.0 mm; fourth, 4.6 mm. Anterior median eyes their diameter apart, 304 BuUeiin Museum of Comparative Zoology, Vol. 146, No. 6 Plate 2. Nuctenea cornuta (Clerck). Above, female (Wisconsin). Horizontal web with flies caught (Minnesota). posterior medians their diameter apart. The height of the clypeus is shghtly less than the diameter of the anterior median eyes. Total length 8 mm. Carapace 4.0 mm long, 3.6 mm wide. First femur, 5.2 mm; patella and tibia, 7.3 mm; metatarsus, 5.3 mm; tarsus, 2.0 mm. Second patella and tibia, 5.9 mm; tliird, 3.0 mm; fourth, 4.7 mm. Diagnosis. Nuctenea umbratica is read- ily separated from the other species of Nuctenea by being flatter (Figs. 93, 105), and by the distinct genitalia (Figs. 43-48). The openings of the epigynum, unlike those of other species, are anterior on the base (Figs. 45, 46) and the median apophysis is spindle-shaped (Figs. 47, 48, 58). Natural history. Nuctenea U7nhratica has its retreat in crevices under bark, between Orb-weaver Araniella and Nuctenea • Levi 305 Nuctenea cor Map 2. North American distribution of Nuctenea cornuta (Clerck). planks, in masonry, in barns, houses and greenhouses, up to 820 m in tlie Alps, but in southern Switzerland to 1200 m (Wiehle, 1931). Mature females ean be found at all seasons, males from June until Oetober. The eggs are laid in a flattened ball sur- rounded by loose, woolly silk. The web is more eccentric than that of JV. patap,i(ita, with the center closest to the retreat. The viscous threads may span 70 cm and there are about 20 radii. There is a line to the retreat. The animal is strictly nocturnal (Wiehle, 1927, 1931). Mating has been described by Gerhardt, 1926. Disirihution. Nuctenea uinbratica is only found in Europe and North Africa. 306 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Nuctenea silvicultrix (C. L. Koch), new combination Figures 50-57, 60, 106 Epeira silvicultrix C. L. Koch, 1845, Die Arachni- den, 11: 131, pi. 932, 933, 9, $. In the Ber- lin Museum are specimens from the L. Koch collection, presumably from Nlirnberg, but no specimens that can readily be interpreted as types of C. L. Koch. The British Museum (Natural History) has specimens from Nlirn- berg belonging to the L. Koch collection and presumably the types. They were not examined. Aranea silvicultrix Wiehle, 1931, in Dahl, Tier- welt Deutschlands, 23: 96, figs. 142-145, $, $. Roewer, 1942. Katalog der Araneae, 1: 792. Araneus silvicultor, - Bonnet, 1955, Bibliographia Araneorum, 2: 598. Epeira (Cyphepeira) silvicultrix, - Archer, 1951, Natur. Hist. Misc., Chicago, 84: 4. Cyphepeira silvicultrix, - Yaginuma and Archer, 1959, Acta Arachnol., 16: 41, fig. 12, $. Proszynski and Stargga, 1971, Katalog Fanny Polski, 16: 85. Description. Female from Schonhaid [near Neustadt, Bavaria]. Carapace red- brown, head lighter, with a double, median, longitudinal darker line. White setae in the head region. The legs are indistinctly banded with narrow, light bands. The dorsum of abdomen has a black folium out- lined by white. The venter is black with a white mark on each side. The diameter of the posterior median eyes is 0.8 diam- eters of anterior medians, laterals about 0.6 diameters. The anterior median eyes are about their diameter apart; the posterior medians slightly less than one diameter. The ocular quadrangle is slightly longer than wide, much narrower behind than in front. The clypeus slants back and its height is about equal to or sHghtly less than the diameter of anterior median eyes. The opening of the epigynum appears to be on the side of the base and is quite difficult to see. Total length 7.0 mm. Carapace 3.1 mm long, 2.8 mm wide. First femur, 3.0 mm; patella and tibia, 3.9 mm; metatarsus, 2.7 mm; tarsus, 0.9 mm. Second patella and tibia, 3.6 mm; third, 2.1 mm; fourth, 3.3 mm. Male from Erlangen, Bavaria. Colora- tion as in female. There is a shallow, round thoracic depression. Posterior and anterior median eyes subequal in size, laterals 0.8 diameters of medians. The anterior median eyes are one diameter apart; posterior ■ medians also one diameter apart. The ' height of the clypeus equals slightly less than the diameter of the anterior median eyes. Total length, 5.8 mm. Carapace 3.2 mm long, 2.6 mm wide. First femur, 3.8 mm; patella and tibia, 5.0 mm; metatarsus, 3.7 mm; tarsus, 1.5 mm. Second patella and tibia, 4.0 mm; third, 2.3 mm; fourth, 3.4 mm. Diagnosis. The epigynum is triangular, anteriorly extended (Figs. 50-54); the terminal apophysis (Figs. 55, 60) and strong setae on the palpal tibia (Fig. 57) separate this species from other Nuctenea. Natural history. In northern Bavaria the species is found among lichens on stunted pines growing on infertile, moist ground; it uses crevices as retreats. Mature males are collected in April, May and again in July and August. The web is similar to that of IV. umbratica (Wiehle, 1931). Distribution. Norway, Finland, Germany, Switzerland to Balkans, Ural Mountains and Turkmen (Bonnet, 1955). Nuctenea cornuta (Clerck), new combination, Furrow Spider Plate 2; Figures 61-76, 94, 97-98, 110-111, 118-119, 126; IVIap 2* Araneus cornutus Clerck, 1757, Aranei Svecici, p. 39, pi. 1, fig. 11, 9 . Female types from Swe- den lost. Locket and Millidge, 1953, British Spiders, 2: 134, figs. 88a, 89b, 90c, ?, $. Bonnet, 1955, Bibliographia Araneorum, 2: 463. Aranea foliata Fourcroy, 1785, Entomologia Pari- siensis, 533. Type specimen from Paris, France, * Correction, July 1974. K. Thaler recently found that what has been called Araneus cornutus in Europe in fact represents two species, a nortliern one and a southern one (Zool. Anz., in press). The American specimens are all hke tlie northern European N. cornuta. I have examined specimens sent by K. Thaler and agree witli his conclusions. Figures 63-66 of this paper thus do not belong to N. cornuta. Orb-weaver Araniella and Nuctenea • Levi 307 immM' Figures 58 62. Nuctenea expanded left male palpus. 58, 59. N. umbratica (Clerck). 60. N. silvicultrix (C. L. Koch). 61, 62. N. cornuta (Clerck). Figures 63-70. N. cornuta (Clerck), epigynum, variation. 63, 65, 67, 69. Ventral. 64, 66, 68, 70. Posterior. 63 66. (Burgenland, Austria.) 67, 68. (Kamtchatka.) 69, 70. (Poland.) Abbreviations. A, terminal apophysis; C, conductor; E, embolus; I, stipes; M, median apophysis; R, radix; S, sub- tegulum; T, tegulum. Scale lines. 0.1 mm. 308 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Nuctenea patagi Map 3. North American distribution of Nuctenea patagiata (Clerck). believed lost. Wiehle, 1931, in Dahl, Tierwelt Deutschlands, 23: 86, figs. 124-127, ?, $. Roewer, 1942, Katalog der Araneae, 1: 800. ?Epeiia frondosa Walckenaer, 1841, Histoire Nat- iirelle des Insectes, Apteres 2: 65. The type is Abbot's manuscript illustration, fig. 326, from Georgia in the British Museum, Natural History, copy in the Museum of Comparative Zoology, examined. Epeira strix Hentz, 1847, J. Boston Natur. Hist. Soc, 5: 473, pi. 31, fig. 5, 9. Type specimens from Pennsylvania and Alabama destroyed. Epeira vicaria Kulczynski, 1885, Pam Akad. Umiej. Krakow, 11: 5. Female holotype from Kamchatka in the Polish Academy of Sciences, examined. Aranea frondosa, - Comstock, 1912, Spider Book, p. 487, figs. 104-106, 128, 186, 194, 509, 513- 516, 9, $, web; 1940, op. cit., rev. ed., p. 501, figs. 104-106, 128, 186, 194, 509, 513-516, ?, $ , web. Epeira cornuta, - Nielsen, 1932, The Biology of Spiders, Copenhagen, vol. 2, figs. 289, 290, retreat. Epeira foliata, - Kaston, 1948, Connecticut Geo!. Natur. Hist. Surv. Bull., 70: 254, figs. 787, 803, 812, 2043, 9, $, web. Ciiphepcira cornuta, - Yaginuma and Archer, 1969, Acta Arachnol., 16: 41. Proszynski and Stargga, 1971, Katalog Fauny Polski, 16: 82. 'Note. Figures 67, 68 were prepared from the holotype of A. vicaria. Variation. Total length of females 6.5- 14.0 mm. Carapace 2.4-5.0 mm long, 1.9- 4.5 mm wide; first patella and tibia, 2.6- 5.8 mm long. The total length of males 4.7-8.5 mm. Carapace 2.1-4.2 mm long, 1.8-3.5 mm wide; first patella and tibia 3.0-6.0 mm. The smallest American sped- Orb-weaveh Araniella and Nuctenea • Levi 309 mens come from Alaska, tlie largest from the area of New England to Texas. Dia<^n(ms. The two anterior lateral lobes of the base of the epigynum are bent over and face posteriorly (Figs. 63, 65, 67, 69, 73, 75). (Those of N. sclopetaria are not bent over.) The posterior face of the epigynum is smooth (Figs. 74, 76), not grooved as in N. patagiota. The palpal terminal apophysis has a prong with a tip wider than its neck (Figs. 61, 110); in N. sciopetarm and N. patagiata it is wide. The embolus of N. cornuta has a wide lamella toward the mesal side (Figs. 61, 110, 118, 119) while in N. sclopetaria and N. patagiata it is simple and cylindrical. Natural history. The web is found around houses, frequently in bushes (Comstock, 1912), often near water (Wiehle, 1931; Kaston, 1948); it has 15-20 spokes and is up to 60 cm in diameter (Kaston, 1948). It is illustrated by figure 516 in Comstock (1912) and by Kaston (1948). The web is made at night, when the spider leaves its retreat. The silken retreat may be in crevices of walls, on railings, or among plants. The retreat has been illustrated by Nielsen ( 1932 ) . Both sexes are mature all summer, from March in the southern part of the range; males are more com- monly mature in spring and late fall (Kaston, 1948). Egg sacs, according to Kaston (1948), are 7-10 mm in diameter, covered with yellowish threads, hidden in the retreat, and contain 50 to 250 eggs. A female may make as many as ten egg sacs. Distribution. Holarctic, perhaps carried by man worldwide. In North America it occurs from the Arctic to Central America, but is most common in the eastern U.S. and Canada, Newfoundland to Florida (Map 2). Nuctenea patagiata (Clerck), new combination Figures 77-84, 100-102, 107, 112-113, 120-123, 127; Map 3 ?Araneus ocellatus Clerck, 1757, Aranei Svecici, p. 36, pi. 1, fig. 9, 9 . Female holotype from Sweden, lost. Bonnet, 1955, Bibliographia Anmeonim, 2: 555. .\iancu\ i>(iia^.i(itus Clerck, 1757, Aranei Svecici, p. 38, pi. 1, fig. 10, 9. Locket and Millidge, 1953, British Spiders, 2: 136, figs. cS9c, 90b, 9, $. Arcnica dumetorum Fourcroy, 1785, Entomologia l^irisiensis, p. .534. Type from Paris, France, belie\ed lost. Wiehle, 1931, in Dahl, Tierwelt Deutschlands, 23: 88, figs. 128, 129, 9, $. U.pcira itJiaca MeCook, 1893, American Spiders, 3: 152, pi. 4, fig. 3, $. Male lectotype from Ithaca, New York, in the Academy of Natural Sciences, Philadelphia, examined. NEW SY- NONYMY. Aranca ocellata, - Comstock, 1912, Spider Book, p. 489, figs. 107, 517-518; 1940, op. cit., rev. ed., p. 50.3, figs. 107, 517-518, 9, $. Epeiia patagiata, - Nielsen, 1932, The Biology of Spiders, Copenhagen, vol. 2, fig. 305, web. Epeiia (luiiictoiiim, - Kaston, 1948, Connecticut Ceol. Natur. Hist. Surv. Bull., 70: 255, figs. 788, 804, 813, 9, $. Cyphcpcim patagiata, - Yaginuma and Archer, 1969, Acta Arachnol., 16: 41. Proszynski and Stargga, 1971, Katalog Fanny Polski, 16: 84. Note. According to Bonnet (1955), C. L. Koch, 1845 (Die Arachniden, 11: 115) was the first revisor, synonymizing the two names of Clerck and choosing the name patagiata. Variation. Total length of females 5.5- 11.0 mm. Carapace 2.5-4.0 mm long, 2.2- 3.3 mm wide; first patella and tibia, 3.5-5.2 mm. Total length of males 5.8-6.5 mm. Carapace 2.9-3.8 mm long, 2.3-3.1 mm wide; first patella and tibia, 4.5-5.5 mm. Diagnosis. Females are separated from the other species by the epigynum: its base is furrowed posteriorly and its scape has a narrow neck (Figs. 78-84). The male dif- fers from IV. cornuta by having the terminal apophysis a flat lobe (as in N. sclopetaria); it differs from both the other species in having a deep division in the heavy median apophysis (Figs. 112, 113, 127) and in having a finger-shaped embolus ( Figs. 112,120-123). Natural history. Kaston (1948) indicates that its habits are similar to those of N. cornuta. According to Wiehle ( 1931 ) the web has 20-24 spokes with about 16 viscid threads above, and 23 below center; 310 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Map 4. North American distribution of Nuctenea sclopetaria (Clercl^). the orb is 25 cm across. The retreat is less silk lined than that of N. cornuta. The web has been illustrated by Nielsen (1932, fig. 305). I think the species prefers more arid, shaded areas than N. cornuta. I have found the retreat under bark in lodgepole pine at Jackson Hole, Wyoming, in a rather dry area. According to Wiehle ( 1931 ) and Kaston (1948) there may be a signal thread to the reti^eat or the spider may use a radius to return to it. Distribution. Some American authors indicate that this is a more northern species than N. cornuta. This may not be quite correct; however, the species is not found in die southeastern states. It appears to be common from the Arctic to North Carolina and Arizona but is much commoner than N. cornuta in the west, the Rocky Moun- tains, and the Pacific northwest states ( Map 3 ) ; also Eurasia. Nuctenea sclopetaria (Clerck), new combination Bridge spider, Gray-cross spider Figures 85-88, 103-104, 108, 114-115, 124, 125, 128; IVIap 4 Araneus sericatus Clerck, 1757, Aranei Svecici, p. 40, pi. 2, fig. 1, 9 . Female type from Sweden lost. Bonnet, 1955, Bibliographia Araneorum, 2: 594. Araneus sclopetaritis Clerck, 1757, Aranei Svecici, p. 43, pi. 2, fig. 3, $ . Type specimen from Sweden lost. Locket and Millidge, 1953, Brit- ish Spiders, 2: 136, figs. 88b, 89a, 90a, 9, $. Aranea undata Olivier, 1789, Encycl. Method. Hist. Nat. Ins. Paris, 4: 206. New name for sclope- tariits Clerck, but preoccupied by DeGeer, 1778. Wiehle, 1931, in Dahl, Tierwelt Deutsch- lands, 23: 90, figs. 130-133. Aranea ovigera Panzer-, 1804, Syst. NomencL, in Schiiffer, Icon. Ins. Ratisb., 1: 244, pi. 174, fig. 3. Aranea sericata, - Comstock, 1912, Spider Book, p. 486, figs. 510-512, 9,5; 1940, op. cit., rev. ed., p. 500, figs. 510-512, 9, $. Aranea ovigera, - Roewer, 1942, Katalog der Araneae, 1: 801. Epeira undata, - Kaston, 1948, Bull. Connecticut Geol. Natur. Hist. Surv., 70: 256, figs. 789, 805, 814-815, 2044-2046, 2, S, web, egg sac. Cyphepeira sclopetaria, - Yaginuma and Archer, 1959, Acta Arachnol., 16: 41. Cyphepeira sericata, - Proszynski and Star§ga, 1971, Katalog Fanny Polski, 16: 84. Note. According to Bonnet (1955), O. P.-Cambridge (1874, Trans. Linnean Soc, 30: 330) first synonymized the two simul- taneously, published names sericatus and sclopetarius, and chose sclopetarius. How- ever, this seems to be an error; the names were first synonymized by Westring ( 1851, Orb-weaver Araniella and Nuctenea • Levi 311 Figures 71-76. Nuctenea cornuta (Clerck), epigynum. 71, 73, 75. Ventral view. 72, 74, 76. Posterior view. 71-72. Cleared. 71-74. (Panama Canal Zone.) 75,76. (Alberta.) Figures 77-84. W. parag/afa (Clerck), epigynum. 77-81,83. Ventral. 82,84. Posterior. 81,82. Cleared. 77. Probably epigynum before last molt. (South Dakota.) 78, 79. (Alberta.) 80. (British Columbia.) 81 84. (On- tario.) Figures 85-88. N. sclopetaria (Clerck), epigynum. 85, 87. Ventral. 86, 88. Posterior. 85, 86. Cleared. Figures 89-92. N. ixobola (Thorell), epigynum. 89, 91. Ventral. 90, 92. Posterior. 89, 90. Cleared. Scale lines. 0.1 mm. 312 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 Goteborg Kongl. Vet. Hand!., 2: 34), who also chose sclopetaria. Thorell (1856, N. Acta Reg. Soc. Sci. Uppsala, p. 22) also lists the synonymy under sclopetaria. Bonnet shows that prior to 1938 the usage of sclopetaria outweighed sericata, although sericata has been uniformly used in North America and also by Bonnet ( 1955 ) . I will follow European arachnologists and use sclopetaria as do Locket and Millidge ( 1953, British Spiders, 2). (See Article 24a of the International Code on Zoological Nomenclature, 1961.) Variation. Total length of females 8-14 mm. Carapace 3.9-4.3 mm long, 3.1-4.0 mm wide. First patella and tibia 5.3-7.0 mm. Total length of males 6-7 mm. Car- apace 3.7-3.2 mm long, 2.9-3.3 mm wide. First patella and tibia 6.2-7.0 mm. Diagnosis. This species can usually be separated from N. cornuta and N. patagiata by the white hairs around the border of the carapace and by the fact that the background of the eye region is lighter brown than the area behind it and than the area of the clypeus (Fig. 108). The female's epigynum has the scape finger- shaped, as in N. cornuta but not as in N. patagiata, and the anterolateral margins of the base lobed and flat ( Fig. 87 ) , not as in N. cornuta. The openings of the epigynum are in dark posterior swellings of the base (Fig. 85). The palpus has a lobe-shaped terminal apophysis (Figs. 114, 115), some- what like that of N. patagiata. The median apophysis is not as deeply divided as that of N. patagiata (Figs. 115, 128). The embolus resembles that of N. patagiata (Figs. 124, 125). Natural history. This is the least common of the three Nuctenea species in North America and is found on houses and other buildings, often near water. One collec- tion from West Virginia came from sweep- ing honeysuckle bushes (Lonicera sp.). Many webs may be found touching one another (Kaston, 1948). In Europe the species is also found on buildings, and especially on bridges and cliffs above water (Wiehle, 1931). The orbs have up to 20 radii, with the viscid spiral separated, and the web reaches 70 cm in diameter (Wiehle, 1931). The web is illustrated in Kaston (1948, figs. 2044, 2046). The spider rests near the end of one of the frame threads rather than in a retreat. The egg sac, according to Kaston, contains 114-337 eggs, and is illustrated in his figure 2045. Distribution. Eurasia. In America this species is probably introduced, judging by its close association with buildings and its limited distribution, which matches that of Araneus diadematus Clerck. It is found from Newfoundland to southern Alaska, south to North Carolina, and is most abun- dant in the Great Lakes states ( Map 4 ) . Nuctenea ixobola (Thorell) new combination Figures 89-92, 95, 109, 116-117, 129 Epeim ixobola Thorell, 1873, Remarks on Syn- onyms of European Spiders, p. 545. Two male and three female syntypes from Austria in the Natural History Museum in Stockholm, not examined. Aranea ixobola, - Wiehle, 1931, in Dahl, Tier- welt Deutschlands, 23: 92, figs. 134-137, 9, $. Roewer, 1942, Katalog der Araneae, 1 : 788. Araneus ixobolus, - Bonnet, 1955, Bibliographia Araneorum, 2: 523. Description. Female from Poland. Car- apace very flat, no thoracic depression. Secondary eyes all about 0.7 diameter of anterior median eyes. Anterior median eyes slightly more than their diameter apart, posterior median eyes their diameter apart. Lateral eyes on tubercles and widely separated from the median eyes. The height of the clypeus equals 0.6 diameters of the anterior median eyes. Total length 13 mm. Carapace 6.5 mm long, 5.3 mm wide. First femur, 6.3 mm; patella and tibia, 8.6 mm; metatarsus, 5.7 mm; tarsus, 2.0 mm. Second patella and tibia, 8.0 mm; third, 4.4 mm; fourth, 6.3 mm. Male from Poland. Carapace and eye arrangement as in female. Total length 12 mm. Carapace 5.4 mm long, 4.7 mm Orb-weaver Araniella and Nuctenea • Levi 313 103 Figures 93 95. Eye region and chelicerae. 93. Nuctenea umbratica. 94. N. cornuta. 95. N. ixobola. Figure 96. Left anterior femora and carapace of N. patagiata. Figures 97-104. Abdomen. 97-98. N. cornuta. 99. N. umbratica. 100-102. N. patagiata. 103 104. N. sclo- petaria. 97, 100, 101, 103. Dorsal. 98, 99, 102, 104. Ventral. Figures 105-109. Carapace and abdomen, dorsal. 105. N. umbratica. 106. N. silvicultrix. 107. N. patagiata. 108. N. sclopetaria. 109. N. ixobola. Scale lines. 1.0 mm. 314 Bulletin Museum of Comparative Zoology, Vol. 146, No. 6 wide. First femur, 8.3 mm; patella and tibia, 11.8 mm; metatarsus, 8.2 mm; tarsus, 3.3 mm. Second patella and tibia, 10.0 mm; third, 5.0 mm; fourth, 7.4 mm. Diagnosis. Females have a narrow scape in the epigynum as does Nuctenea sclopefaria, with which it has been con- fused. However, the anterior lateral end of the base differs (Figs. 89, 91) and posterior lobes of the base do not extend ventrally. The male differs from that of N. sclopetaria in having a differently shaped embolus (Fig. 116). Natural history. According to Wiehlc (1931), this species is similar in habits to N . sclopetaria and replaces it in eastern Europe. It lives on buildings, fences, and bridges near water. Distribution. From Central Europe to Iran (Roewer, 1942; Bonnet, 1955). Speci- mens examined came from Leopoldshall (Anhalt, German Democratic Republic), Biolowieza, Distr. Hajnowka, and Distr. Kosice, Poland. REFERENCES CITED Berman, J. D., AND H. W. Levi. 1971. The orb-weaver genus Neoscona in North America (Araneae: Araneidae). Bull. Mus. Comp. Zool., 141: 465-500. Cambridge, F. O. Pickard-. 1903. Araneidea, In Biologia Centrali-Americana, 2. CoMSTOCK, J. H. 1912. The Spider Book. Garden City, New York: Doubleday, Doran and Co. . 1940. The Spider Book, rev. ed. Ithaca: Conistock Publ. Co. Gerhardt, U. 1926. Weitere Untersuchungen zur Biologie der Spinnen. Z. Morphol. Okol. Tiere, 6: 1-77. Gertsch, W. J. 1934. Further notes on Amer- ican spiders. Amer. Mus. Novitates, No. 726: 1-26. Levi, H. W. 1971. The Diademattts group of the orb-weaver genus Araneus north of Mex- ico (Araneae: Araneidae). Bull. Mus. Comp. Zool., 141: 131-179. . 1973. Small orb-weavers of the genus Araneus north of Mexico (Araneae: Aranei- dae). Bull. Mus. Comp. Zool. 145(9): 473-552. IX PREPARATiox. The presence of the cap on palpal emboli and mating behavior. Locket, G. H., and A. F. Millidge. 1953. British Spiders, 2: 1-449. Petrunkevitch, a. 1925. External reproductive organs of the common grass spider Agelena naevia Walckenaer. T- Morphol., 40: 559- 573. Proszynski, J., and W. Starega. 1971. Katalog Fauny Polski, 33: 1-382. ' Seyler, p. J. 1940. The generic and specific status of four spiders of the genus Agelenop- sis. Ohio J. Sci., 41: 51-69. Wiehle, H. 1927. Beitrage zur Kenntnis des Radnetzbaues der Epeiriden, Tetragr.athiden und Uloboriden. Z. Morphol. Okol. Tiere, 8: 468-537. . 1931. Araneidae. In F. Dahl, Die Tierwelt Deutschlands, 23: 1-136. Yaginu.ma, T., and a. F. Archer. 1959. Genera of the Araneine Argiopidae foimd in the Oriental region and generally placed under the comprehensive genus Araneus. Acta Arachnol., 16: 34-41. INDEX Valid names are printed in italics. Page numbers refer to main references, starred page numbers to illustrations. alba, Epeira 296 alpica, Aranea 299 alpica, Araniella 299* alpica, Epeira 299 alpicus, Araneus 300 Araniella 292 cornuta, Cyphepeira 308 corniita, Nuctenea 304*, 306, 307*, 311*, 313*, 315* cornutus, Araneus 306 croaticus, Araneus 296 cucurbitina, Aranea 298 cucurhitina, Araniella 297*, 298 cucurbitina, Epeira 296 cucurbitinus, Araneus 298 Cyphepeira 300 decipiens, Epeira 294 displicata, Aranea 296 (lisplicata, Araniella 293*, 294, 295*, 297* displicata, Epeira 294 displicatus, Araneus 296 dumetorum, Aranea 309 dumetorum, Epeira 309 OlUJ-\\'KA\'KH ArANIKLLA AND NuCTENEA • Lcvi 315 126ij Figures 110-117. Left male palpus. 110, 111. Nuctenea cornuta (Clerck). 112, 113. N. patagiata (Clerck). 114, 115. A/, sc/ope/ar/a (Clerck). 116,117. N. ixobola (ThoreW). Figures 118-125. Embolus of male palpus. 118,119. N. cornuta. 120 123. N. patagiata. 124, 125. N. sclope- taria. 118,120,121,124. Probably virgin. 119,122,123,125. Probably mated. 118. Mesal-apical. 119 125. Mesal. Figures 126-129. Male palpus, apical view. 126. N. cornuta. 127. N. patagiata. 128. N. sclopetaria. 129. N. ixobola. Scale lines. 0.1 mm. 316 Bulletin Museum of Coinparative Zoology, Vol. 146, No. 6 foliata, Aranea 306 foliata, Epeira 308 frondosa, Aranea 308 frondosa, Epeira 308 inconspiciia, Aranea 298 inconspiciia, AranieUa 298, 299* inconspicua, Epeira 298 inconspicuus, Araneus 298 ithaca, Epeira 309 ixobola, Aranea 312 ixobola, Epeira 312 ixobola, Nuctcnea 311*, 312, 313*, 315* ixobolus, Araneus 312 Niictenea 300 ocellata, Aranea 309 ocellatus, Araneus 309 octopunctata, Araniella displicata 296 opisthographa, Araneus cucurbitina 298 ovigera, Aranea 310 patagiata, Cyphepeira 309 patagiata, Epeira 309 patagiata, Nuctenea 309, 311*, 313*, 315^ patagiatus, Araneus 309 proxima, Aranea 298 proxima, Epeira 298 sclopetaria, Cyphepeira 310 sclopetaria, Nuctenea 310, 311*, 313*, 315* sclopetarius, Araneus 310 sericata, Cyphepeira 310 sericatus, Araneus 310 sexpunctata, Aranea 302 sexpunctata, Epeira 294 sihicultor, Araneus 306 sihicultrix, Aranea 306 silvicultrix, Cyphepeira 306 sil\icultrix, Epeira 306 silvicultrix, Nuctenea 303*, 306, 307*, 313* strix, Epeira 308 lunbratica, Chinesteha 302 umbratica, Epeira 302 umbratica, Nuctenea 302, 303*, 307*, 313* undata, Aranea 310 undata, Epeira 310 vicaria, Epeira 308 i us ISSN 0027-4100 SuUetin OF the Museum of Comparative Zoology The Anatomy of Saurosuchus galilei and the Relationships of the Rauisuchid Thecodonts WILLIAM D. SILL HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS, U.S. A VOLUME 146, NUMBER 7 21 NOVEMBER 1974 PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952- BULLETIN 1863- Memoers 1864-1938 JoHNSONiA, Department of Mollusks, 1941- OccAsioNAL Papers on Mollusks, 1945- SPECIAL PUBLICATIONS. 1. Whittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredini- dae (Mollusca: Bivalvia), 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms. 284 pp. 4. Eaton, R. J. E., 1974. A Flora of Concord. 250 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C, P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mammalian Hibernation. Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15. Proceedings of the New England Zoological Club 1899-1948. (Complete sets only.) Publications of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publications Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massa- chusetts, 02138, U.S.A. © The President and Fellows of Harvard College 1974. THE ANATOMY OF SAUROSUCHUS GALILEI AND THE RELATIONSHIPS OF THE RAUISUCHID THECODONTS WILLIAM D. SILL' Abstract. Saitrosuclius galilei was a large quadrupedal carnivorous thecodont from the Ischigualasto Formation of western Argentina, which is of approximately Carnian age. Its skull anatomy indicates that it descended from an erythrosuchid t\'pe of primitive thecodont. Sauro- suchus, together with Luperosuchtts, Prestosuchiis, Ticinos'UcJius, "Mandasiichus" and possibly some other less well known genera, form a well-defined lineage that can be trticed throughout most of the Triassic. Rauisuchus is considered a member of the same family, and thus the earlier name Rauisuchidae is retained for the group. Two other thecodont lineages, the Proterochampsidae and the Ornithosuchidae, are traced throughout the Tri- assic. The relationships of the three families strongly suggest that they are independent deri- vations of the three Early Triassic primitive families. Dinosaur origins remain unclear. There is no good evidence for associating the Raui- suchidae with early dinosaurs; on the contrary, there is an unexplained time oxerlap of large carnivorous dinosaurs and thecodonts that have nearly identical adaptations. INTRODUCTION Saurosiichus g,aUlei is one of the 18 or more genera of reptiles found in the now legendar}' Ischigualasto Basin of western Argentina. Its significance for paleonto- logic studies lies in the excellent preserva- tion of the material, particularly of the skull and tarsus, which makes possible the clari- fication of the anatomy of the closely related Brazilian thecodonts, and generally aids interpretation of the family Raui- suchidae on a worldwide basis. Together ' Universidad N'acional de San Juan, Dept. Geo- logia, San Juan, Argentina. Bull. Mus. Comp. with Ticinosuchus from the Middle Triassic of Switzerland, Saurosiichtis provides a key for tracing a thecodont lineage that was world-wide in distribution throughout most of the Triassic Period. Most of the specimens used for this study were collected in the Ischigualasto For- mation by expeditions from the Instituto Miguel Lillo of Tucuman, Argentina. The first specimen was collected in 1959, under the direction of Dr. Osvaldo Reig. Sub- sequent expeditions, led by Jose Bonaparte, recovered parts of four additional individu- als. From these various parts, most of the skeleton can be reconstructed, although the forelimb is not represented in any of the specimens. Saurosiichus was one of the largest the- codonts of its time, and no doubt competed with the emerging dinosaurs for the large carnivore role. Thecodonts, of course, lost the competition, and contemporary dino- saurs, both saurischian and ornithischian, from the Ischigualasto Formation indicate that superior locomotion was a factor related to dinosaurian dominance. At pres- ent, although Saurosuchus appears to be the most advanced member of the family yet described, it is less progressive ana- tomicalK' than its dinosaiuian contempora- ries. The lineage of Saurosuclius proxides e\idence to support the premise that pro- gressive thecodonts were competitors rather than progenitors of the dinosaurs. Abbreviations for the institutions referred to in this report are as follows: Zool., 146(7): 317-362, November, 1974 317 318 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 PVL Institute Miguel Lillo, Tucuman, Argentina DGM Division of Mines and Geology, Brazil T University of Tubingen, Ger- many PIMZ Paleontological Institute, Zurich, Switzerland MSJ Museum of Natural Sciences, San Juan, Argentina Acknowledgements. This study was made during a year's stay at the Institute Miguel Lillo in Tucuman, Argentina. Special thanks are due to Jose Bonaparte and the directors of the Institute, whose help and generosity made the study possible. I am also greatly indebted to A. W. Crompton of Harvard University and John Ostrom of Yale for their technical help and personal assistance. Many colleagues offered sug- gestions and gave perspective to the re- search; among them were A. S. Romer, Bernard Krebs, Alan Charig, A. Keyser, and Alick Walker. Drawings were made by Alexander Gavriloff. Funds for the Research were provided by NSF Grant GB-4435X1. Geologic Setting The Ischigualasto Basin (Hoyada de Ischigualasto or Valle de la Luna) fomis a depression on the western limb of a large syncline whose axis runs northwest-south- east. Differential erosion of the soft clay- stones of the Ischigualasto Formation cre- ated a prominent depression at the base of the cliff-foiTning red sandstones of the Los Colorados Formations (see Fig. 1). The Triassic sediments extend approximately one hundred kilometers, and are bounded on the south by the Valle Fertil mountains and on the north by the Sierra de Mas range. Within this area of outcrop there are numerous minor flexures, principally anticlines. One such saddle-shaped anti- cline divides the basin into a northern and southern portion; this division coincides with the boundary between the provinces of San Juan and La Rioja. The southern, or San Juan, portion is the larger of the two and has produced most of the fossils known from the basin. East of the depression, the opposite limb of the large syncline has ex- posed the type area of the earlier Chaiiares Formation. Interpretation of the time-stratigraphic relationships of the sedimentary units in the Ischigualasto basin has varied considerably. For many years the whole succession was considered "Rhaetic," or uppermost Tri- assic. With the discovery of vertebrate fossils that were more primitive than the classic Upper Triassic fauna, vertebrate paleontologists assigned the Ischigualasto Formation to the Middle Triassic. As new discoveries are being made a consensus is forming that the Ischigualasto Formation is most probably of Camian age, possibly Late Ladinian, with the underlying Los Rastros Formation closely equivalent in time to the Santa Maria Formation of Brazil. (I have elsewhere summarized the various interpretations of the South Ameri- can Triassic: Sill, 1969.) Although the general geologic relation- ships between the various formations are quite straightforward, no attempt has yet been made to study sedimentary cycles within the Ischigualasto Fonnation, or to correlate the occurrence of specific fossils with different sedimentary regimes. TAXONOMY AND MORPHOLOGY Introduction Taxonomic history of the Rauisuchidae began with Huene's work on the specimens he found in the Triassic of Brazil. In a short paper on thecodont relationships (Huene, 1936), he proposed the subfamily Rauisuchinae as a part of the family Stagonolepidae to include tlie genera Rauisuchus and Prestosuchus from Brazil. Later, (Huene, 1942) the group was ele- vated to familial rank and the genus Rhadinosuchus, also from the Triassic of Brazil, was included. At about the same time (Huene, 1938), he described Stagono- Saurusuciius and the Rauisuchid Thecodoxts • Sill 319 comes 10 K ilometers Figure 1. Generalized geologic map of the southern portion of the Ischigualasto Basin. Saurosuchus localities. marks 320 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 suchus from the Manda Beds of East Africa and noted its similarity to the Brazilian forms. However, not until 1956 did Huene formally place StagonosucJius in the family Rauisuchidae, at which time he also in- cluded a number of poorly known theco- donts that are no longer considered to be closely related to the family. Since then, intei"pretations of tlie broader relationships of the Rauisuchidae have followed the general pattern of uncertainty that has been the hallmark of thecodont taxonomy. Huene ( 1956 ) continued to maintain the family in close association with the stagonolepid-aetosaurid groups and included in the family such diverse genera as Cerritosaurus and Episcoposau- riis. Romer (1956) was the first to separate most of the genera of the Rauisuchidae from the armored thecodonts, and tenta- tively placed Raiiisuchus, Prestosuchus, StagonosucJius, RJmdinosuchus, and Pro- cerosuchus in the Ornithosuchidae. Hoff- stetter ( 1955 ) retained the family in the Stagonolepoidea, but removed Stagono- suchus to the Stagonolepidae. Reig (1961) presented a comprehensive review of the family and showed beyond reasonable doubt that the family Rauisuchidae should consist only of the genera Rauisuchus, Prestosuchus, Stagonosuchiis, and the then recently discovered Saurosuchus from Is- chigualasto. He also presented convincing evidence showing that the family is not closely related to the Stagonolepidae, and placed it in the "traditional" thecodont group which he termed Ornithosuchia (the equivalent of Pseudosuchia of most au- thors ) . Hughes ( 1963 ) , on the other hand, tentatively placed Rauisuchus and Sauro- suchus in the primitive thecodont group Proterosuchia as members of the Erythro- suchidae, a ranking that has not been accepted by the majority of paleontologists. Ticinosuchus, on the basis of a complete skeleton, was added to the family by Krebs ( 1965 ) ; its affinities with the other mem- bers of the family as described by Reig are evident, A further genus, Luperosuchus, from the Chaiiares Formation, was added to the family by Romer (1971a), and a closely related form has recently been found in the Los Colorados Formation (Bona- parte, personal communication). These latter discoveries are especially significant, for they permit the Saurosuchus lineage to be traced through the major part of the Triassic in a single basin of deposition. Romer ( 1966 ) followed Hughes in tenta- tively associating Rauisuchus and Sauro- suchus with the Erythrosuchidae, and adopted the term Prestosuchidae from Charig's unpublished thesis for the remain- ing genera Prestosuchus, Procerosuchus, "Mandasuchus"^ and, tentatively, Stagono- suchus. However, he later ( 1968 ) replaced Rauisuchus and Saurosuchus with the above mentioned forms, but did not sup- press Prestosuchidae. Meanwhile Presto- suchidae was carried on by Charig- ( 1967), who notes that the group is essentially the same as the Rauisuchidae of Huene (1942) but with the genus Rauisuchus excluded. In a more recent work on thecodont taxonomy Romer ( 1972a ) continued to use the family name Prestosuchidae on the grounds that Rauisuchus was too poorly known. However, he included Rauisuchus within the family Prestosuchidae (see Dis- cussion with regard to the affinities of Rauisuchus) . Assignment of tlie Rauisuchidae to a suborder is difficult given the present un- stable nature of thecodont taxonomy. Romer ( 1972a ) places the family with the primitive thecodonts in the Proterosuchia; other authors, Charig (1967) and Bonaparte (1971) place it in the usual "catch-all" sub- order Pseudosuchia. Rauisuchids certainly appear to have been derived from the erythrosuchid lineage of the Proterosuchia (see discussion on thecodont phylogeny). ^ Mandasuchus is technically a nomen nudum, as it has never been described in print. - In Charig's paper, origin of the Prestosuchidae was ascribed to Charig 1967. This paper has not been published. In an erratum, the family name was given as Romer 1966. Saurosuchus and the Rauisuchid Thecodonts • Sill 321 but they are much more spcciaHzed and progressive than any of its known members. On the other hand, they do not ha\'e a great deal in common with the "tv'pical" ornithosuchid pseudosuchians. As thecodont relationships become more clearly under- stood, a new suborder will probably have to be erected for this and perhaps other lineages descended from the erx'thro- suchids, but at present such a step would be premature. Discover}' of nearly complete remains of Ticinosuchtis and Saiirosrichus, represent- ing what appear to be the earliest and the latest members of the lineage so far de- scribed, has provided the means for an accurate characterization of the family. Basically, the new evidence tends to con- firm the definitions of the family given bv Krebs (1965) and by Reig (1961): Reig's paper provides an excellent summary of the taxonomic histor\' of the family and of the Thecodontia in general. The family ma}' be defined as follows: Medium- to large-sized carnivorous qua- drupedal thecodonts. Skull large, deep, orbit keyhole-shaped, large elongate antorbital fenestra, small crescent-shaped accessor}' antorbital fenestra present in some genera, teeth flattened, recurved, serrated. Pelvis triradiate, acetabulum closed, ischium elongated and rodlike, fused at the midline along most of its length. Femur long, slightly sigmoid, without a well-defined fourth trochanter. Calcaneum and astraga- lus articulate by a ball and socket joint, the socket on the calcaneum, the ball on the astiagalus. Five digits, fifth metatarsal short and hooked. Many of these features are generalized characteristics of the primi- tive thecodonts which have been carried over in the familv and are retained throughout their knowii history. Family distribution. Middle and Late Triassic; Argentina, Brazil, East Africa, Switzerland, possibly China. Family Raui- suchidae Huene 1936 (as a subfamily); genera Rauisuchus Huene 1936 Brazil, Pres- tosuchus Huene 1936 Brazil, Stagonosuchus Huene 193S East Africa, Saurosuchus Reig 1959 Argentina, Ticinosuchus Krebs 1965 Switzerland, Luperosuclius Romer 1971 Argentina, "Mamlasuchus" unpublished thesis Charig 1956. A number of additional genera are sometimes included in the family (see Romer, 1966 and 1972), but they are not well known. These additional genera are: Cuijosuchus, HopUtosaurus, Rhadinosuchus, Pallisteria, Spoiulylosoma, Procerosuchus, Fenhosuchus. Saurosuchus Reig 1959 Type species. Saurosuchus galiJei. Di.striJ)ution. Late Ladinian or Carnian, Ischigualasto Basin, Western ^\i-gentina. Diagnosis. As for the species. Saurosuchus galilei Reig 1959 Type. PVL 2062, nearly complete skull, posteriormost portion missing. Hypodigm. The t}'pe and: P\T. 2198, partial maxilla, left ilium, both ischia, nine articulated dorsal vertebrae and fragments, part of the dermal armour, associated ribs and teeth. PVL 2557, two dorsal vertebrae, both sacrals, nine caudals, right ilium and ischium, partial pubis, parts of right femur, tibia, fibula, complete right tarsus and foot, associated ribs and chevrons. PVL 2267, poorly preserved partial ilium, partial femur, tibia, fibula, well-preserved tarsus, partial foot. PVTL 2472, poorly preser\'ed cervical vertebra, tibia, astragalus. MSJ 102, fragment of maxilla and lower jaw. Horizon. Apparently all levels of the Ischigualasto Formation, San Juan province, Argentina. The five specimens of Sauro- suchus were collected from four localities, all in the soutliern portion of the outcrop area. The t\pe, P\'L 2062, consists of a nearly complete skull and was found in the upper third of the strata. The more complete skeletons, P\T. 2198 and PVL 2557, came from the middle part of the section, and the remaining two indi\'iduals, PVL 2267 and 2472, wer(> found in the lower third of the strata, as was MSJ 102 (see map for specific localities). 322 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Emended diagnosis. Large carnivorous l^ut the occipital region and braincase are thecodonts, up to six meters in length. Skull lacking. A fragment of the right maxilla deep, elongate, finely sculptured, with of PVL 2198 is identical to the correspond- keyhole-shaped orbit, large antorbital ing region of the type and allows assign- fenestra elongated anteriorly, small cres- ment of the specimen to the genus with a cent-shaped accessory antorbital fenestra considerable degree of confidence. The present between premaxilla and maxilla, lower jaw is known only from a fragment. Large elongate, nearly vertical external The skull is long, approximately 65 centi- nares bordered only by premaxilla and meters, triangular in shape, and .stiu-dily nasal. Teeth robust, recurved, laterally com- constructed. The cranial table is high and pressed with serrate edges. Four teeth on narrow. Orbits are large, keyhole-shaped premaxilla, ten on maxilla. Strong orbital openings, of which the upper part is a arch fomied by the frontal, small supra- well-defined circle high up the side of the temporal fenestra lying in dorsal plane of skull. A large antorbital fenestra is present, the skull below the crest of the orbital arch, subtriangular in shape and slightly smaller Vertebrae amphicoelous, spines broad and than the orbit. It is surrounded by a well- flat with prominent interspinous notch on defined smooth border set in from the anterior face. Cervicals apparently elon- sculptured surface of the maxilla. An un- gated, dorsals strongly compressed laterally, expected feature is the presence of a nar- rib facets well separated and on different row accessory antorbital fenestra located levels throughout column. Two sacral between the maxilla and the premaxilla, vertebrae. Shoulder girdle and forelimb extending from above the tooth-bearing unknown. Pelvis with closed acetabulum, surface to the posterior tip of the external pubis almost excluded. Ilium with broad nares. brevis shelf, ischium long, rodlike, ex- Like the antorbital fenestra, the external panded at the tip and fused at the midline nares are subtriangular in shape, relatively along most of its length. Femur slightly large, and situated principally in the verti- sigmoid, without a large greater trochanter, cal plane of the skull. Notable for their and with a small fourth trochanter. Fibula small size are the supratemporal fenestrae, bears a prominent iliofibularis tubercle, which lie in the horizontal plane of the Tarsus of the "crocodiloid" type, calcaneum skull roof just behind and slightly below bearing a large tuber and a prominent the heavy orbital arch. Only the anterior medial socket for articulation of the astra- border of the infratemporal fenestra is galar ball. Facets for articulation of the preserved, but it indicates a triangular or tibia and fibula close together. Fourth subrectangular shape approximately the tarsal large, subtriangular with prominent same size as the orbit, rounded facet for articulation of fifth meta- The large size of the skull and its sturdy tarsal. Five digits on foot, first two most construction indicate that Saurosiichus was robust, third is the longest, fifth is broad, an active predator. Using the head size flat, and oriented outward. Demial armour index of skull length to length of the pre- present, two rows of small scutes along sacral vertebral column, a value of either each side of most of the vertebral column, .27 or .34 is obtained, the latter calculated leaf-shaped and imbricating. on the assumption that neck vertebrae were _ , _ . ^, approximatelv the same length as the General Description / , , .; ,, „ ^. dorsals, while the smaller ratio assumes Skull elongated cervicals. Both indices are in Cranium. Cranial material is represented the range of the large predaceous dino- almost exclusively by the type, in which saurs; AUosaurus is .28, Tyrannosaunis most of the dermal elements are preserved, is .41, Saurosuciius and tiie Rauisuchid Thecodonts • Sill 323 Prcinaxilld. Both promaxillae of tlic type arc complete and well piescived. The main body of the bone is a massive rectangle from which a slender process extends up- ward and backward around the external naris to a long o\'erlapping contact with a similar process of the nasal, and a second rodlike extension that forms the entire lower border of the naris and terminates wx^dged between the nasal and the maxilla. At its anterior border the premaxilla forms a straight \'ertieal line from the tip of the naris to the first tooth position. Below the narial opening the bone swells to a thick, slightly undulating ridge that bears four large teeth. At the most anterior part, just above the toodi row, lie three foramina. No sculpturing is present. The rodlike process that forms the lower border of the naris is an isolated structure that separates the accessory antorbital fenestra and the external naris. Medially, the premaxillae meet in a long sturdy symphysis. The ah'eolar margin is thick and slightly vaulted behind the first two teeth. Of the four teeth, the third is the largest. Two deep pits are present in the ^ aulted area, one beside the second tooth, the other between the third and the fourth. A large foramen is present above the third alveolus. The interalveolar septum between the third and fourth teeth is ex- panded on the lingual surface to form a small interdental plate. Posteroventrally, a clearly defined suture is not present between the maxilla and the premaxilla, but above the thick tooth-bear- ing portion of the bone the accessory antor- bital fenestra serves to separate the two elements. Maxilla. The maxilla is a large platelike bone that slopes posteriad and upward from its suture with the premaxilla to meet the nasal and lacrimal dorsally and the jugal ventrally. It is deeply emarginated by the antorbital fenestra, around which runs a broad smooth shelf. Outside the shelf area the maxilla is heavily sculptured by an irregular network of grooves. It fonus Tahle 1. Measurements of the skull (in centimeters) of SAUROSUCHUS GALILEI BASED ON the type PVL 2062. Note, further preparation HAS modified some OF THE MEASUREMENTS MADE HY ReIG (1959) IN HIS Pl^ELlMINAUY ACCOUNT. Total length of the skull (estimated) 67 Length from tip of snout to anterior border of the supratemporal lenestra 54 Lengtli from lower anterior corner of infra- temporal fenestra to tip of snout 47 Diameter of tlie upper portion of the orbit ... 10 Maximum lieight of the orbit _._ 17 Maximum lengtli of the antorbital fenestra - . 19 Maxinnun height of the antorbital fenestra .- 8 Maximum length of the depression smround- ing the antorbital fenestra 21.5 Maximum height of the depression surround- ing tlie antorbital fenestra __- 10 Nhiximum lieight of the skull betA\'een top of the rim of the orbit and bottom of jugal — . 20.5 Length of nasals along tlie midline 32 Length of tlie preniaxil!ar>' tooth row 9 Length of tlie niaxillar\' tooth row 27 Length of the external naris 12 Distance from tip of snout to anterior border of the antorbital fenestra 21 W'idtli of skull across the supratemporal fenes- trae ---.. 17 Widtli of skull in front of the orbits 10 Length of teeth alveoli Premaxilla Left Right 1. 1.5 — 2. 1.5 1.6 3. 2.0 1.8 4. 1.3 1.5 Maxilla 1. 1.8 1.5 2. 2.4 2.3 3. 3.0 3.0 4. 2.6 2.3 5. 2.7 2.4 6. 2.2 2.3 7. 2.3 2.1 8. 1.9 — 9. 1.8 — 10. 1.8 — Length of maxillar\- teetii, left side, from lateral edge of the maxilla to die tip of the teeth Anterior Posterior Tooth No. curvature curvature 3. 4.6 3.5 5. 5.8 4.7 6. 3.5 2.5 7. 5.0 3.9 8. 3.9 3.1 324 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 CO o Q. ■1 Q. To c s (D a CO E o CO c o (0 D) 3 CO m CO c I Z X E X (0 E eo 3 u 3 0} o 3 CO CO (0 ^ o (D D. ^i Q- 5, <= D) o Saurosuciius and ttik Rauisuciiid Thecodonts • Sill 325 the entire ventral, and half of the dorsal union with the frontals, from .5 to 1.5 cm. borders of the antorbital fenestra, meeting Tlie lateral component is not extensive and the lacrimal in a broad overlapping snture disappears entirely at the beginning of the on the smooth shelf portion, and the jugal antorbital fenestra. At their maximum in a broad zig-zag digitate union. Ten teeth width the joined nasals are approximately were pr(\sent on the maxilla, of which seven seven centimeters wide, an indication of were apparently functional at any one time, the narrowness of the anterior portion of Numerous foramina are present on the the skull. Sculpturing on the nasal is in the lateral surface just above the tooth row. fonn of irregular longitudinal grooves. On the medial surface the most promi- Union with the maxilla and lacrimal is in nent feature of the maxilla is the formation a straight sloping line. The suture with the of the alveoli by large interdental plates, frontal is an inverted V located at the level The plates are leaf-shaped extensions of the of the posterior border of the antorbital alveolar septa and slightly overlap one fenestra. another at the middle of the tooth body. Prefrontal. The area corresponding to Above the plates a prominent groove runs the prefrontal is badly fractured, but this to the dental lamina, which slopes down- element appears to be a small platelike ward posteriorly to terminate on the ventral bone lying in the horizontal plane above surface of the maxilla just behind the last the lacrimal. It does not participate in the tooth. From the groove foramina represent- orbit, but sutures are difficult to distinguish, ing the replacement teeth open directly Lacrimal. Most of the lacrimal lies in above each tooth. This morphology ap- the depression surrounding the antorbital parently represents a standard pattern of fenestra and is therefore completely smooth, tooth replacement, analyzed by Edmund as is that part of the maxilla that partici- (1957, 1960), in which the fibrous con- pates in the same depression. The lacrimal nective bone that surrounds the tooth is is an extensive thin plate, forming most of partially resorbed during the replacement the smooth shelf around the upper part of process to form the shield-shaped inter- the antorbital fenestra. Anteriorly it is dental plates. overlapped by the maxilla. Posteriorly it Above the tooth row, in the anterior forms a ventral prong that overlies the portion of the maxilla, a massive buttress dorsal extension of the jugal to fonn the projects medially to meet the vomer and preorbital bar. The border between lacrimal form part of the vault of the premaxillary and prefrontal is not discernible, but must chamber. Dorsally, on the medial surface, lie in the zone behind the smooth depres- the maxilla terminates in a straight sloping sion of the fenestra. This area is thick and contact with the nasal and the broad over- heavily sculptured, and from it arises a lap of the lacrimal. Posteriorly the jugal is prominent lateral ridge that runs down the laminated to the maxilla just above the surface of the preorbital bar, terminating tooth row. A large maxillary foramen is at the tip of the ventral prong of the bone, present just anterior to the jugal suture The lacrimal forms the upper third of the midway between the tooth row and the posterior border of the antorbital fenestra, ventral border of the antorbital fenestra, on and virtually all of the anterior border of the medial surface. the orbit. There is no definite lacrimal Nasal. Anteriorly, the nasal is a thin bar foramen, but a rounded depression is above the external naris overlapping the present on the ill-defined internal border similar element of the premaxilla. From between the lacrimal and the frontal. There this position it broadens to a dorsal and is no transverse component of the lacrimal, component thickens considerably near its Jugal. The large skull openings of Satiro- lateral plate of bon(>. Posteriorly the dorsal suchiis have reduced the jugal to a hori- 326 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 CO o Q. I o Q. c o in o Q. CO E *^ u CO c o I (0 D) 3 CO CO CO c I z CO E JO ■>< CO E 0) 00 o 3 CO O CO CO 3 CO a> 5 o "> 15 Xl 0) ;S X CO a? M 3 *- O) J3 U- o Saurosuchus and the Rauisuchid Thecodonts • Sill 327 zontal rod \\'itli two dorsal prongs projecting npwards to form parts of the pre- and postorbital bars. It is most ex- panded anteriorly where it is platelike and overlapped by the maxilla in a prominent zig-zag snture. Immediately behind this union, the dorsal projection of the pre- orbital bar reaches up medial to the narrow ridge of the lacrimal. Behind the orbit the second prong of the jugal extends upward and backward to form a strong, sloping, abutted contact with the \'entral expression of the postorbital. At the ventral border of the infratemporal fenestra, the jugal is a relatively narrow uniform bone. It thins out to a fine edge on its lateral surface, showing clearly the area where it was overlapped by the quadratojugal. Sculptur- ing is present only in the anterior portion of the bone, where it meets the maxilla, and even that is light. Only the ventralmost part of the orbit is formed by the jugal, but it constitutes nearly all of the anterior border of the infratemporal fenestia. Di- rectly below the postorbital bar there is an outward bulge in the jugal, fomiing a distinct pocket on the internal surface, possibly the contact for the ectopterygoid. Frontal. The frontal is a thick strong bone dominated by the massive supra- orbital arch. Medially it curves down from the arch to the midline where its posterior portion meets the anterior projection of the parietal. Anteriorly it joins the nasal and prefrontal in a zig-zag suture. Sculpturing is present, principally on the arch, and is of the pit and groove variety. The thickest part of the frontal is the area of the mid- line, which in the type is two centimeters deep. Internally there does not appear to be an interorbital septum, but the orbit is well defined by the medial continuation of the orbital arch. Anteriorly the arch forms the previously mentioned pocket at its junction with the lacrimal. Anterior to the orbit the frontal thins to slightly over one centimeter in thickness, and bears a down- ward-projecting ridge near the midline. This ridge, presumably the border of the olfactory tract, is eight millimeters high at its maximum and tapers off to the level of the bone at the anterior end of the frontal. Behind the orbit, at the junction of the frontal, parietal, postfrontal, and post- orbital, a prominent circular pocket is present. This most probably received the anterior process of the laterosphenoid. Postfrontal. This is a small semicircular bone lying on the dorsolateral surface of the skull between the frontal and the post- orbital. It does not enter into the supra- temporal fenestra. On the ventral surface of the skull it is not possible to distinguish the borders of the postfrontal. Postorbital. The postorbital forms nearly all of the posterior border of the orbit, and the upper third of the anterior border of the infratemporal fenestra. Dorsally, just behind the orbital arch, it bears a promi- nent, sculptured tuberosity. Ventrally, it meets the ascending process of the jugal in a long diagonal contact. The anterior border of the postorbital bar is emarginated and beveled at the delimitation of the circular part of the "keyhole" orbit. On the cranial table the postorbital fomis most of the lateral and anterior border of the small supratemporal fenestra. A well-defined, smooth margin surrounds this fenestra, otherwise the upper region of the post- orbital is sculptured by linear grooves. Internally, the anteromedial portion of the postorbital forms the rear part of the socket for the laterosphenoid articulation. The posterior part of the postorbital is not pre- served in the type. Nothing remains of the cranial table be- hind the frontal and postorbital bones in the available material. Palatal Complex Palatal remains of Saurosuchus are not well preserved, but allow reconshuction of the major features. A primiti\'e character of the palate is the long triangular inter- pterygoid vacuity. The internal nares are somewhat displaced toward the rear and close to the sides of the maxillae. No traces 328 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Figure 4. Palatal view of the skull of Saurosuchus. Ec — ectopterygoid, Pt^ — pterygoid, PI — palatine, V — vomer. X 1/4. Saurosuchus and the RAtnsucHiD Thecodonts • Sill 329 of teeth are found on the palatine or on the pterygoid. Although crushed, the palate appears to have formed a deep vault rather than a shelf. The basicranium is not known. Pterygoid. As usual, the pterygoid is the largest of the palatal bones, and is divided into the customary three components: flange, palatal, and quadrate rami. The palatine ramus consists of a broad thin plate of bone that extends forward from the base of the flange portion and narrows anteriorly to a V-shaped ridge that meets the vomer near the midline. The medial border of the pterygoid is formed by a rounded ridge and steep shelf of bone that form the edge of the interpterygoid fe- nestra. Only at the anteriormost tip do the pterygoids join at the midline. On the dorsal surface of the palatal ramus a deep groove is present just lateral to the wall of the intei-pter)'goid vacuity. This groove may continue onto the vomer. The flange portion of the pter>^goid is massively con- structed, and bears a thick, rounded posterior border that curves out to fonii the "wing." Where the wing meets the heavy ridge that borders the interpterygoid fenesti'a a deep pocket is formed. Postero- medial to this pocket lies a thick remnant of the basipterygoid articular bar. An- teriorly the flange thins considerably, be- coming the same thickness as the palatine. At the posterior margin the flange is 15 mm thick, while anteriorly it is only 4 mm. The angle of inclination of the flange is approxi- mately 45 degrees. Ectoptenjgoid. Only the massive portion of the ectopterygoid that forms the lateral border of the pterygoid flange is preserved. This portion forms a strong buttress along the entire lateral edge of the pterygoid "wing." There is no identifiable scar on the maxillae or jugal to indicate the articulation of the ectopterygoid, although it seems probable that the bulge just below the postorbital bar was for reception of the ectopterygoid strut. The massive nature of the preserved portion of the bone indi- cates that the ectopterj^goid served to strengthen the lateral part of the pterygoid. Palatine. As preserv^ed, the palatine is a thin plate, not possessed of unusual char- acteristics. Anteriorly it forms tlie posterior half of the internal naris; the suture with the vomer is well preserv^ed. Laterally it is applied to the side of the maxilla, opening posteriorly into the pterygoid fenestra. The medial border is not well preserved, but appears to have been of the usual platelike contact with the pterygoid. Vomer. The vomer is poorly preser\'ed and represented only by a distorted and ill- defined mass of bone anterior to the in- ternal nares. As near as can be detemiined, the vomer formed the anterior half of tlie internal naris, above which it expanded considerably in the form of a laminar sheet of bone applied to the medial side of the massive maxillary buttress. Possibly, a por- tion of the vomer behind tlie maxilla formed a secondary buttress behind tlie laminar part. Dentition. Most of the 14 sturdy teeth in the upper jaw are of equal or nearly equal size. In the premaxilla the teeth are not preserved, but to judge from the size of the alveoli, the first and fourth teeth are slightly smaller than the second and third. In the maxilla, the last three teeth show a slight reduction in size compared with the anterior ones. All of the maxillary alveoli show the presence of functional teeth, with the possible exception of the first two, although at least two and possibly three growth stages are represented. The third, fifth, and seventh teeth are the largest, with the fifth slightly larger than the others. The fourth, sixth, and eighth are approximately the same size. The ninth and tenth teeth are broken off at the alveolar border, but were similar to the eighth in size. All of the teeth are of similar shape, heavily constructed, laterally compressed, sharply pointed, and recurved. The last three or four of the maxillary series seem to be more stiongly recurved than the anterior mem- bers, but this may be due to defomiation. 330 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Roots of the teeth are approximately twice parallels the development found in some as long as the crown. Near the alveolar prosauropods, and seems to be related to margin, the teeth are much more com- size. pressed and elongate than in the main body Cervical vertebrae. Only one cervical is of the crown, and on the fully erupted teeth known, PVL 2472. It is poorly preserved a slight depression is present on the lingual and of questionable reference to Sauro- face of the tooth near the margin formed by suchus. It was found in association with a the alveolar septum (see Plate 1). The teeth tibia and astragalus, also poorly preserved, are essentially symmetrical, but the plane that appear to be identical to those of PVL of symmetiy, taken between the anterior 2267. However, the unusual features of the and posterior serrations, is slightly rotated vertebrae warrant its inclusion in this study anteromedially-posterolaterally. Enamel on even though its association with Saurosuchus the crown is thin and not striated. is not completely reliable. Only the centrum Serrations are present on the distal three- is preserved; it is an elongated, flattened quarters of the anterior edge and along all structure generally constricted in the of the posterior margin. However, this con- middle. The anterior ( ? ) face is sti'ongly dition can be fully appreciated only on the concave and bears a proti-uding lower mar- fully erupted teeth; in those teeth that have gin that would seem to indicate a cervical not reached maturity the serrations con- flexure. The rear (?) surface is only tinue to the alveolus. Form of the serrations slightly concave. There is no keel. In the is the same on both edges; they consist of middle portion, the body of the centrum is simple crosscuts perpendicular to the long not only constricted laterally, but is also axis of the tooth. Density of the serrations greatly flattened, which transforms the is 12-14 per 5 mm, and is the same on both whole into a very lightly built structure, the anterior and posterior edges. There are Prominent pleurocoels are present just be- no wear facets on the teeth, although the hind the flared articulating surfaces. Thus larger ones have a somewhat more rounded the lateral border of the centrum is almost apex. a horizontal plate that curves inward to the narrow waist (see Fig. 5). The char- AxiAL Skeleton acteristics of the vertebrae represent an The exact number of presacral vertebrae ^^^"^^ development of a strong, lightweight is not known. Two vertebral series are support for the cervical region. As such it preserved, PVL 2198 and PVL 2557. The ^l,^°^P\^^^/^ "^u rl ^°^^^V . former consists of nine dorsals, all of which ^^ i' f^'l presence of bore ribs; the latter series is from the sacral Pleurocoels are not found on any of the and caudal region and does not duplicate ^^^°^^^ ^^^^f^ vertebrae. This condition any of the vertebrae of the PVL 2198 series. ^'''^^^' f ^^^^^^^^ ^^ ^^^^^ t^'lV In PVL 2557 the first two presacrals are ^^"^^ °^ ^^'' ^?^^"g^, ^^^T''i c Measure- I i 1 .1 . .1 . ments are as follows: length, 18 cm; width preserved and show that ribs were not ,- , , . ^ ,, .i i r i . .1 A 4.U *- • u of the anterior surface, 11 cm; width of the present on these. As the anterior members , . . i . j , of the PVL 2198 series do not show char- fo^^tricted waist 4 cm; and approximate acteristics of cervical vertebrae, it seems ^'^'^^^ ^} ^^^^t, 2 cm. reasonable to assume that not more than ^^^^«^ veiiebrae. Vertebrae of the dorsal two vertebrae represent the gap between series are represented by the first two pre- the presacrals of the two series. Assuming sacrals of PVL 2557 and by nine articulated the usual presence of seven or eight cervi- members of PVL 2198. The anterior mem- cals, the vertebral count would fall into the bers of the PVL 2198 series are poorly 23 to 25 characteristic of archosaurs. In preserved. The most striking feature of the general, structure of the vertebral column Saurosuchus vertebrae is the reduction of Saurosuchus and the RAuisucmo Thecodonts • Sill 331 B Plate 1. A. Type of Skull of Saurosuchus, PVL 2062. X Vs. B. Lingual view of left maxilla, note interdental plates. X 1. C. Enlarged view of a recently erupted tooth, showing serrations on the posterior edge. X 8. 332 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 i Table 2. Measurements of the vertebral column of Saurosuchus galilei (in centimeters). PVL 2472 cervical (?) Maximum lengtli 20.0 Transverse width of posterior face 10.5 Transverse width of anterior face 9.0 Minimum width of constricted waist measured on ventral surface 4.0 Height of anterior face 5.3 Height of centrum body at waist 2.0 Dorsal vertebrae PVL 2198 Anterior Posterior Maximum length of centrimi 7.5 8.5 Height of anterior rim 8.7 Widdi of anterior rim 7.2 Width of posterior rim 6.0 Total height of vertebrae 20.0 22.0 Minimum width of constricted waist 1.2 1.3 Lateral extension of transverse process from the midline 5.5 4.8 Width of neural spine table 3.0 2.4 Diameter of parapophysis 2.5 2.5 Diameter of diapophysis 2.7 2.7 Height of neural arch above centrum 12.6 13.0 Lumbar and sacral vertebrae PVL 2557 Presacral 1 Sacral 1 Sacral 2 Length of centnmi 9.0 10.5 10.0 Height posterior rim 12.0 9.5 10.0 Width posterior rim 10.5 10.0 8.5 Height anterior rim 11.0 9.5 Width anterior rim 12.0 10.0 Minimimi width of constricted waist 4.0 3.7 3.1 Width of neural spine table 4.5 3.6 Height of neural arch above centrum 17.6 19.5 Total height of vertebrae 28.0 29.5 Caudal vertebrae PVL 2557 Caudal 3 Caudal 9 Length of centrum 8.0 7.5 Height posterior rim 9.5 7.0 Height anterior rim 9.0 7.0 Widtli anterior rim 9.0 6.0 Width posterior rim 9.0 6.0 Minimum width of constricted waist 3.9 2.3 Width of transverse process from midline 11.5 7.0 Chevrons PVL 2557 No. 1 No. 2 No. 3 No. 4 No. 5 Length 18.0 21.0 21.5 21.2 18.5 Width between articulations 7.5 7.5 6.7 6.2 6.0 Width of articular facets 4.0 4.0 4.0 3.0 2.5 Length from facet to fusion with opposite side 4.5 3.8 4.0 3.5 3.2 the centrum to a thin vertical plate between specialization of the genus. Another notable the flared rims. This condition is not as feature of the dorsals is the complete sepa- well developed in other thecodonts and ration between the diapophysis and the seems to be a unique weight-reduction parapophysis along the entire known series; Saurosuchus and the Rauisuchid Thecodonts • Sill 333 tlic latter is always found on the neural arch, never on the centrum. Transverse processes are larger in the anterior region than in the posterior, but all are rather short and stubby. The neural arches sit high on the centra and bear flat rectangular spines that are not inclined posteriorly. Size and shape of the vertebrae appear to be uniform throughout the series. All centra are uniformly amphycoelus and do not bear keels. Morphologic changes along the series are not prominent and consist principally of the reduction of the transverse processes in the lumbar region. In end view the centra are oval-shaped with the long axis in the vertical plane. The rims are flared and rounded, not beveled. Reduction of the body of the centrum took place by expansion towards the rims of the common "hour-glass" constriction. The re- sults are a steeper angle of the constriction behind the rims and the formation of a narrow plate between them (see Fig. 6). No ridges, rugosities or excavations are present on the body of the centrimi. Length of the centrum is 7 to 8 cm, width 6 to 7 cm, height is around 9 cm. The body of the centrum expands slightly to receive the neural arch and form the floor of the neural canal. The neural arch is a large structure that sits high up on the centrum. Contact with the centium is a simple butt union, without the formation of pedicels. Prezygapophyses are not well preserved, but form short processes that sweep forward on either side of the prominent interspinous notch of the neural spine, just above the articular facet for the capitulum. Apparently the prezyg- apophyses did not overhang the border of the centrum. The postzygapophyses lie on the same level as the transverse process and are formed from lateral expansions that diverge from the base of the neural spine, creating a wedge-shaped cleft behind it. The zygapophyseal facets are relatively small smooth areas facing downward with a .slight inclination toward the midline. Figure 5. Supposed cervical vertebra of Saurosuchus. Top, ventral, Bottom, dorsal. X Va. Rib articulations are restricted entirely to the neural arch. The parapophysis is a round facet on the anterior vertebrae of the dorsal series, but becomes laterally ex- panded into a peduncle on the posterior vertebrae. On all of the dorsals the parapophysis lies below and in front of the transverse process. These processes ai'e short and robust; those of the shoulder region are larger than those of the lumbar series and project posteriorly approximately 30 to 45 degrees. The diapophysis forms as an expanded foot at the tip of the trans- verse process. On the anterior dorsals this expansion is considerably larger than the parapophyseal facet, while in the lumbar region it is of the same size. A notable featui'e of the transverse process is the presence of strutlike ridges on the under- 334 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Figure 6. Three views of a dorsal vertebra of Saurosuctius. Left, right lateral; middle, posterior; right, ventral. X 1/4. side and edges. In the shoulder region, where the transverse process is largest, four struts are present. One extends to the prezygapophysis, another to the postzyg- apophysis, a third down to the parapophysis, and a fourth down and back to the rim of the centrum. All of the ridges extend the entire length of the transverse process, and form a strong supporting structure. In the posterior dorsals the strut structure is modi- fied by a reduction to three struts. The parapophysis has moved slightly dorsal, al- most to the level of the prezygapophysis and the transverse process is smaller. Only one ridge is present in the anterior portion, extending from the transverse process to the parapophysis. The shorter transverse process and the lateral expanded para- pophysis change the aspect of the support- ing strut from that of a ridge to a sheet of bone (see Fig. 6). The neural spine on all of the dorsal vertebrae is a robust rectangular blade, slightly higher than the centrum. As well as can be determined, the blade is not in- clined posteriorly on any of the vertebrae. On its dorsal surface the spine is expanded into a spine table, presumably for the at- tachment of dermal armour. At the anterior and posterior borders the spine does not attenuate, but bears prominent grooves for the interspinalis musculature. On the lead- ing edge the groove occupies the lower half of the length and is deeper at the base. The groove on the posterior margin is shal- lower but extends the entire length of the blade. A distinct lumbar region was present in Saurosuchus, but it is not possible to de- termine the number of vertebrae involved. Specimen PVL 2557 has preserved the t\vo vertebrae immediately anterior to the sacrum, and these vary from the other dorsals principally in their lack of normal ribs. It is not possible to determine whether the short downcurved processes are ribs or transverse processes. They appear to be transverse processes, arising from the same Saurosuchus and the Rauisuchid Thecodonts • Sill 335 position on the neural arches as those of the anterior dorsals. The processes are oval in cross section and heaxily constructed. Their origin on the arch is considerabh' broader than that of the dorsals of PVL 219(S. From the arch they curve slightly for\\'ard, then strongly downward. Sacral vertebrae. The sacral \'ertebrae are known exclusixely from the well-preserved representatives of PVL 2557. Two sacrals are present in Saurosuchus. The centra are slightly more elongate than the other dorsals, but otherwise are not different. The sacrals are not fused, but there is a con- siderable reduction of the rims where the two meet, with the posterior rim of the first sacral flared out at the sides and a corresponding reduction and slight forward extension of the anterior rim of the second sacral. This imparts a slightly V-shaped configuration to the union between the two vertebrae. This condition is repeated in the junction between the last sacral and the first caudal. Such a union must have essentially immobilized the three vertebrae in\^olved, providing a partial substitute for the fusion of the sacrals. The transverse processes of the first caudal vertebrae are not preserved, so it is not possible to de- termine if it participated in supporting the pelvis. Position and shape of the transverse processes of the sacrals are essentially of the type found in primitive archosaurs; the first is large, oval-shaped, and positioned near the anterior border of the centrum, while the second is more crescent-shaped and arises from the center of the centrum. Both are impressive structiu-es, greatly en- larged and heavily constructed. Neural spines and arches of the sacral vertebrae are not significantly different from those of the presacrals that form the lumbar region. The spines are heavily con- structed and expanded, but form a well- matched series with those of the lumbars. The same is true for the neural arches. It should be pointed out that the verte- brae of specimen PVL 2557 do not have the centra constricted nearly as much as those of specimen PVL 2198.' Whether this dif- ference is due to the difference in size be- tween the two animals (PVL 2557 is considerably larger than PVL 2198) or to their different positions in the vertebral column cannot be ascertained. Caudal vertebrae. In general the caudals of Saurosuchus are of shorter length than the other vertebrae, and ha\'e large rounded rims. The first three caudals do not bear chevrons. Diameter of the centia of the first five caudals is essentially equal to that of the sacrals. Beginning with the sixth caudal, there is a gradual reduction in size. Rims of all of the caudal ^'ertebrae are broad and rounded compared to the some- what thinner rims of the other vertebrae. The area between the rims is not reduced as in the dorsals; the centra are more "typical" in their squat rounded shape. Beginning with the seventh caudal, a slight groo\'e appears on the ventral surface of the centrum. At the eighth or ninth the shape of the centrum changes to the more elongate and spool-like shape characteristic of tail vertebrae in general. Large lateral processes are present on the nine articu- lated caudals preserved in specimen PVL 2557. The processes of the first four or iive caudals are large bladelike stnictures that extend outward and backward from the level of the dorsal surface of the centra. In caudals numbers six and seven, the out- ward extensions of the processes are greatly reduced, but they retain the blade shape. In caudals eight and nine, the lateral pro- cess loses the bladelike expansion and be- comes a simple short lateral process. Neural spines of the first four caudals are large, cover the entire length of the centrum, and in general are like those of the dorsals. Beginning with the fifth caudal there is a relatively sharp reduction in the anteroposterior length, and in the height of the spine. The spine becomes more in- clined caudad and develops a more promi- nent interligamentum cleft in tlie anterior border near the base. The sides of the cleft 336 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Table 3. Measxjrements of the pelvis of Saurosuchus galilei (in centimeters). Ilium Length along dorsal border Lengtli of anterior spine from the ventral curvature Length of posterior spine from tlie ventral curvature Height of dorsal border above acetabulum Maximum height of ilium Widtli of dorsal border Ischium Total length along curvature Width of shaft Height of shaft Height of terminal expansion Widtli of temiinal expansion PVL 2198 PVL 2557 36.0 41.0 4.2 6.5 17.0 23.5 6.5 10.5 23.5 29.0 1.8 2.1 42.5 50.0 3.7 4.1 3.9 4.6 5.3 9.3 5.9 6.1 develop progressively into prominent ridges that sweep forward to form the prezyga- pophyses. Position of the zygapophyses undergoes a slight progressive shift towards the front; the prezygapophysis begins to overhang the centrum and is accompanied by a corresponding anterior displacement in the postzygapophysis. Chevrons. Chevron bones first appear on the fourth caudal. The first four chevrons are Y-shaped and bear large disclike pedi- cels for articulation with the vertebrae. The arms of the "Y" become progressively closer together until they join at the fifth chevron (eighth caudal), leaving a small opening of which only a vestige remains in the sixth chevron (ninth caudal). Construction of the chevrons is simple and not unusual; the expanded pedicel is followed by a long sturdy shaft fused with its opposite just below the centrum. A slight ridge is pres- ent on the posterior surface of the shaft. The six preserved chevrons are all approxi- mately the same length (equal to the total height of the vertebrae) and are strongly inclined caudad. Ribs. Few ribs are preserved. Those available are fragmentary and are covered with a thick iron-rich matrix. They appear to be heavily constructed, thick-bodied, and with a prominent ridge on the upper third of the anterior edge. The posterior surface (at least in the proximal section) is flat with a slight depression down the middle. Rib articulations appear to be well ossified. As may be expected, the largest ribs were the anterior members (size inferred from the relative size of the articular surfaces on the vertebrae). Appendicular Skeleton Pelvic girdle. The pelvis of Saurosuchus is well represented except for the distal portion of the pubis. Elements available are: left ilium and paired ischia of PVL 2198; right ilium, complete right and par- tial left ischia of PVL 2557; and a poorly preserved fragmented ilium of PVL 2267. The proximal portion of the right pubis of PVL 2557 is articulated with the ilium of that specimen. The usual elements of the pelvis were present, in typical triradiate forms. There is no indication of perforation of the acetab- ulum. A notable feature of the pelvis, apparently common to the Rauisuchidae, is the high position of the pubic articu- lation and the limited participation of this element in the acetabulum. Ilium. Two major structural divisions are present on the ilium; the acetabulum and the iliac blade. Most of the ilium is Saurosuchus and the Rauisuchid Thecodonts • Sill 337 •.'.v.'-'i n-'f'fi Hi!Ail!l-A ur B?HjWv^'«;V^«!>.^^■;.*^K'S^^^..■^■«■T«'■'^''^^''.^.^■^;!. ■' Figure 7. Ilium of Saurosuchus. x y4. incorporated into the acetabulum, which is a large deep depression that faces shghtly downwards. The dorsal margin of the acetabulum is formed by a thick lateral flange positioned just below the anterior emargination of the iliac blade. Ventrally the acetabulum wall thins considerably at its borders with the ischium and pubis. Anteriorly it expands transversely where it meets the dorsal border of the pubis, below which the bone thins, presenting a tear- drop shape in cross section. A notable feature of the ilium is its articulation with the pubis and ischium; the suture of the pubis occupies nearly all of the anterior border, starting from a level almost at the dorsal border of the acetabulum, whereas the ischium meets the ilium more in the ventral plane. The ilium is not constricted above the acetabulum. Rather, the anterior origin of the iliac blade arises from an emargination immediately above the thick dorsal margin of the acetabulum, while the posterior portion of the blade sweeps up- ward and backwards from a level slightly above the midline of the acetabulum. The anterior tip of the blade is short and tliick; it does not reach the anterior border of the acetabulum. The posterior portion of the blade consists of tliree prominent elements: 1) a rounded dorsal ridge, 2) a horizontal shelf on the medial side, midway between the dorsal and ventral borders, and 3) the very thick rounded ventral border of the blade. The internal shelf con-esponds to the structure termed "brevis shelf" by Romer (1927) and originates just behind the acetabulum, becoming considerably heavier and thicker at the terminal end of the blade. At its posterior tip the iliac blade is heavily constructed with the brevis shelf lying perpendicular to the blade. Rugosities present in the tip region indi- cate that it was probably continued in carti- lage. Facets for the sacral ribs lie just above the level of the acetabulum. Total 338 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 B Figure 8. Two views of \he paired isciiia of Saurosuchus. A, ventral; B, dorsal. X ''A. length along the dorsal border of PVL 2198 is 36 cm of which 16.5 lie below the acetab- ular rim. Thus the blade above the acetabulum is only 6 cm high. Ischium. The ischium of Saurosuchus consists of a broad flange followed by a relatively long shaft that bears a mild terminal expansion. In general it resembles somewhat that of the dinosaurs in that it is rodlike rather than platelike. Proximally the ischium bears a large expanded head with a prominent lateral lip. As usual, the anterior portion of the head is considerably thinner than the posterior. Anteriorly, be- low the lip is a deep concavity, where the bone becomes a thin plate that angles to- Saurosuchus and the Rauisuchid Thecodonts • Si7/ 339 Table 4. MEAsuREJ\rENTs of the hind limb of Saurosuchus galilei (in centimeters). PVL 2267 PVL 2557 Femur Appio.ximatc total lenj:;th Maximum width of proximal articulation Distance of 4tli trochanter from head Thickness of shaft at midpoint Approximate width of distal articulation Tibia Length Minim mn shaft \\'idth Widtli distal articulation Widdi proximal articulation Fibula Length approximate Minimum shaft width Anteroposterior \\'idth of distal articulation Transverse width of distal articulation Distance between distal articulation and ilio-fibiJaris trochanter Astragalus Maximum width across anterior face Maximum anteroposterior length Height on anterior face Calcanium Maximum anteroposterior lengtli Maximum height of tuber Maximum height of anterior face Maximum transverse width 65.0 7.0 17.0 26.0 5.0 9.0 PVL 2472 PVL 2267 PVL 2557 46.5 45.0 4.0 3.5 6.5 5.5 6.8 12.0 PVL 2267 43.0 2.7 8.5 4.0 21.0 PVL 2472 10.5 8.5 3.8 PVL 2267 12.5 7.5 3.8 8.0 PVL 2557 3.0 8.2 5.3 22.0 PVL 2557 10.6 9.6 5.0 PVL 2557 16.0 11.0 5.5 8.5 wards the midline, terminating in a smooth rounded border that does not touch its opposite member. Posterodorsally, the ischium is a heavy rounded strut arising from the thick buttress that forms the posteroventral rim of the acetabulum. Distal to the expanded head region the two ischia are solidly fused. The symphysis forms a slight ridge down the ventral (an- terior) surface of the paired bones. At the distal termination the ischium flares out to a moderately expanded foot, similar to that of Ticinosuchus, but more rodlike. In cross section the rod portion is tear-drop shaped, the thin portion being fused to its opposite. This fusion created a channel along the dorsal midline and a correspond- ing ridge along the ventral surface. The ischium makes an approximately 45-degree angle with the iliac blade, and the strut portion is slightly concave upwards. Pubis. Only the proximal portion of PVL 2557 is known. It shows that the dorsal portion of the articulation with the ilium was a very thick continuation of the an- terior border of the ilium beneath the blade. Below this thick rounded border, the pubis thins rapidly, matching the thick- ness of the ilium. As noted previously, only the edge of the pubis actually participates in the acetabular depression. From the cross section of the broken portion of the 340 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Figure 9. Femur of Saurosuctius, composite drawing. X 1/3- pubis, it would appear that the bone thinned considerably in its anterior portion below the rounded dorsal margin. Femur. The femur is known from two nearly complete specimens. The complete proximal half of the femur is well preserved in specimen PVL 2557, and was found articulated with the corresponding pelvis. It is well-preserved material but appears to be slightly compressed. PVL 2267, the other femur, consists of a complete shaft but lacks the extreme articular surfaces at both ends. This specimen was figured by Reig (1961) and shows a slight intertro- chanteric depression. The depression is a deformation of the particular specimen and not a true anatomical feature. In its overall aspect, the femur of Sauro- suchus is of the crocodile type rather than like that of the dinosaurs. The proximal portion is a flange with a wedge-shaped articular head. The shaft is gently sigmoid and oval-shaped in cross section. Distally the termination flares out to what must have been large articular condyles. Owing to the defoi-mation of PVL 2267, it is not possible to determine the degree of rotation of the two extremes, but it appears to have been slightly greater than that of crocodiles. The proximal articulation consists of a rugose tear-drop-shaped surface, the broad portion of which forms a continuation of the thick anterior border of the femur. Be- hind this section the bone thins rapidly to the posterior edge. In PVL 2557 the broad portion is 5.5 cm thick, the tapered poste- rior edge is 2 cm. Curvature of the head in toward the acetabulum takes the form of an arc along the anterior border and reaches a maximum of 4 cm of inward displacement from the shaft. There is no fonnation of structures that can be defined as greater or lesser trochanters. The an- terior border of the femur is uniformly thick below the articular head, but in the upper portion it thins rapidly, forming a slight depression on the posterior flange area. Ventrally the bone is smooth, de- creasing in thickness from the expansion Saurosuchus and the Rauisuchid Thecodonts • Sill 341 of these; there is no intertrochanteric fossa. The posterior edge of the fhmge area thick- ens rapidly, becoming part of the shaft at the level of the fourth troclianter. Midway between the articular head and the fourth trochanter a slight expansion is present on the posterior edge. The fourth trochanter is a relatively small rugose bulge arising in the center of the ventral surface, ap- proximately one third of the way down from the proximal articulation, very much like that of crocodiles. Below the fourth trochanter, at approximately half the total length, the proximal flattened expansion disappears into the oval-shaped shaft. Distally the shaft expands evenly into the distal condyles. These are not pre- served, but a remnant of the intercondylar fossa on the dorsal surface indicates that the posterior condyle was the larger of the two. Tibia. This bone is known from the com- plete but poorly-preserved specimens PVL 2472, PVL 2267, and the well-preserved distal half of PVL 2557. The tibia is a ro- bust bone approximately twenty percent shorter than the femur. Proximally, the head expands to a triangular shape, the narrow point of which projects anteriorly and medially to form the cnemial crest. This crest extends down one third of the length before merging with the shaft. Posteriorly, the proximal surface is sepa- rated by a prominent depression into the condyles for articulation with the femur. This area of PVL 2472 is shattered, but from the area surrounding the depression it would appeal- that the two condyles were of nearly the same size. The medial surface of the proximal expansion formed the shortest leg of the triangle and bears a slight depression, probably indicating the contact for the fibula. Anterolaterally a broad flat area was present, separating the cnemial crest from the lateral condyle. The shaft is long, subround in cross section and slightly flattened on the anterolateral sur- face. Distally, the tibia flares out to a transverse expansion equal in size to the articulating surface for the femur. How- ever, it should be noted that in actual artic- ulation with the astragalus the tibia was rotated approximately thirty degrees, orient- ing the cnemial crest directly forward. Thus the lateral side of the distal termi- nation rested on the anterolateral portion of the astragalus and the medial portion on the posteromedial. The lateral expansion is broad and oval-shaped, the medial is nar- row and tapering. Separating the two areas of expansion is a narrow groovelike depres- sion on the posterior face that extends up the shaft to the midpoint (see Plate 2). The major surface of articulation is con- cave on the underside. All articulations are well ossified and have a shiny surface. Fibula. The fibula is known from the right distal half of specimen PVL 2557. The shaft is oval in cross section, the long axis oriented anteroposteriorly, and is flat on the medial surface facing the tibia. The most prominent feature of the shaft is the large tubercle on the anterolateral face, just above the midpoint. Presumably, this was for the insertion of the iliofibularis muscle. Above the tubercle the shaft curves slightly outward; below, it is characteristically con- cave toward the tibia. Distally the tibia has a flared surface for articulation with the calcaneum and astragalus. The articu- lating surface is lower on the lateral side than on the medial, and bears two grooves corresponding to the two tarsal elements. Aiticulation with the calcaneum occurs on the large lateral groove behind an anterolateral expansion of the bone. The astragalar articulation occupied a smaller diagonal groove on the anteromedial side of the distal termination. Tarsus. The tarsus of Saurosuchus was of the "crocodilian" type, in which the cal- caneum was functionally a part of the foot and the astragalus rotated with the crus. Four elements were present: proximally the large ti-iangular astragalus and the equally large tuberous calcaneum, distally a large lateral tarsal and a much smaller medial one. Elements preserved are: left and right 342 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 •*wr" Plate 2. A. Ilium and Ischium of Saurosuctius, PVL 2198. X Vs. B. Dorsal view of the paired Ischia. X Va. C. Distal portions of the tibia and fibula of Saurosuctius specimen, PVL 2557. X Va. Saurosuchus and the RAmsucHiD Thecodonts • Sill 343 Plate 3. A. Exploded and stereo view of the foot and tarsus, PVL 2557. B. Proximal view of the articulated metatarsals. C. Articulated foot. 344 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 D Plate 4. Stereo views of Tarsal elements of: A. Saurosuchus, B. Crocodylus, C. Undescribed rauisuchid from Los Colorados Fm. (courtesy of Jose Bonaparte), D. Neoaetosauroides, E. Riojasuchus. (All to same scale.) Saurosuciius and the RAUisucino Thecodonts • Sill 345 r calcaneum of PVL 2262, left astragalus of PVL 2472, all poorly preserved hut easily identifiable, and the extremely well-pre- served complete tarsus and foot of PVL 2557. Adequate description of these com- plex irregular bones is difficult, and the reader is referred to the stereo-photographs (Plates 3 and 4). The tarsus of Sauro- siichus appears to be virtually identical to that of Ticinosiichus, as described by Krebs ( 1965), except for minor details. However, Saiirosuchus was a much larger animal and the tarsal elements are naturally much larger and more heavily constructed. All tarsal elements were well ossified. Astrafialus. The astragalus is an irregular triangular block of bone. On its dorso- medial surface it bears a large, triangular, saddle-shaped area for articulation with the tibia. On the lateral side and separated from this area by a small, steep, forward- inclined ridge, lies the much smaller facet for articulation with the fibula. This sur- face is much more steeply inclined than that of the tibial articulation, and lies at approximately seventy degrees to it. An- teriorly, the surface of the bone bears a deep excavation, common to most reptiles that have a crocodiloid tarsus, medial to which is the bulbous convexity for articu- lation with the first metatarsal. A notable feature of the astragalus is its shallow depth. Thus the anteriormost border of the tibial facet is practically on the same level as the first metatarsal articulation. Posterolaterally the face of the astragalus is inclined downward from the peak of the ridge separating the epipodial articulations to the rounded ball that articulates with the calcaneum. Just behind the ridge peak a deep groove is present, which opens up posteroventrally to a curved depression in front of the ball joint. This depression fits over the anteromedial rounded border of the calcaneum. When thus articulated the fibular facets of both astragalus and cal- caneum are brought together and a more or less double ball and socket joint is fonued. The posteromedial border of the astragalus forms a rather featureless thick rounded border. Calcaneum. Basically, the calcaneum is a rectangular block of bone that bears a posterior upturned tuber and a medial process that forms the rear border of an anteromedially directed! hemispherical socket. Four polished articular surfaces are present on the bone. The anteriormost border is formed by the rounded, slightly ginglymoid articular surface for the fibula. This area is clearly marked and by its terminal position indicates that the cal- caneum must have been strongly rotated through its transverse axis, elevating the anterior end and depressing the tuber por- tion. Medial to the fibular facet is the rounded convexity that faces anteromedi- ally at a forty-five degree angle from the face of the fibular surface, which articu- lated with the previously described con- cavity of the astragalus. Again, the area of movement is well marked by the polished surface. Immediately behind this area is the small excavation that forms the anterior expression of the hemispherical socket that constitutes the major articulation between the two proximal tarsal elements. This excavation is continued medially onto the anterior face of the medially projecting process mentioned above, the whole form- ing a well-developed spherical depression directed inward at an angle of approxi- mately forty-five degrees from the anterior face. The fourth articular surface is a small rounded depression ventral to the fibular facet. This was for the reception of the large fourth tarsal bone. On the dorsal sur- face of the calcaneum, behind the fibular facet and lateral to the socket, lies a raised molding of bone that did not function as an articular surface and is not marked by muscle or tendon scars. It appears to have been an artifact of ossification. Continuing dorsally, the large tuber calcaneum projects upward and rearward. The dor.sal and posterior siuface of the tuber is rugose, indicating ligament attachment. The lateral surface is a flat wall, slightly depressed in 346 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Table 5. Measurement of the pes of Saurosu- chus galilei ( in centimeters). Lateral tarsal 7.0 Transverse width of anterior face Lengtii of lateral articulation with fifth metatarsal 5.5 Maximum height 4.0 Medial tarsal Height Width Metatarsals PVL 2557 Length Minimum shaft width Width proximal end Height proximal end Width distal end Height distal end I II 3.3 2.2 III IV 13.7 17.2 17.7 16.5 12.0 2.5 2.6 2.2 2.2 — 4.0 3.5 3.9 3.9 8.0 6.3 7.6 7.2 7.0 — 3.5 4.5 3.8 4.0 4.0 4.7 4.6 3.8 3.5 — Phalanges PVL 2557 Ij I, 11^ II„ III^ III^ IVj Vj Length 5.6 8.5 5.8 4.0 5.8 3.4 4.3 3.7 Height pro.ximal 4.3 3.4 4.1 2.7 3.9 2.9 3.0 3.0 Height distal 2.8 1.2 2.7 2.2 2.5 2.0 2.1 2.2 the center portion. Ventrally the surface is also flat, but a small pitlike depression is present at the base of the tuber. Distal tarsals. Two distal tarsals were present in the foot of Saurosiichus, ap- parently corresponding to numbers III and IV of the primitive reptilian tarsus. The lateral one is the largest of the two and is tetrahedral in shape; the ventral surface is flat, the other three sides form a rounded pyramid dorsally. The dorsal surface is slightly divided into a concavity for re- ception of an expansion on the astragalus, and a convexity that fits into a shallow pit on the oalcaneum ventral to the fibular articulation. Laterally the fourth tarsal bears a large, saddle-shaped, convex, articu- lar surface for the fifth metatarsal. Antero- medially are two convex surfaces, separated by a prominent groove, for articulation of the third and fourth metatarsals. At the extreme medial tip, beside the convexity for the third metatarsal, lies a small concave facet for reception of the third tarsal bone (see Plate 3). This element is a small rounded bone wedged between the lateral side of the second metatarsal and the astragalus. Pes. The pes of Saurosiichus had five sturdily constructed digits in the usual reptilian fashion. Metatarsal V was widely separated from the others, hooked, and bore a broad medial expansion. The re- maining four metatarsals were directed straight out from the foot, with a prominent transverse arch in the "instep" region. It is perhaps notable that the expansion of the proximal articulation surfaces of the meta- carpals lies in the vertical rather than the horizontal plane (see Plate 3B). Virtually all of the information available comes from the well-preserved right foot of PVL 2557, which is complete except for some of the distal phalanges. Additional elements of the foot are represented by poorly pre- served portions of left and right members of PVL 2267. Apparently the phalangeal formula was 2, 3, 4, 5, 3 in the usual primi- tive fashion. However, the fifth toe may have been reduced to but one or two phalanges. Metatarsal No. 1 is shorter than 2, 3, or 4, is thick bodied, and bears a pulley- shaped distal articulation behind which a prominent diagonal groove traversed the dorsal surface. Proximally, a concave facet is present on the medial side of the articulating surface, the remainder of the surface being smooth. The lateral margin of the proximal tip is vertical, its shape matching the medial border of the second metatarsal, with which it makes a very close fit. The first phalanx is relatively large, almost half the length of the metatarsal, and bears a proximal concavity with a ventrally projecting "heel" for articulation with the rolling surface of the metatarsal. Distally, the joint with the ungual is a shallow ginglymus, narrow at the tip and expanded ventrally. The ungual is a thick- bodied pointed claw, narrow at the top, wider on the bottom, and is half the length of the metatarsal. Largest of the metatarsals Saurosuchus and the Rauisuchid Thecodonts • Sill 347 is tlie second, although numbers 3 and 4 are ventral. The medial margin is expanded of similar length. It bears a large narrow at the top to form a bulge, with the afore- proximal articulation, expanded almost ex- mentioned groove lying just below it. Later- clusi\'i'l\' in the \ertical plane. On the ally the proximal articulation bears a con- medial side of the expansion are two facets cavity on the dorsal portion and a small for the first metatarsal. Laterally, the proxi- convexity ventrally, corresponding to op- mal articulation forms a straight vertical posite features on the close-fitting fourth surface with no overlapping contact for the metatarsal. Distally the articular surface third metatarsal. Midway down the side is of metatarsal number 3 is similar to that of a prominent pit, corresponding to a simi- number 2 but smaller. The rounded flange larly sized notch on the medial side of the is more expanded on the medial side than adjacent metatarsal. Presumably this formed on the lateral, and a groove is present be- a channel for nerve and blood supply. The hind the flange on the medial side. Only shaft is thickly built, similar to the con- the first two phalanges are preserved; they struction of the first metatarsal, and is are virtually identical to those of the second concave on the lateral margin but straight digit, but somewhat more slender. The on the medial side. Distally the articulation fomth metatarsal is slightly shorter and is a large rolling surface with a prominent more heavily constructed than the third, groove on the \'entral border. Just behind Its proximal articulation is diagonal in the the articular surface, on the lateral side, an vertical plane like that of the third, but on indentation is present between the flange the surface itself a prominent excavation is of the articulation and the body of the present below the side dorsal border for shaft. Shape and articulation of metatarsals the reception of the bulge of the fourth show that the axis of the transverse "instep" tarsal. A major feature of the fourth meta- arch ran between the second and third tarsal is its bowed shape; it is concave on metatarsals. Two phalanges of the second the lateral side, with the convex medial side metatarsal are preserved. As might be ex- fitting closely against the side of metatarsal pected, they are the largest and most number 3. This curvature also serves to hea\'ily constructed of the digits. The first rotate the plane of the promixal articulation bears a large concave flange proximally, a approximately twenty degrees from the short shaft, and a distal articulation similar vertical, toward the lateral side, from the to that of the metatarsal. However, the plane of the distal articulation. On the groove is considerably larger than that of lateral surface of the shaft in the proximal the metatarsal. The second phalanx is sub- region anterior to the articular surface is a rectangular in shape, and has a smooth prominent triangular depression, apparently conca\ity proximally and a pulley-shaped for muscles and flesh related to the lateral articulation distally. Although the ungual plantar pad of the foot. Distally, the is missing, the size and shape of the distal articular surface consists of a pulley-shaped articulation indicates that the claw was convexity somewhat different from that of approximately the same size as that of the the other metatarsals. The groove runs first ungual. Metatarsal number 3 is ap- diagonally across the articulation from preciably more slender than the others and ventrolateral to dorsomedial. \^entrally, is slightly longer than the second or the medial to the groove a prominent heel pro- fourth. Its proximal expansion is of similar jects downward. Laterally, just behind the size and shape to that of metatarsal number articular surface lies an expanded process 2, but whereas that of the second is a that continued onto the shaft, making a straight vertical surface, the third has a pronounced curvature of the lateral border diagonal proximal surface with the dorsal of the shaft, and giving thc^ distal articu- portion extended more posteriorly than the lation the aspect of being offset towards 348 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Figure 10. Two views of a posterior scute of Sauro- suchus. Left, dorsal; right, ventral. X V2. the medial side (see Plate 3). Only one phalanx of the fourth digit is present; it is rectangular, heavily constructed, and in general similar to that of the second digit, although .somewhat smaller and flatter. The proximal surface is more clearly divided into lateral and medial concavities than in the other digits. Distally the articular sur- face is considerably flatter, and lacks the downward extension of the rolling surface found on the first phalanges of the other digits. These are indications that the fourth toe was probably long and relatively slender. Specimen PVL 2267 has three iso- lated articulated phalanges that probably belonged to the fourth digit. These show a rectangular shape that rapidly diminishes in length distally with the last of the series, probably the pre-ungual, little more than a transverse rectangular chip of bone. How- ever, the association of these three phalanges (PVL 2267) is not certain. In Ticinosuchus all of the phalanges are longi- tudinally rectangular, as are all of the proximal ones preserved in PVL 2557. Metatarsal number five is a massive hook- shaped element that bears a large hemi- cylindrical articular surface on its medial side for the matching concavity of the fourth tarsal bone. On the anterodorsal face of the surface is a small facet for the lateral edge of the fourth metatarsal. Be- hind the large ball surface, the posterior border curves laterally and posteriorly to terminate in a rounded point at the rear lateral edge. From this point the lateral margin curves out and forward to the distal tip. A small expanded process is present on the lateral edge one third of the way back from the distal articulation. The medial surface of the "shaft" curves smoothly from the anterior tip of the major proximal articulation to terminate in the blunt surface of the distal articulation. This articular surface bears neither flanges nor grooves, but is a simple, slightly convex surface. The first phalanx is rectangular in shape, broader at the proximal end, and bears an expanded concave articular sur- face that partially envelopes the convexity of the metatarsal. Distally, the phalanx terminates in a simple flat vertical surface devoid of rounded features. No other phalanges are known for the fifth digit. The fifth toe was widely separated from the other digits. Dermal Armour Scutes have been found associated only with PVL 219(S. These were found partially articulated with the vertebral column, and like most of the vertebrae, are poorly pre- served. Three articulated scutes, much smaller than the others, were found in as- sociation with the other bones of the speci- men, but not in a definable position. As they are very well preserved, and in general the degree of preservation becomes better caudally in PVL 2198, it is assumed that these scutes were from the posterior dorsal region. Two paramedian rows of scutes were present on the dorsal region of Saiiro- siichus, the total width being 10 cm on specimen PVL 2198. As preserved, the two rows do not appear to have been joined by a strong sutural contact. The dorsal scutes are slightly asymmetrical and leaf-shaped in outline, drawn to a point in front and truncated at the rear. They are imbricated, the wide rear margin overlapping the point of the scute just caudal to it. Although the two rows join at the midline, the medial border is only slightly thicker than the Saurosuchus and the Raltisuchid Thecodonts • Sill 349 lateral. The anterior point is slightly asym- metrical; it is off center toward the medial border. Dorsally the scutes are gently arched in cross section, slightly more so on the lateral side than on the medial. A keel as such is not present, but there is a slight longitudinal ridge. Possibly a small inden- tation was present on the posterior border. Ventrally there appears to be but a slight indentation in the posterior portion to re- ceive the point of the following scute. A significant change in size takes place along the length of the series; the posterior scutes are smaller than the anterior ones, changing from approximately 5 cm in width to 4. The three isolated scutes differ consider- ably from the otliers, but are of the same pattern and certainly belong to the same specimen. They are, however, perfectly symmetrical, with each edge tapered to a very thin border (see Fig. 10). Anteriorly the point is longer and more tapered than in the other scutes, and fits into a wedge- shaped groove in the preceding scute. The dorsal surface is prominently ridged in the center, leading to the point anteriorly and to an indentation posteriorly. These char- acteristics suggest that these were members of a single row of scutes, rather than paired. A similar condition is reported for Ticino- suchus by Krebs (1965), and is to be expected given the other similarities of the two genera. The greatest difference be- tween the dorsal and the lumbar scutes is size; the former are 5 cm wide and ap- proximately 7 cm long while the latter are 3 cm wide and approximately 4 cm long. This condition differs from that of Ticino- suchus in which the scutes of the unpaired row are larger than the paired. However, the overall aspect of the armour of Sauro- suchus is that it is more reduced relative to the size of the animal than is that of Ticinosuchus. DISCUSSION Origin of the Rauisuchidae The anatomical characteristics of the figure 11. Pelvis of: A, S/7ans/st;c/,us (after Young); B, Ticinosuchus (after Krebs); C, Rauisuchus (from a known members of the family strongly photograph in Huene, 1942); D, Saurosuchus. B 350 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Figure 12. Left, calcaneum; Shansisuchus {after Young). right, astragalus of suggest direct derivation from the erythro- suchids, rather than from ornithosuchids or from a common erythrosuchid-ornitho- suchid ancestry. Cranial anatomy is but little modified from the erythrosuchid con- dition (see Fig. 13). Within the Erythro- suchidae, the most advanced member (both anatomically and stratigraphically) appears to be Shmisisiichus from the Ehrmaying Series of China (see Young, 1964; Reig, 1970; and Charig and Reig, 1970). This genus provides a rather good intermediate between the two families, and indeed was tentatively included in the "Prestosuchidae" by Romer ( 1972a ) . However, it still retains the primitive pelvic girdle and simple tarsal structure common to the Erythrosuchidae. As locomotory abilities seem to have been a principal evolutionary factor within the Rauisuchidae, it would seem appropriate to consider the less advanced Shansisuchus as an erythrosuchid. Major characteristics of the rauisuchids that can be traced with a reasonable degree of confidence through the lineage are: 1) Skull configuration: a keyhole-shaped orbit, large antorbital fenestra surrounded by a smooth depression, small supra- temporal fenestra, high narrow cranial table, and a posterior prong on the pre- maxilla. Some of the genera have an ac- cessory antorbital fenestra between the premaxilla and the maxilla. 2) Vertebrae: high neural arch, straight rectangular spine with distal expansion, deep interspinous clefts. 3) Pelvis: prominent posterior spine, presence of a brevis shelf, styliform ischium with an expanded tip, greatly reduced pubic plate, pubis with slight participation in the acetabulum. :-.-.:-.--'s>.-A. .«;_ ./ Figure 13. Comparison of cranial morphology in A, Stiansisuctius (after Young); B, Ticinosucfius (modified from Krebs); C, Luperosuchus (from Romer); D, Sauro- suctius. Not to same scale. 4) Femur: crocodilelike, without rounded medial expansion. 5) Tarsus: ball and socket crocodiloid type, fifth metatarsal hooked. The several genera that make up the Saurosuchus and the Rauisuchid Thecodonts • Sill 351 Rauisuchidae can be separated into three morphologic groupings that reflect both their stratigraphic position and their prob- able phylogeny: 1) an early group, repre- sented by Ticinosticlius from the earliest Middle Triassic; 2) an extensive intermedi- ate group represented by Luperosiichus, StagonosucJnis, "Mandasuchus", Prestosu- chus, and Raiiisuchus from the later Middle Triassic; and 3) Saurosuchus and the unde- scribed form from the Los Colorados, of earlier and later Late Triassic respectively. Ticinosuchus, the earliest member of the family, and the only one known from a com- plete skeleton, has a sk-ull that has been higlily fractured and compressed to a largely two-dimensional state. As recon- structed by Krebs, the skull is similar, but not strikingly so, to Saurosuchus and Lu- perosuchus. However, using the more com- plete knowledge afforded by the Argentine specimens, it is possible to reinterpret to some degree the skull of Ticinosuchus on the basis of the published photographs. Two modifications of Krebs' reconstruction ap- pear feasible: the antorbital fenestra was probably smaller than shown and was sur- rounded by a smooth shelf, and the anterior border of the maxilla was inflected just above the tooth row, possibly indicating a small accessory opening similar to that of Saurosuchus. Cervical vertebrae represent the only anatomical character that shows a consider- able degree of variation among the several genera of the family. In Ticinosuchus the cervicals are elongated, but otherwise un- specialized. A similar condition appears to be present in "Mandasuchus" but not in Stagonosuchus, Prestosuchus, or Raui- suchus. Only one cervical vertebrae is known from Saurosuchus; it is a highly specialized elongate structure so different from other known forms that it is assigned to the genus with reservation. In the other comparable features char- acteristic of the family, there is a remark- able similarity among the genera definitely assigned. More subtle differences dis- tinguish Saurosuchus as the most progres- sive of the described rauisuchids^; centra of the vertebrae are constricted, the ischium is longer and more rodlike, and the femur is more gracile than the corresponding features of the other genera. Within the Ischigualasto Basin three rauisuchids are found in the sequential continental sediments. The earliest of these is Luperosuchus from the Chanares For- mation (Romer, 1971a). It has already attained the large size characteristic of most of the family, but is known only from an incomplete skull. Changes in the skull from Luperosuchus to Saurosuchus to the Los Colorados form were slight; the orbit be- came more circular in the upper portion and the smooth shelf around the antorbital fenestra is larger in the later genera. It seems reasonable to assume that these three forms were continuous members of a single regional lineage. Very possibly Prestosuchus from Brazil should be included in the lineage. Prestosuchus is very comparable to Saurosuchus; apparently the only signifi- cant difference is that the femur of the former appears to be more heavily con- structed and less gracile than that of the latter. Relationship of the Ischigualasto Basin forms to other members of the family is not as close. The vertebrae of Stagono- suchus are somewhat constricted like those of Saurosuchus, but the pelvis is more primitive. "Mandasuchus" is quite similar to Saurosuchus, and the two may be con- generic or they may be closely related forms similar to Prestosuchus and Raui- suchus. Rauisuchus itself is less like the other members of the family and its associ- ation with the group has been questioned (Charig 1967, Romer, 1972a, Walker, per- sonal communication). Walker (personal communication) has suggested that Raui- suchus may be an ornithosuchid. His sug- gestion is based principally on some aspects ^ The undescribed rauisuchid from the Los Colorados Formation is larger than Saurosuchus; it had a considerably more advanced tarsus, but a very similar skull (Bonaparte, personal com- munication ) . 352 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 of the skull fragments and on the dermal armour. However, the premaxilla bears the posterior projecting prong that separates the external naris from the maxilla, and its overall shape is similar to that of Sauro- suchus and Luperosuchus. The ilium figured by Huene (1942, plate 27) is re- markably like that of Prestosuchus and Saurosuchus (see Fig. 7). Other elements are not as closely comparable, giving rise to the doubts about the affinities of the genus. However, the morphology of the vertebrae and dermal armour are not in- consistent with that of the other members assigned to the family, and their resem- blance to ornithosuchids may be superficial, as are a number of the resemblances be- tween the two groups (see discussion of vertebrae and tarsus). For the present, I would leave Rauisuchus in the family as- sociation that is termed "Prestosuchidae" by some authors, but recognize that it is less comparable to the larger genera Presto- suchus, Saurosuchus, and "Manclasuchus" than these are to each other. It would seem likely then that the Brazilian and Argentine genera were part of a South American radiation, perhaps from a Luperosuchus-liVe stock. The Afri- can forms, Manclasuchus and Stagono- suchus, may represent a separate but closely related line. A summary of the evolutionary history of the Rauisuchidae would then be: origin in the early Middle Triassic from a progres- sive group of erythrosuchids, the first members of the family probably near the Ticinosuchus level; adaptive radiation in the Ladinian and Carnian; survival of specialized members that could compete with dinosaurs in the uppermost Triassic, and extinction of the group by the Early Jurassic (see Fig. 14). Habits of the Rauisuchidae On the basis of the known remains, the rauisuchids can be described as large quadrupedal animals ranging in total length from three to six meters. The sharp serrated dentition leaves no doubt that they they were carnivores, and the deep narrow skull would suggest predaceous habits. During the Middle and Late Triassic they were probably among the largest of the terrestrial carnivores. Regarding locomo- tion, the hind limbs were of the crocodiloid grade of evolution, and as such the raui- suchids were reasonably good runners, although no doubt less agile that the later dinosaurs and probably less agile than the contemporary Ornithosuchidae.^ Rise of the rauisuchids may have been parallel to the rise of the rhynchosaurs and the gom- phodont cynodonts during the Ladinian and Carnian in a predator-prey relation- ship. It is usually assumed that the large thecodont predators disappeared during the Late Triassic owing to the competition from dinosaurs. However, the presence of a very large, advanced rauisuchid in sedi- ments considered to be Late Norian in age (see Bonaparte, 1972a and Sill, 1969 for details on the stratigraphic relationships of the Argentine Triassic), would indicate that these thecodonts had become adapted to prey on the early saurischians, many of which were herbivores. The last known rauisuchid was a very large animal and had an advanced digitigrade foot. Nevertheless, the femur remained at the crocodiloid stage of development, namely, without the for- mation of a medial condyle or a shift to the parasagittal plane of the body. Assuming that the vertical position of the limbs was an important adaptation, the rauisuchids would have been at a disadvantage with regard to the emerging carnivorous dino- saurs. Such a relationship presumably would explain the extinction of the group as the dinosaurs became dominant. Thecodont Taxonomy and Phylogeny Although thecodonts have long been recognized as the key group in the rise of ^ However, Bakker ( 1972, and in press ) has shown by experimental data that the physiologic cost of locomotion is dependent only on speed and body weight, entirely independently of limb posture. Saurosuchus A^a) the Rauisuchid Thecodonts • Sill 353 o 00 CO < en CD CL X) CD o g c o O c D C O C o CO 'c < Undescribed Saurosuchus Stagonosuchus Mandasuchus i Prestosuchus Rauisuchus / Luperosuchus Ticinosuchus \ Shansisuchus Figure 14. Suggested phylogeny of the Rauisuchidae. the archosaur faunas that dominated the later Mesozoic, they have been a poorly known and confusing group. As new dis- coveries have been made in the last few years there has been a renewed interest in the order, and at last the prospect emerges of unraveling the many and varied theco- dont lineages. Traditionally, thecodonts have been divided into three groups: 1) the very primitive forms from the Early Tri- assic, 2) the highly specialized taxa of the Late Triassic, phytosaurs and aetosaurs, and 3) the main stream, Pseudosuchia, somewhat of an "everything else" suborder. The n(>w discoveries ha\'e permitted the clarification of some relationships, and have added a new lineage, Proterochamp- sidae, to the order. But the major relation- ships are still far from settled, and there is a considerable number of genera that do 354 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 LATE TRIASSIC MIDDLE TRIASSIC EARLY TRIASSIC LATE PERMIAN Figure 15. Suggested phylogeny of some thecodont lineages. not fit into known families or even sub- by Romer (1972a) and by Bonaparte (1971), orders. as follows (the sequential order followed Current thinking on thecodont taxonomy by these authors has been changed to facili- is perhaps best reflected in recent papers tate comparison): Saurosuchus and the Rauisuchid Thecodonts • Sill 355 Ronier Order Thecodontia Suborder Proterosuchia Family Proterosuchidae Family Erythrosuchidae Family Prestosuchidae ( =Rauisuchidae) Family Proterochampsidae Suborder Pseudosuchia^ Family Ornithosuchidae Family Scleromochlidae Suborder Aetosauria Family Aetosauridae ( =Stagonolepidae) Suborder Parasuchia ( Phytosauria ) Family Phytosauridae Bonaparte Order Thecodontia Sul)order Proterosuchia Family Proterosuchidae Family Erythrosuchidae Suborder Pseudosuchia Infraorder Ornithosuchia Family Ornithosuchidae Family Rauisuchidae Family Pallisteridae Family Teleocrateridae ( ? ) Family Scleromochlidae Infraorder Sphenosuchia Family Sphenosuchidae Family Triassolestidae Infraorder Proterochampsia- Family Cerritosauridae Family Proterochampsidae Suborder Aetosauria Family Stagonolepidae (= Aetosauridae) Suborder Parasuchia Family Phytosauridae ^ The family Sphenosuchidae was placed by Romer in the suborder Protosuchia of the Crocodilia. Teleocrater and T liassolestes, together with other poorly known genera, are not assigned to families. "The suborder Archeosuchia was previously erected for the Proterochampsidae (Sill, 1967). Both of these authors retain tlie usual categories mentioned previously, but it is interesting to note the different inteipre- tations given to the newly defined line- ages Rauisuchidae and Proterochampsidae. Romer considers them to be continuations of the primitive radiation, while Bonaparte would suggest they are offshoots of the pseudosuchian stock. It is perhaps still premature to restructure thecodont taxonomy, but the new dis- coveries do make it possible for the first time to trace some of the lineages through- out the Triassic. Primitive thecodonts consist of three families; the ancestral stem Proterosuchidae (see Cruickshank, 1972), the large terrestrial Erythrosuchidae, derived from the Protero- suchidae, and the progressive Euparkeri- idae, usually considered the first of the Pseudosuchia (see Ewer, 1965 and Charig and Reig, 1970). The proterosuchids were probably aquatic or semi-aquatic carni- vores that somewhat resembled crocodiles. Erythrosuchids show many characters that relate them to the stem group, but were fairly large terrestrial carnivores. Euparkeria was apparently derived from an early line- age that separated from the Erythrosuchidae and evolved rapidly towards a more agile locomotory system. It has usually been assumed that it was the euparkeriid stock that produced the later thecodont radiation (Romer, 1966, and other textbooks). The new discoveries of fossil thecodonts, in particular those from South America, make it possible to connect some evolutionary lines of all three primiti\'e groups from the Early to the Late Triassic. As has been noted previously, the origin of the Rauisuchidae almost certainly lies in the Er\throsuchidae. Rauisuchids can be 356 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 traced through much of the Triassic with stones (Newton, 1894; Walker, 1964). To closely related forms present in every stage these Romer ( 1972b ) recently added of the period from the Anisian to the Gracilisuchus from the Chaiiares For- Norian (see Fig. 14). The Proterochamp- mation. These genera in turn show reason- sidae represents a newly defined lineage at ably close affinities to Euparkeria, and present known only from South America, appear to represent a descendant lineage Earliest members of the family are from from the euparkeriid type of early theco- the Chaiiares Formation, Chanaresuchus dont. and Giialosuchus, probably of Early In tracing these families from their Ladinian or Late Anisian age (Romer origins in the early history of the Theco- 1971b). Later forms occur in the Santa dontia, mention has been made only of Maria Formation of Brazil, Cerritosaunis those genera that are well enough known (Price, 1946; Bonaparte, 1971), and in the to show definite relationships; there are, of Ischigualasto Fonnation of Argentina, Pro- course, still many thecodonts whose system- terochampsa (Reig, 1959; Sill, 1967). Most atic associations are not clear at present members of the family show semiaquatic and who are usually assigned to families tendencies, the Brazilian form Cerritosau- on a rather uncritical basis. rus less so, and Froterochampsa itself more There remains the two well-known so. The apparently more aquatic habitus of specialized suborders, the Aetosauria and Proterochampsa was used by Bonaparte to the phytosaurs. In general these groups are separate the other genera from it as the limited to the Late Triassic, although an family Cerritosauridae, but the genera are isolated phytosaur has long been noted, no doubt closely related and probably and disputed, from the Early Triassic of should be placed in the same family. Plac- Europe (Jaekel, 1910; Gregory, 1962). ing the Proterochampsidae as an infraorder Phytosaurs are well known morphologi- of the Pseudosuchia implies a common cally, except for the tarsus, but no sure origin after the acquisition of the pseudo- indication exists regarding their relation- suchian adaptive characteristics. It appears ship to the primitive groups. In general it more likely that the Proterochampsidae has been assumed that they were pre- were independent derivatives of the primi- crocodile derivatives of the Pseudosuchia, tive stem proterosuchians, as suggested by driven into extinction by the appearance of Romer's classification, but they had ad- the true crocodiles (see Gregory, 1962). vanced beyond tlie level common to the Howe^'er, phytosaurs were basically primi- proterosuchids and erythrosuchids. On the tive animals, retaining additional skull basis of the skull, I previously (Sill, 1967) elements that were lost early in the de- believed them to be primitive crocodiles, velopment of the other thecodonts. Also, but the posteranial material of the earlier the pelvic girdle consisted of large platelike forms described by Romer renders this bones similar to the pattern of the primi- interpretation unlikely. tive groups (see Camp, 1930; Gregory, The third lineage to be well documented 1962, 1969). On the basis of the recently is not new at all, but is the "mainline" fam- described proterochampsids, it seems pos- ily Ornithosuchidae. As redefined by Bona- sible that phytosaurs may have been de- parte (1972a) this family would be re- rived from an earlier continuation of the stricted to the following well-defined aquatic forms of the Proterosuchia. genera: Venaticosuchus from the Ischi- Aetosaurs are the other closely-knit group gualasto Formation (Bonaparte, 1972b), of specialized thecodonts. Like phytosaurs Riojasiichiis from the Los Colorados For- they are known principally from the Upper mation (Bonaparte, 1969, 1972a), and Triassic, the earliest ones coming from the Ornithosuchtis itself from the Elgin Sand- Ischigualasto Formation of Argentina (Car- I Saurosuchus and the Rauisuchid Thecodonts • Sill 357 nian).' Those from Iscliigualasto are fully specialized members of the family, bearing littl(> indication of primiti\'eness. Aetosaurs were probably an early specialization for a rooting, pig-like habit (see Walker, 1961). Aside from their obvious specializations, aetosaurs retain many primitive character- istics common to the Erythrosuchidae and Euparkeriidae. As noted by Ewer (1965), Euparkeria was already more advanced in its locomotory apparatus than the aetosaurs. Therefore, the origins of the Aetosauria must have been from a progressive line of erythrosuchids or an early member of the Euparkeriidae. If it is true that the Euparkeria lineage represents an early de- parture from the Erythrosuchidae, based largely on limb specialization, then it would be more likely that the aetosaurs were an independent derivation from the erythro- suchid stem, perhaps from the same group that produced the rauisuchids. Indirect anatomical evidence supporting the affinity of Aetosauria with erythro- suchids is found in the tarsus. It has long been noted that the astragalus and cal- caneum of aetosaurs is of the "crocodile- type" in common with a number of other thecodonts. The closest comparison of these elements seems to be with the Raui- suchidae (see Plate 4). Another group of thecodonts, which has long been particularly difficult to interpret consists of those that share a number of characteristics of the crocodiles, but are not true crocodiles. These have been an enigma since they were first discovered around the turn of the century. They have been considered alternately as stages in the evolution of crocodiles (Huene, 1925), independent lineages (Haughton, 1924) and aberrant or primitive members of the Crocodylia (Sill, 1967; Romer, 1972a). ^ It is possible that an aetosaur was present in the earher Ehnnaying Series of China. A cal- caneum figured by Young (1964:81) is very much hke that of the Ischigualasto aetosaur, and quite unlike that of ornithosuchians. Walker ( 1970 ) has recently separated out the crocodilelike thecodonts and placed them as a suborder, Paracrocodylia, of equal rank with the Crocodylia in a new order Crocodylomorpha. Walker's work, based largely on re-examination of Spheno- siichus and HaUopus, indicates the presence of a possibly unified lineage that shared many anatomical characteristics of croco- diles, but were not ancestral to them. Whether or not a new order should be erected to place this group in juxtaposition with the Crocodylia will be decided by future discoveries. At the moment it does not seem to be justified. The Crocodylia are a well-defined group. Walker's Para- crocodylia is based on the Triassic family Pedeticosauridae (or Sphenosuchidae), the genus HaUopus — an apparent Jurassic de- rivative of the earlier family — and the Baurusuchidae, which he removes from the crocodilian suborder Sebecosuchia. Such a classification does not reflect the same degree of natural grouping that is found in the present category Crocodylia. It would seem more reasonable at present to consider the Pedeticosauridae as either a derivation of the thecodont line that gave rise to the true crocodiles, or as aberrant crocodiles from the early radiation of the Crocodylia. An alternative possibility is that croco- diles arose from an early branch of the Ornithosuchidae, possibly a derivative of the Euparkeria line, or from a continuation of the Erythrosuchidae, perhaps from the same stock that produced the Rauisuchidae (and possibly aetosaurs). Evidence sug- gesting the possibility of such a relationship is found in the similarity of the crocodilian tarsus to that of thecodonts in the above- mentioned categories. The so-called croco- dilian tarsal joint, in which the calcaneum bears a prominent tuber and is functionally part of the foot while the astragalus is fixed to the crus, appears to have been better developed in these lines than in either Proterosuchus or the Proterochampsidae. In addition, there appears to be a funda- 358 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 mental difference between the construction of the tarsus in ornithosuchids and the groups presumably derived from erythro- suchids. In the Ornithosuchidae the major joint between the proximal tarsal elements is formed by a ball on the anteromedial surface of the calcaneum and a correspond- ing socket on the astragalus. On the other hand, in rauisuchids, aetosaurs, and croco- diles, the main socket is on the calcaneum and the ball is on the astragalus. Both forms appear to be functionally the same, but possibly represent parallel evolutionary paths. Recognition of this condition, first noticed by Bonaparte ( 1971 ) , tends to diminish the difficulty noted by Krebs (1963) and Reig (1970) of explaining ap- parently unrelated thecodonts that possess very similar complicated tarsal joints. The "true" crocodile tarsus then becomes an impressive argument against derivation of this group from the Ornithosuchidae- Euparkeriidae type of pseudosuchian, and would tend to suggest a closer affinity with the erythrosuchid lineage, and the pre- sumed derivatives of that line. Neverthe- less, not enough is known about the tarsal joint of the Proterosuchidae, Proterochamp- sia, or Phytosauridae, to exclude them from a common ancestry with the Crocodylia. Tarsal joints of various members of the Thecodontia are currently under study by a number of paleontologists, some of whom feel that the structure may represent a 4cey to both thecodont and dinosaur phylogeny. There remains a considerable number of thecodonts that are not members of any of the groups mentioned in this paper. Some of these are almost certainly cladogenetic derivatives of these groups ( see the generic list in Romer, 1966, 1972a). The various phylogenetic possibilities of these forms have been discussed recently by Reig (1970) and little more can be said until additional fossil material is available. In addition there are a number of "ghost thecodonts," forms that have been named and placed in the ordinal hierarchy, but have never been duly described.^ These forms, largely from critical Middle Triassic strata, should provide additional insights into the thecodont radiation. Dinosaur origins remain unclear. Both saurischian and ornithischian representa- tives are present and clearly recognizable in the Ischigualasto Formation of Argentina (Late Ladinian-Early Carnian); saurischi- ans occur in the earlier Santa Maria For- mation of Brazil. There is no solid evidence for linking saurischians with either ornitho- suchid or rauisuchid thecodonts. However, Charig ( 1967 ) suggested the possibility of prosauropods arising from the latter group (Prestosuchidae in his usage). Reig (1970) considered it more likely that saurischians had descended directly from an erythro- suchid lineage than from a Eiiparkeria type of thecodont. No clues at all exist regard- ing the origin of the ornithischian dinosaurs; the earliest representative {Pisanosaurus from the Ischigualasto Formation) is a fully developed member of the group. It seems to be an inescapable conclusion that dinosaurs separated from thecodonts earlier than has usually been assumed, and that most thecodonts were competitors of dinosaurs rather than their progenitors. Thecodont-Dinosaur Transition It is perhaps paradoxical that the more we learn about thecodont evolution the less we know about dinosaur origins. Theco- ^ Mandasuchiis and Teleocrater were described by Charig in his doctoral thesis of 1956 and the names then published in an abstract in 1957. The names were incorporated into the literature by Huene (1956) and Romer (1966), but no formal descriptions have ever been published. In a later paper Charig, Attridge, and Crompton ( 1965 ) referred to the genera, but added a footnote to the effect that they were nomina nuda. Charig ( 1967 ) mentions both genera, an additional one from the same area, Pallisteria, and also two fami- lies, Pallisteriidae and Teleocrateridae. As author of all three genera and both families, he cites Charig ( 1967 ) , a paper which has not yet been published. All of these names, except Pallisteria and its family, are listed in Romer (1966), but all appear to be without proper foundation. Saurosuciius anu the Rauisuciud Thecodonts • Sill 359 O LD 00 < q: < or o THECODONTS o (J) if) < a: q: < Ornithischians < CO 2 3 Aetosourids :^-.r^j Ornilhosuchids ond crocodile'like forms Rauisuchids and other lorge forms Phytosours Proterochompsids -4=- ■:-.v:-: DINOSAURS I Aquolic Carnivores 2 Small Herbivores 3 Large Herbivores 4 Lorge Cornivores t) Small Carnivores Figure 16. Time-habitat relationships of thecodonts and dinosaurs (see text). donts evidently were successful, wide- spread, and diversified during the major part of Triassic time. Yet dinosaurs, usually considered as more or less the end result of thecodont evolution, had their origins well into the Middle Triassic (see Fig. 16). Thecodonts and dinosaurs apparently lived side by side during at least the hist half of the Triassic. This situation naturally raises some questions about the selective forces involved and the nature of the competition that presumably existed between the two groups. The superiority of dinosaurs relative to thecodonts is usually ascribed to a shift from a semi-erect to a fullv erect body stance (Bakker, 1971; Charig, 1972). In this case the more agile dinosaur loco- motion supposedly would have driven the thecodonts into extinction (but see foot- note, p. 352). However, an early or tran- sitional stage of dino.saurian limb posture is not found in any of the known thecodonts, and in particular there is no evidence of the shift to the simple hinge t\'pe of foot characteristic of dinosaurs. Charig (1972) postulated an as yet unknown thecodont ancestor in which the calcaneum was re- duced and rotated with the cms rather than with the pes. Reig (1970), on the other hand, would have the dinosaurs origi- nate directly from a primitive thecodont of an ervthrosuchid level in the Earlv Triassic, and evolve essentially independently of the 360 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 major thecodont radiation of the Middle and early Late Triassic. However, if this were the case it would be expected that dinosaurs rather than thecodonts would have dominated the Middle Triassic. The earliest dinosaur remains currently known come from the Manda and Santa Maria Formations of approximately Anisian or Ladinian age (Charig, 1967; Colbert, 1970). These genera, "Nyasasaunis" (un- described) and Staurikosaiirus are con- temporaries of rauisuchid thecodonts, found in the same sediments ( "Mandasuchus" and Prestosuchus). Staurikosaurus was more- over a predator of approximately the same size as Prestosuchus. A similar situation obtains in the Ischigualasto Formation, where the carnivorous dinosaur Herrera- saiirus is found with the same size carnivo- rous thecodont Saiirosuchiis. The earliest ornithischian, Pisanosaurus, is found in the Ischigualasto Formation and, although poorly preserved, shows that the basic features of the group had been acquired by that time (Casamiquela, 1967). The first theropods occur at approximately the same time (Charig, 1967), apparently oc- cupying an ecologic role parallel to that of the ornithosuchid thecodonts. Nevertheless, the thecodonts were con- siderably more abundant and varied in the sediments of the Middle and lower Late Triassic. They apparently took over the carnivore niche previously occupied by the carnivorous cynodonts, but did not extend into the herbivore field ( with the exception of the aetosaurs ) . Dinosaurs produced both carnivores and herbivores early in their history. The origins of both categories are still virtually unknown. Actual data from the fossil record allow three well -supported concepts to be stated: 1) dinosaurs were in existence at least during the last half of the Triassic; 2) thecodonts were abundant and diverse dur- ing the Middle and first half of the Late Triassic, becoming less so during the latter part of the Late Triassic; 3) although dino- saurs existed earlier, their major expansion did not begin until the last half of the Late Triassic. The reasons for the difference in expansion phases between the two groups are not clearly understood, nor can the apparent ecologic overlap between the large carnivores be explained on the basis of current data. However, it seems an in- escapable conclusion that the more agile mechanical condition of the dinosaur limbs was a factor in their eventual replacement of the thecodonts. It is also possible that the dinosaurs were undergoing more ex- tensive physiologic changes, perhaps related to the changes in locomotion (see Bakker, 1972). LITERATURE CITED Bakker, R. T. 1971. Dinosaur physiology and the origin of mammals. Evolution, 25: 636- 658. . 1972. Locomotor energetics of lizards and mammals compared. The Physiologist, 15(3): 278. -. In press. Lizard locomotor energetics and the Reptile-Mammal transition. Bonaparte, J. F. 1969. Dos nuevas "faunas" de reptiles Triasicos de Argentina. Gondwana Stratigraphy, lUGS Sumposium, Buenos Aires, 1-15 October 1967, UNESCO, pp. 283-284. . 1971. Cerritosaiinis binsfeldi Price, tipo de una nueva familia de Tecodontes (Pseudo- suchia Proterochampsia ) . An. Acad. Brasil. Cienc. (1971), 43: 417-421. . 1972a. Los tetrapodos del sector superior de la Formacion Los Colorados, La Rioja, Argentina. Opera Lilloana, 22: 1-183. 1972b. Annotated list of the South American Triassic tetrapods. Proc. II Gond- wana Symposium, South Africa, 1970, Pre- toria, pp. 665-682. Camp, C. L. 1930. A study of the phytosaurs, with description of new material from North America. Mem. Univ. Calif., 10: 1-174. Casamiquela, R. M. 1967. Un nuevo dinosaurio ornitisquio Triasico {Pisanosaurus viertii: Ornithopoda) de la Formacion Ischigualasto, Argentina. Ameghiniana, 4(2): 47-64. Charig, A. J. 1957. New Triassic archosaurs from Tanganyika including Mandasuchus and Teleocrater. Abstr. Diss. Univ. Cambridge, 1955-56: 28-29. . 1967. Archosauria. The Fossil Record. Geological Soc. London, pp. 695-731. . 1972. The evolution of the archosaur pelvis and hindlimb: an explanation in func- tional terms. In Joysey, K. A. and T. S. Kemp Saurosuchus and the Rauisuchid Thecodonts • Sill 361 (Eds.), Studies in Vertebrate Evolution. Edinburgh: Oliver and Boyd, pp. 121-155. — , J. Attiuu(;e, and A. W. Crompton 1965. On the origin of the sauropods and the classification of the Saurischia. Proc. Linn. Soc. London, 176: 197-221. -, AND O. Reig. 1970. The classification of eines Pseudosuchiers aus der Trias des Monte San Giorgio (Kanton Tessin, Schweiz). Paleont. A., 37: 88-95. 1965. Ticinosuchtis ferox nov. gen. nov. the Proterosuchia. Bio. Jour. Lin. Soc, 2: 125-171. Colbert, E. H. 1970. A saurischian dinosaur from the Triassic of Brazil. Am. Mus. Novi- tates. No. 2405: 1-39. Cruickshank, a. R. I. 1972. The proterosuchian thecodonts. In Joysey, K. A. and T. S. Kemp (eds.), Studies in Vertebrate Evolution. Edinburgh: Oliver and Boyd, pp. 89-119. Edmund, A. G. 1957. On the special foramina in the jaws of many ornithischian dinosaurs. Cont. Roy. Ontario Mus. Zool. and Palaeo., No. 48: i-14. . 1960. Tooth replacement phenomena in the lower vertebrates. Roy. Ontario Mus., Life Sci. Div., 52: 1-64. Ewer, R. R. 1965. The anatomy of the thecodont reptile Etiparkeria capensis Broom. Phil. Trans. Roy. Soc. London, B, 248: 379-435. Gregory, J. T. 1962. The genera of phytosaurs. Amer. Jour. Sci., 260: 652-690. , AND F. Westphal. 1969. Remarks on the phytosaur genera of the European Trias. Jour. Paleont., 43: 1296-1298. Haughton, S. H. 1924. The fauna and stratig- raphy of the Stonuberg Series. Ann. So. Africa Mus., 12: 323-497. Hoffstetter, R. 1955. Thecodontia. In Traite de Paleontologie, 5: 665-694. HuENE, F. V. 1925. Die Bedeutung der Spheno- suchtis Gruppe fiir Ursprung der Krokodile. Z. Indukt. Abstamm.-Vererbungsl., 38: 307- 320. . 1936. The constitution of the Theco- dontia. Amer. Jour. Sci., Ser. 5, 32: 207-217. . 1938. Ein grosser Stagonolepide aus der jiingeren Trias Ostafrikas. Neues Jahrb. Min. Geol. Pal., Beilage-Bd., 80: 264-278. . 1942. Die fossilen Reptilien des Siida- merikanishen Gondwanalandes. Ergebnisse der Sauriergrabungen in Siidbrasilien, 1928/ 1929. Miinchen: C. H. Beck'she. 332 pp. -. 1956. Paliiontologie und Phylogenie der niederen Tetrapoden. Jena: G. Fischer. 716 pp. Hughes, B. 1963. The earliest archosaurian rep- tiles. S. Afr. Jour. Sci., 59: 221-241. Jaekel, O. 1910. Ueber einen neuen Belodonten aus dem Buntsandstein von Bemburg. Sit- zungsber. Ges. naturf. Freunde, Berlin, 1910: 197-229. Krebs, B. 1963. Ban und Funktion des Tarsus sp. Ein neuer Pseudosuchier aus der Trias des Monte San Giorgio. Schweiz. Palaont. Abh., 81: 1-140. Newton, E. T. 1894. Reptiles from the Elgin sandstone. Description of two new genera. Phil. Trans. Roy. Soc. London, B, 185: 573-607. Price, L. I. 1946. Sobre um novo pseudosuquio deo Triassico superior do Rio Grande do Sul. Bol. Ser. Geol. Min. Brasil, 120: 7-38. Reig, O. A. 1959. Primeros datos descriptivos sobre nuevos reptiles arcosaurios del Triasico de Ischigualasto (San Juan, Argentina). Rev. Asoc. Geol. Argentina, 13: 257-270. . 1961. Acerca de la posicion sistematica de la familia Rauisuchidae y del genero Saurosuchus (Reptilia, Thecodontia). Publ. Mus. Munic. Cien. Nat. Trad. Mar de la Plata, 1: 73-114. . 1970. The Proterosuchia and the early evolution of the archosaurs; an essay about the origin of a major taxon. Bull. Mus. Comp. Zool., 139: 229-292. RoMER, A. S. 1927. The pelvic musculature of ornithischian dinosaurs. Acta Zoologica, 8: 225-275. . 1956. Osteology of the Reptiles. Chicago: Univ. Chicago Press. 772 pp. . 1966. Vertebrate Paleontology. Chicago: LTniv. Chicago Press. 468 pp. . 1968. Notes and comments on vertebrate paleontology. Chicago: Univ. Chicago Press. 304 pp. . 1971a. The Chaiiares (Argentina) Tri- assic reptile fauna. VIII. A fragmentary skull of a large thecodont, Luperosiichus fractus. Breviora, Mus. Comp. Zool., No. 373: 1-8. . 1971b. The Chaiiares (Argentina) Tri- assic reptile fauna. IX. Two new long-snouted thecodonts, Chanaresuchus and Gualosuchus. Breviora, Mus. Comp. Zool., No. 379: 1-22. . 1972a. The Chaiiares (Argentina) Tri- assic reptile fauna. XVI. Thecodont classi- fication. Breviora, Mus. Comp. Zool. No. 395: 1-24. . 1972b. The Chaiiares (Argentina) Tri- assic reptile fauna. XIII. An early ornitho- suchid pseudosuchian, Gracilisuchus stipani- ciconini, gen. et sp. nov. Brexiora, Mus. Comp. Zool., No. 389: 1-24. SfLL, W. D. 1967. Frotcrochanipsa harrionucvoi and the earh' evolution of the Crocodilia. Bull. Mus. Comp. Zool., 135: 415-146. . 1969. The tetrapod-bearing continental 362 Bulletin Museum of Comparative Zoology, Vol. 146, No. 7 Triassic sediments of South America. Am. Jour. Sci, 267: 805-821 Walker, A. D. 1961. Triassic reptiles from the Elgin area: StagonoJepis, Dasijgnatluts and their allies. Phil. Trans. Roy. Soc. London, B, 244: 103-204 . 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carno- saurs. Phil Trans. Rov. Soc. London, B, 248: 53-134. . 1970. A revision of the Jurassic reptile HaUopus victor (Marsh) with remarks on the classification of crocodiles. Phil. Trans. Roy. Soc. London, B, 257: 323-372. Young, C. C. 1964. The pseudosuchians in China. Palaeont. Sinica, New Ser. C, 19: 106-205. us ISSN 0027-4100 Sulletln OF THE Museum of Comparative Zoology The Cranial Foramina of Protrogomorphous Rodents; An Anatomical and Phylogenetic Study JOHN H. WAHLERT HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS, U.S.A. VOLUME 146, NUMBER 8 18 DECEMBER 1974 PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952- BULLETIN 1863- Memoirs 1864-1938 JoHNSONiA, Department of Mollusks, 1941- OccASioNAL Papers on Mollusks, 1945- SPECIAL PUBLICATIONS. 1. Whittington, H. B., and E. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution o£ Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredini- dae (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echinoderms. 284 pp. 4. Eaton, R. J. E., 1974. A Flora of Concord. 250 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. Creighton, W. S., 1950. The Ants of North America. Reprint. Lyman, C. P., and A. R. Dav^^e (eds.), 1960. Symposium on Natural Mammalian Hibernation. Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15. Proceedings of the New England Zoological Club 1899-1948. (Complete sets only.) Publications of the Boston Society of Natural History. Price list and catalog of MCZ publications may be obtained from Publications Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massa- chusetts, 02138, U.S.A. © The President and Fellows of Harvard College 1974. THE CRANIAL FORAMINA OF PROTROGOMORPHOUS RODENTS; AN ANATOMICAL AND PHYLOGENETIC STUDY JOHN H. WAHLERr Dedicated to Katherine Alexander and James Carol and Daniel CONTENTS LIST OF FIGURES 363 ABSTRACT 363 INTRODUCTION 364 CRANIAL FORAMINA OF RODENTS WITH SPECIAL REFERENCE TO MARMOTA 366 Systems: Cranial nerves 367 Arteries . 368 Veins 369 Foramina of the rodent skull 369 PARAMYIDAE Param ys 374 Leptotomus 379 Reithroparannjs 380 Ischyrotumus 381 Fseudotomus 383 Manitsha 384 SCIURAVIDAE .-... 385 ISCHYROMYIDAE 388 CYLINDRODONTIDAE 393 PROSCIURIDAE 397 APLODONTOIDEA 400 CONCLUSIONS 405 REFERENCES 408 LIST OF FIGURES 1. Marmota monax 370 2. Paramys copei _ _ 375 3. Paramys delicattis 376 4. Auditory and pterygoid regions of Paramys copei 377 ^ American Museum of Natural History, Verte- brate Paleontology Department, Central Park West at 79th Street, New York, N. Y. 10024. 5. 6. 9. 10. 11a. lib. 12. 13. Reithroparamys delicatissimus 380 Auditory and pterygoid regions of Ischyrotomus oweni 382 Sciiiraviis nitidus .__. 385 Auditory region of Sciuravus nitidus 387 Ischyromys typus 389 Ardyitomys occidentalis 394 Prosciurus sp. 398 Prosciurus aff. saskatchewaensis 398 Allomys nitens 400 Mijlugauhis laevis 401 Abstract. The cranial foramina and the blood vessels and nerves passing through them are de- scribed in detail for the sciurid genus Marmota; this data serves as the basis for understanding structures seen in the fossils. The cranial foramina are described and compared in North American specimens of the protrogomorphous rodent families Paramyidae, Sciuravidae, Ischyromyidae, Cylindro- dontidae, Prosciuridae, Aplodontidae, and Myla- gaulidae. The least variable foramina are those that transmit nerves; the most variable, veins. Presence or absence, relative position, number, and relative size of foramina are useful characters in determining relationships. Within the Para- myidae differences indicate an early radiation of lineages. Paramyids and sciuravids ha\'e man\' primitive features in common, but differ in several details; of especial interest in these families are the pathways of the internal carotid artery and its branches. Peculiarities common to the foramina of ischyromyids and cylindrodontids suggest that the two groups can be made subfamilies of the family Ischyromyidae. The Prosciuridae are included like- wise with the Aplodontidae and Mylagaulidae in the Aplodontoidea. Bull. Mus. Comp. Zool., 146(8) : 363-410, December, 1974 363 364 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 INTRODUCTION The origin of the Rodentia and their successful radiation can be attributed to a unique design for gnawing and chewing. Perfection of the design has involved modi- fication of the jaw and skull for more efficient muscle configuration, and special- ization of the incisors and cheek teeth in response to the multitude of specific niches into which rodents have diversified. The masticatory system has been subjected to great selection pressure and has been modi- fied from the original design in ways that were limited by genetic potential and by the efficiency of certain modifications relative to others. These are the principal reasons for the parallelism so typical of rodent phylogeny. To date, the classification of rodents has been based primarily on the structure of the masticatory muscles, the infraorbital foramen, the lower jaw, and the cheek teeth. These characters are all part of the masticatory system, and, when traced through time, their observed modifications reveal a complex phylogeny. Gaps in the sequence, however, cannot always be filled. Whole families of rodents stand in uncertain relationships to proposed phylog- enies. This situation is not surprising; in a phase of rapid evolution a gap of a few million years is enough to permit a discrete group to appear full-blown in the fossil record. The ancestry of such a group is often unclear because of parallelism among the earlier lineages from which it could have descended. The cranial foramina, unlike the com- ponents of the masticatory apparatus, are not part of a single functional system. There is no reason to suppose that selection acts on them as a unit or that selection pressure from the external environment acts on them directly. In the main, foramina serve a passive function; they permit nerves and blood vessels to pass through the bones of the skull. It is reasonable to suppose that foramina may vary in position and soft- part content so long as tliey satisfy the re- quirements of the circulatory and nervous systems. Within these limits selection is unimportant, and changes fixed in a small population by random genetic processes will characterize a new lineage arising from it. Fusion and division of foramina are pos- sible examples. The position or the existence of foramina may be changed as they are impinged upon by other structures. Foramina in the orbit are modified to lead around the roots of high-crowned cheek teeth. In the temporal region they may be closed off by enlarged bullae, and some other combination of foramina then acquires their function. A foramen may be taken over by a dif- ferent functional system. The infraorbital foramen has been seized upon in the hystricomorphous and myomorphous ro- dents for transmission of a part of the medial masseter muscle. From the moment of seizure it ceased to behave solely as the foramen it was and came under the in- fluence of the selective forces acting upon the masticatory system. The tough con- nective tissue around a foramen may change to accommodate a new stmcture. In those sciurids which lack an infraorbital canal, a tough membrane shielding the transmitted nerves and vessels from the lateral division of the masseter takes its place. Hill (1935 and 1937), Guthrie (1963 and 1969), and Bugge (1970, 1971a, b, and c) have been the principal contributors to knowledge of cranial foramina and the cephalic nervous and vascular systems in living rodents. They describe differences that appear to have a systematic basis. But the very nature of their work, limited pri- marily to modern examples, precludes dis- cernment of the primitive and derived conditions for each aperture. The pattern of evolution can be seen with certainty only when the time-dimension of paleontology is added. Detailed consideration of the fossils indicates which features in a group are primitive and eliminates the need to » rely on a so-called, but not in fact, primi- tive living genus such as Aplodontia. This paper on the protrogomorphous North American rodents is the first half of my Ph.D. dissertation (Wahlert, 1972), which included the scimomorphs also. De- scription of the cranial foramina in the latter families will be presented elsewhere, and I hope to extend the work to myo- morphous and hystricomorphous forms. The Protrogomorpha as defined by Wood (1937 and 1955) contain the families Paramyidae and Sciuravidae, which are parts of the initial rodent radiation, and the derived families Cylindrodontidae, Ischyromyidae, Prosciuridae\ Mylagaulidae, Aplodontidae, and Protoptychidae. Protoptychus was found to be both hystricomorphous and hystricognathus; a separate paper deals with its cranial and dental morphology (Wahlert, 1973). This approach to the study of rodent evolution brings with it a special set of problems. The number of fossil skulls adequately preserved is very small in com- parison with the number of fomis known from teeth. The forms whose skulls can be examined include representatives of every family, but they may be from specialized side branches and not from the main lines of evolution. Most specimens are incom- plete. The task of assembling data may be compared with that of a man in the dark who attempts to describe an exquisite topiary arabesque with only the aid of an unreliable flashlight. The text is divided into sections, each dealing with a single taxon; for extinct lineages this is the family, and for surviving lineages, the superfamily. Sections are sub- divided according to the importance of the included material and the completeness of the specimens. Paramyid genera are considered sepa- ^I have followed Wilson (1949c) and assigned the prosciurids to a taxon of rank eqnal to the paramyid group. Wood places them in the Para- mvidae as a subfamily. Cranial Foramina • Wahlert 365 rately because many rodent lineages may have originated from within the family. Differences between genera may be critical in determining relationships to later forms, and it is important to recognize that in- formation about cranial foramina in the fossils is quite uneven. The genera within several families and even within a superfamily are enough alike that a single section describing each group is sufficient. The ischyromyids, cylindro- dontids, prosciurids, and aplodontoids are treated in this manner. Discussions at the end of each section compare features within the groups de- scribed and compare the most interesting features of the entire assemblage with those considered in preceding sections. The bearing of the evidence provided by cranial foramina on the phylogeny and relationships of North American protrogo- morphous rodents is discussed in the con- clusion. A list of the specimens examined is presented at the beginning of each section or subsection. Definitions of the strati- graphic names can be found in Wood (H. E. Wood et ah, 1941) and Keroher (Keroher et al, 1966; Keroher, 1970). Abbreviations are as follows: AMNH American Museum of Natural History CM Carnegie Museum of Natural History F:AM Flick Collections, American Mu- seum of Natural History FMNH Field Museum of Natural His- tory KU University of Kansas Museum of Natural History LACM Los Angeles County Museum (CIT) (California Institute of Tech- nology Collection) MCZ Museum of Comparative Zool- ogy, Harvard University USNM National Museum of Natural History 366 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 SDSM South Dakota School of Mines and Technology UCMP University of California Mu- seum of Paleontology UNSM University of Nebraska State Museum UOMNH University of Oregon Museum of Natural History YPM Peabody Museum of Natural History, Yale University A letter code, which follows each num- ber, indicates the completeness of the fossil specimens: s - whole skull o - orbit n - snout t - pterygoid region p - palate c - cranium A code such as npo indicates that the snout, palate, and orbit of the particular speci- men are preserved and provided informa- tion for this study; the pterygoid region and cranium are either gone or are dam- aged and the detail destroyed. Measurements of length were taken with a dial caliper calibrated to 1/20 mm. The diastemal length is a straight line measure- ment from the back of the incisor alveolus to the anteriormost edge of the alveolus of the first cheek tooth. The sizes of foramina smaller than 1.0 mm were estimated with a Dunlap spark-plug gauge. Most of the figures were drawn with the aid of proportional dividers; enlargements of detail and outlines of small specimens were traced with a camera lucida micro- scope. I have made no attempt to show crenulations in the sutures but have taken care to illustrate the relationship of sutures to foramina. I have omitted detail from the teeth because excellent figures of the dentitions of all species studied are avail- able in the literature. Solid lines indicate structures and sutures that I have seen in at least one specimen of the genus illus- trated. Dashed lines indicate details that are less certain but probable in view of similar features in closely related fonns. Dotted lines represent guesses. To some degree all the figures are restorations. I have attempted to eliminate distortions and to reconstruct all broken elements; the figures are not copies of the specimens. The key to abbreviations in the illustrations is given in the caption of Figure 1. Acknowledgements. I would like to thank Bryan Patterson for his assistance and criticism throughout the course of this study, and Albert Wood for his suggestion of the topic and his assistance in the initial stages of the work. I am indebted to the staff members of the institutions, listed above, for making such a wealth of material available, and to many of these same people for their kind hospitality when I toured museums. I appreciate greatly the comments of Craig C. Black, Mary R. Dawson, Robert J. Emry, and T. Mylan Stout, all of whom read parts of the manu- script, and of Parish A. Jenkins, Jr., who read the entire thesis and had many excel- lent suggestions for improvement of this manuscript. Barbara Lawrence and Charles Mack of the Mammal Department, Mu- seum of Comparative Zoology, provided me with skulls for sectioning and specimens for dissection. Travel was financed by the Departments of Geological Sciences and of Biology at Harvard University. Other expenses were generously sustained by Katherine H. Wahlert. Special thanks are due to Carol C. Jones for incisive criticism of my grammar and for advice on the figures, to Daniel C. Fisher and James M. Labaugh, HI, for help in preparing the final manuscript, and to Katherine H. Wahlert for devoted typing. CRANIAL FORAMINA OF RODENTS WITH SPECIAL REFERENCE TO MARMOTA Hill (1935) was the first to attempt a complete listing of the foramina in rodent skulls. From dissections and prepared skulls he described the position and contents of each foramen and stated how thev differ Cranial Foramina • Wahlcrt 367 among scxcral grncra, but he gave no ac- count of the circulatory and nervous systems themselves. Unless tliese systems are understood it is not possible to interpret and name sex'cral of the foramina. Tandler (1899. 1901, and 1902), Guthrie (1963 and 1969), and Bugge (1970, 1971a, b, and c) have examined the cephalic arterial circula- tion of various rodents, but, apart from Guthrie, these authors pay little attention to the foramina involved. As an introduction to what follows, I present an account of the foramina and the circulatory system and cranial nerves in Marmota monax (Fig. 1). I follow Hyman (1942) and Greene (1935) for terminology of the soft parts. Marmota has several ad- vantages for this purpose. Although fully sciuromorphous, it retains most of the cranial foramina met with in the earliest rodents, and the bones of the skull do not fuse in the adult. Systems Cranial Nerves The hypoglossal (XII) emerges from one or more hypoglossal foramina and runs anteromedially into the base of the tongue. The foramina are situated just anterior to the occipital condyle on the ventral side of the skull. The vagus (X), accessory (XI), and glossopharyngeal (IX) emerge from the jugular foramen. It is between the bulla and the basioccipital and is lenticular in shape. The facial (VII) emerges from the stylo- mastoid foramen deep between the mastoid process and the bulla. The main part of the nerve runs anteriorly and diversifies over the masseter muscle. The chorda tympani branch of the facial emerges from a tiny slot, the canal of Huguier, in the front surface of the bulla. It runs antero- medially to join the lingual branch of the trigeminal (V) nerve. The mandibular division (3rd) of the trigeminal nerve (V) emerges from the large foramen ovale in the pterygoid region. Initially it runs anterolaterally through a notch in the lateral pterygoid flange. A strut of bone may cross the notch to form a foramcMi which I am calling the foramen ovale accessorius. The strvit separates the mandibular nerve and its internal ptery- goid branch. The auriculotemporal branch diverges just outside the foramen ovale. The mandibular nerve continues around the outside of the external pterygoid muscle and splits into three parts. The inferior alveolar branch enters the mandibular canal of the lower jaw. The mylohyoid and lingual, which is joined by the chorda t>anpani, run around the muscle and turn medially into the soft tissue. Two portions of the mandibular division, the masseteric and buccinator nerves, run dorsolaterally from the foramen ovale through a canal in the alisphenoid bone; the canal shares its posterior opening with the alisphenoid canal, but runs through the bone above it. These nerves emerge on the side of the head. Both nerves may pass through one foramen or each through its own (masticatory and buccinator foram- ina); the two cases can occur together on opposite sides of a single skull. The bucci- nator nerve runs anteriorly, but a small branch turns back on leaving the foramen. The masseteric nerve also has two branches; the smaller runs dorsally to the temporal muscle. The main part remains against the alisphenoid region in a shallow vertical channel. When this branch reaches the front of the posterior root of the zygoma, it turns laterally, passes through the mandibular notch of the jaw, and descends to the masseter muscle. The maxillary division (2nd) of the trigeminal nerve enters the orbit through the sphenoidal fissure. On cutting away the lateral surface of the alisphenoid region, the alisphenoid canal is exposed. Two large branches of the maxillary enter the canal dorsally through two large foramina; the small zygomatic branch emerges in some cases from a separate small foramen be- i 368 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 tween them. The posterior foramen (or two foramina) seems comparable to the foramen rotundum in other mammals. The two branches of the maxillary division unite to form the infraorbital nerve. The vidian nerve could not be separated or distin- guished from these. The main trunk of the infraorbital nerve enters the infraorbital canal; inside the canal a twig, the anterior superior alveolar nerve, descends into the maxillary bone. The trunk continues out onto the side of the snout. A small medial branch, the sphenopalatine, comes off the infraorbital nerve where it enters the orbit; it re-enters the skull through the spheno- palatine foramen. As it crosses the orbital floor it gives off a descending palatine branch, which enters the dorsal palatine foramen, runs through the palatine canal, and emerges on the palate through the posterior palatine foramen. In company with the anterior portion of the maxillary division, the ophthalmic division ( 1st ) of the trigeminal, and the abducens (VI), trochlear (IV), and oculo- motor (III) nerves enter the alisphenoid canal. The foramen through which they pass is comparable to the orbital fissure in other mammals. The nasociliary branch of the ophthalmic re-enters the skull through the more anterior of the two ethmoid foramina. The frontal branch of the ophthalmic ascends the medial wall of the orbit and exits onto the top of the skull through the supraorbital notch. The optic nerve (II) enters the orbit through the large, oval optic foramen. It is situated entirely within the orbitosphenoid. Arteries The common carotid artery gives off three branches when it reaches the back of the larynx. The superior thyroid artery diverges medially; the stapedial and oc- cipital arteries branch off on the lateral side. The main trunk continues as the external carotid. The occipital artery turns posteriorly and crosses ventral to the sta- pedial. It runs through a channel between the condyle and the paroccipital process to the back of the head and neck. The stapedial artery in company with the vagus, accessory, and glossopharyngeal nerves passes through the jugular foramen, and it enters the stapedial foramen in the bulla. It exits from the middle ear and enters the cranial cavity via the stapedial artery canal in the periotic. A dorsal branch from it continues out the temporal foramen to the temporal muscle. The main portion runs anteriorly and exits via the spheno- frontal foramen into the orbit; this is the ophthalmic artery, which supplies the eye and eye muscles with blood. One branch, the ethmoidal artery, enters the postero- dorsal ethmoid foramen. Another, the superior ophthalmic artery, ascends the medial wall of the orbit with the frontal branch of the ophthalmic nerve and goes through the supraorbital notch onto the top of the head. The external carotid artery bends later- ally and gives rise to auricular, internal maxillary, and other branches which supply the lower jaw, jaw muscles, and ear region with blood. At the bend, the external maxillary artery diverges and nms an- teriorly between the masseter and digastric muscles. It gives off a lingual artery and a glandular branch. In this region a third branch proceeds dorsally, gives off a tiny meningeal twig to the foramen ovale, enters the alisphenoid canal, and passes as the internal maxillary into the orbit where it divides into three branches. The outermost branch of the internal maxillary artery, the posterior superior alveolar, runs anterolaterally to the cheek region. The middle branch, the infraorbital, gives off minute branches tliat enter the nutritive foramina. It passes through the infraorbital foramen where a miniscule orbital twig pierces the bone dorsally, emerges from the malar foramen, and goes into the tissue anterior to the eye; a ventral branch, the anterior superior alveolar, in company with the nerve of the same name, enters a foramen below. The main trunk Craxial Foramina • Wahlert 369 continues out onto the snout. The inner- most branch of the internal maxillar}' arter>' gives rise to the descending pahitine artery and continues on into the sphenopalatine foramen. The descending palatine artery enters the dorsal palatine foramen, runs through the palatine bone, and emerges from the posterior palatine foramen; it diversifies in the tissue of the palate and disappears again into the incisive foramen. Veins Three distinct ti'unks carry blood from xarious parts of the head. These are the anterior and posterior facial veins, which unite in the neck to form the external jugular vein, and the internal jugular vein. The posterior facial recei\'es blood from the temporal and orbital-palatine regions. The infraorbital vein begins on the snout and passes through the infraorbital canal where it picks up a small tvvig from the anterior alveolar foramen. In the orbit, as the internal maxillar\' \'ein, it collects tsvigs from the nutritive foramina and small branches from veins passing through the sphenopalatine foramen and palatine canal. The descending palatine vein ascends through the posterior maxillary notch and joins it at the back of the maxilla. There are t\vo ethmoid foramina, and the ethmoid \ein exits through the posterodorsal one. It joins the ophthalmic, which then unites with the internal maxillary just before it enters the sphenoidal fissure. The internal maxillary occupies most of the space within the alisphenoid canal, the internal maxillary artery filling only a small dorsal portion of the canal. The vein communicates through the transverse canal in the basisphenoid with the same vein on the opposite side. It empties into the pterygoid plexus. The superficial temporal vein gatliers the posterior deep temporal, transverse facial, masseteric, and auricular branches. It is joined by a large vein from the temporal foramen and condylar area. This broad \'essel also continues into the pterygoid plexus. The inferior alveolar vein enters the plexus from the mandibular foramen in the jaw. The plexus anastomoses dorsally with the internal maxillary vein and ven- tralh' with the submental vein. A meningeal branch enters it through a small foramen bet^veen the bulla and the basisphenoid bone; this aperture may be a remnant of the middle lacerate foramen. The pterygoid plexus changes from a .sack-like structure into a large vein that proceeds posteriorly and is called the posterior facial vein. The anterior facial vein begins on the snout. It gathers tributaries from the masseteric and submental regions. It passes back into the neck where it unites with the posterior facial vein to form the external jugular. The internal jugular vein is quite small. It collects a branch from the inferior petrosal sinus in the carotid canal, leaves the cranium through the jugular foramen in company with the ners^es and the sta- pedial artery, turns posteriorly with them, and passes into the neck. Foramina of the Rodent Skull I have followed, as far as possible, the temiinology used by Hill ( 1935 ) and ha\'e attempted to find names commonh- used in the literature for foramina he did not de- scribe. My main points of departure from Hill are in the temporal and pterygoid regions. I have retained the term postgle- noid foramen but haxe abandoned the names subsquamosal, postsquamosal, supra- squamosal, and squamosal in favor of the general temi temporal foramina. In the Rodentia the temporal foramina are quite variable and cannot be categorized. The new terms post-alar fissure, squamoso- mastoid foramen, and foramen o\'ale ac- cessorius are used for apertiu'es that are different from anything in Hill's list. The fossils demonstrate that Hill's distinction betvveen alisphenoid and sphenopterygoid canals is not universal in the order; only one canal, the alisphenoid, is present in the earliest rodents, and the sphenopterygoid 370 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 appears to be unique to geomyoids among the groups examined. The following topographic list of foram- ina and their contents is based mainly on the woodchuck (Marmota mormx) unless otherwise stated. I have indicated also those foramina present but not figured, because they are hidden by another struc- ture. Foramina lacking in Marmota are described from the rodents in which they occur. No rodent possesses all of the foram- ina listed. 1. The unpaired interpremaxillary fora- men does not occur in the woodchuck. When present it is situated just behind the incisors on the median premaxillary suture, and it transmits a branch of the palatine op eth suo uml chu fo -t- hiL dpi a. 1 cm 1 cm msc + bu -poTTi .occ hy cc etic Figure 1. Marmota monax (MCZ B9911). Cranial Foramina • Wahlert 371 artery. As Hill (1935:122) states, it is relatively large in Aplodontia. 2. The incisive foramina flank the midline of the diastema. The lateral margin of each is intersected posteriorly by the premaxil- lary-maxillaiy suture. Each transmits a duct from the nasal passage, a branch of the palatine artery, and a branch from the palatine vein. 3. The major posterior palatine foramen is usually .situated in the maxillary-palatine suture. It transmits the descending palatine arteries and nerves and a small vein. In many rodents a posterior pair is present in the palatine. 4. The posterior maxillary notch is situ- ated between the end of the maxilla and the pterygoid extension of the palatine; it transmits the descending palatine vein. In many forms tlie notch is enclosed as a foramen. 5. The infraorbital foramen opens on the side of the snout in the maxilla. It is the anterior opening of the infraorl)ital canal and transmits the infraorbital nerve, artery, and vein. Protrogomorphous rodents lack the canal. 6. The anterior alveolar foramen (not figured) occurs in die floor of the infra- orbital canal and transmits the anterior superior alveolar nerve plus a .small artery and vein. Key to figures: Foramina and related structures (numbers correspond to those in text): aa — anterior alveolar (6) asc — alisphenoid canal (21) bu — buccinator (24) bup — posterior aperture, buccinator nerve canal (24p) cc — carotid canal (30) oca — anterior end, carotid canal (30a) chu — canal of Huguier (40) dpi — dorsal palatine (16) euc — Eustacian canal (29) eth — ethmoid (12) fac — facial canal (43) fco — fenestra cochleae (41) fo — foramen ovale (26) foa — foramen ovale accessorius (27) fro — foramen rotundum (20) fv — fenestra vestibuli (42) hy — hypoglossal (33) ifo — infraorbital (5) in — incisive (2) iom — depression, origin of inferior oblique eye muscle ipm — interpremaxillary (1) ito — interorbital (13) ju — jugular (32) ma — malar (7) mif — middle lacerate (28) mn — meningeal ms — mastoid (38) msc — masticatory (23) msp — posterior aperture, masseteric nerve canal (23p) nl — nasolachrymal (8) nu — nutritive (17) of — orbital fissure (19) op — optic (14) paf — post-alar fissure (35) pgl — postglenoid (34) pom — posterior maxillary notch or foramen (4) ppl — posterior palatine (3) spf — sphenofrontal (15) spl — sphenopalatine (11) spn — sphenoidal fissure (18) spt — sphenopterygoid canal (22) sqm — squamoso-mastoid (39) St — stapedial (31) stc — stapedial artery canal (44) sty — stylomastoid (37) suo — supraorbital notch (10) t — temporal (36) trc — transverse canal (25) uml — unossified area in maxillary-lachrymal (9) suture Bones: ab — auditory bulla as — alisphenoid bo — basioccipital bs — basisphenoid f — frontal i — jugal 1 — lachrymal m — maxillary mst — mastoid n — nasal occ — occipital OS — orbitosphenoid P — parietal pet — petrosal pi — palatine pm — premaxillary ps — presphenoid Pt — pterygoid sq — squamosal stippled areas: cut through bone solid line: seen in specimen dashed line: probable position dotted li ine: hypothetical position 372 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 7. The malar foramen (not figured) is situated in the orbit where the orbital and zygomatic portions of the maxilla meet above the infraorbital foramen. It transmits the malar artery from the infraorbital canal to the tissue in front of the eye. It is rarely present; presumably the artery is usually external to the bone. 8. The nasolachrymal foramen is situated in the lachrymal bone and is bounded anteriorly by the zygomatic portion of the maxilla. It transmits the lachrymal duct. 9. An unossified area between the lachrymal bone and the orbital and zygo- matic portions of the maxilla is not a fora- men, but the area of origin of the inferior oblique eye muscle. It has occasionally been confused with the nasolachrvmal foramen. 10. The supraorbital notch is an indenta- tion in the supra-orbital flange of the frontal bone. It permits passage of the frontal branch of the ophthalmic nerve and the superior ophthalmic artery to the top of the head. No superior ophthalmic vein was found accompanying them; it may have been too small to see. The notch occurs in the Sciuridae, only. 11. The sphenopalatine foramen is situ- ated at the front end of the orbital process of the palatine bone above the junction of the second and third molars. The maxilla forms the rest of its margin. It transmits the sphenopalatine nerve, artery, and vein. The bones participating in the margin of the foramen differ among rodent groups. 12. The two ethmoid foramina are entirely within the orbital lamina of the frontal bone. The anterior one faces ventrally into a shallow channel and transmits the nasocil- iary branch of the ophthalmic nerve. The posterior is larger and more dorsal; it transmits the ethmoid artery and vein. A single ethmoid foramen, which carries the nerve, artery, and vein, is present in the orbitosphenoid-frontal suture in most ro- dents. 13. A single or multiple interorbital fora- men pierces the orbitosphenoid in many rodents; it is absent in Marmota but present in some other sciurids. In geomyids it transmits a sinusoid vein between the orbits (Hill 1935:124). 14. The optic foramen is large and is en- tirely within the orbitosphenoid. It trans- mits the optic nerve. 15. The sphenofrontal foramen is situated between the orbitosphenoid and alisphe- noid. It is not quite separate from the orbital fissure in some specimens. It trans- mits the ophthalmic artery. The foramen is absent in many groups. 16. The dorsal palatine foramen leads into the palatine canal, which runs from the orbit downward through the palatine bone and out the posterior palatine foramen. It transmits the descending palatine artery and nerve and a small vein. 17. Many nutritive foramina (not figured) occur in the orbital surface of the maxilla above the cheek tooth roots. They transmit minute branches of nerves and arteries, and are present in all the specimens examined. 18. The sphenoidal fissure has as its outer wall the alisphenoid bone. The nerves and vessels transmitted by the orbital fissure (no. 19), the foramen rotundum (no. 20), and the alisphenoid canal (no. 21) exit from it. 19. The orbital fissure (not figured) is bounded anteriorly by the orbitosphenoid, and posterolaterally by the alisphenoid. It transmits the oculomotor, trochlear, and abducens nerves, and the ophthalmic di- vision and part of the maxillary division of the trigeminal nerve. In most rodents the fissure is united with the foramen rotun- dum. 20. The foramen rotundum (not figured) is completely concealed within the ali- sphenoid canal. It pierces the inner wall formed by the alisphenoid and transmits the remainder of the maxillary nerve; the I Cranial Foramina • Wahlert 373 zygomatic branch may have a separate foramen. The foramen rotundum and orbital fissure are united in most rodents. 21. The alisphenoid canal passes length- wi.se through the alisphenoid bone. It trans- mits the internal maxillary artery and vein. 22. The sphenopterygoid canal is absent in Marmota. I have found it only in geomyoids; it leads from the pterygoid fossa to the sphenoidal fissure. It transmits the internal maxillary artery, and its walls are the area of origin of the internal pterygoid muscle. 23. The masticatory foramen is situated in the alisphenoid and is often confluent with the buccinator (no. 24). It transmits the masseteric branch of the maxillary nerve. 23p. The posterior aperture of the masse- teric nerve canal (not figured) can be seen just anterior to the foramen ovale in some rodents. 24. The buccinator foramen is antero- ventral to the masticatory or confluent with it. It transmits the buccinator division of the maxillary nerve. 24p. The posterior aperture of the bucci- nator nerve canal (not figured) can be seen in a specimen of Paramys; usually this canal and the masseteric share a common aperture, as in Marmota. 25. The single transverse canal (not figured) runs between the alisphenoid canals through the basisphenoid. It trans- mits a vein connecting the two internal maxillary veins. 26. The foramen ovale is situated postero- laterally in the pterygoid region. It transmits the mandibular branch of the trigeminal nerve and a minute meningeal artery. 27. I define as new the foramen ovale accessorius that is lateral to the foramen ovale and transmits the mandibular branch of the trigeminal nerve to the side of the head. It is present in forms having a sub- stantial lateral pterygoid flange that reaches the auditory region. 28. The middle lacerate foramen is be- tween the pterygoid region and the an- terior end of the tympanic bulla or periotic as the case may be. The foramen is absent in Marmota; a minute aperture in the region (not figured) transmits a meningeal vein.^ 29. The Eustachian canal emerges dorsal to the anteromedial portion of the tympanic bulla. It transmits the Eustachian tube. 30. The carotid canal begins at or in front of the anterior end of the jugular foramen and runs anteriorly between the basioccip- ital and the periotic and tympanic. In many rodents having a canal it transmits the internal carotid artery. In Marmota, however, it transmits a vein, the inferior petrosal sinus, which joins the internal jugular vein. 30a. In some fossil rodents there is a foramen leading into the cranium antero- medial to the periotic. It seems to be the anterior end of the carotid canal. 31. The stapedial foramen (not figured) is dorsolateral to the jugular foramen and shares a common aperture with it. It enters the middle ear probably in the fused suture between the tympanic and periotic, and transmits the stapedial artery. 32. The lenticular jugular foramen is be- tween the basioccipital and the postero- medial part of the bulla. It transmits the vagus, accessory, and glossopharyngeal nerves, the stapedial artery, and the in- ternal jugular vein. ^ The function of the foramen is uncertain; no description exists of its contents in any of the Recent forms I have examined. In muroids it transmits the portion of the stapedial artery which emerges from the anterior part of the middle ear ( C'.uthrie, 1963). In the dog the internal carotid artery passes through the foramen into the cranium (Gregory, 1910:430). 374 Bulletin Museum of Comparative Zoology, VoJ. 146, No. 8 33. The hypoglossal foramen in the basi- occipital is anterior to the condyle and may be subdivided into two or more parts. It transmits the hypoglossal nerve. 34. The postglenoid foramen pierces the squamosal bone ventral to the zygomatic root and posteromedial to the glenoid fossa. It is absent in many Marmota skulls. When present, it transmits a large vein that drains most of the cranial cavity. 35. The post-alar fissure is absent in Marmota. I introduce this tenn for an opening between the alisphenoid wing and the tympanic bulla; it probably serves a function similar to that of the postglenoid foramen. In some forms it separates a part of the squamosal from the tympanic. 36. The temporal foramen is absent in Marmota. When present it is within the squamosal bone or in the squamoso-parietal suture, usually posterodorsal to the root of the zygomatic arch. It serves the same function as the postglenoid foramen and can take over the entire function of that opening. In some forms there are two or more temporal foramina. 37. The stylomastoid foramen is between the external auditory meatus and the mas- toid process. It transmits the facial nerve, and is constant in all rodents that possess a bulla, 38. The mastoid foramen is on the occip- ital surface between the occipital bone and the medial portion of the mastoid bone. It transmits a small vessel which, according to Hill (1935:128), is a vein from the neck muscles to the transverse sinus. 39. The squamoso-mastoid foramen is ab- sent in Marmota. I introduce the term for the foramen, which is present in many rodents, on the occipital surface between the squamosal and the mastoid. It trans- mits a vein. 40. The canal of Huguier is a minute slit in the anterior surface of the bulla. It transmits the chorda tympani division of the facial nerve. The following, which are not, strictly speaking, cranial foramina, have been shown in figures of several early rodents. They are useful as points of reference, and the canals are, of course, associated with soft parts intimately related to cranial foramina. 41. The fenestra cochleae (rotundum) is a round, membrane-covered aperture lead- ing into the scala tympani of the cochlea. 42. The fenestra vestibuli (ovale) is an oval, membrane-covered aperture leading into the scala vestibuli of the cochlea. The footplate of the stapes rests on this mem- brane. 43. The facial canal is in the periotic dorsolateral to the promontorium and is the canal in which the facial nerve traverses the middle ear. 44. The stapedial artery canal is also situ- ated in the periotic dorsolateral to the promontorium, and is die canal by which the stapedial artery exits from the middle ear. In many of the fossils it appears to be united, in part, with the facial canal. PARAMYIDAE Par amy s Specimens examined: Paramys copei (Figs. 2 and 4): Lysite Member, Wind River Formation: PU 16564 p. Lost Cabin Member, Wind River Fonnation: AMNH 4755 (type) npot, 4756 potc. P. delicatior: Twin Buttes Member equivalent, Bridger Fonnation: AMNH 55675 po. P. delicatus (Fig. 3): Blacks Fork Member, Bridger Formation: AMNH 12506 s, 13090 s; USNM 23556 s; YPM 13381 npo. P. sp.: Willwood Formation: PU 17421 np. Cranial Foramina • Wahlert 375 L elh i ii> .V hy ju fi' f ^ ^ bu spf ifo I cm Figure 2. Paramys copei (composite of AMNH 4755 and 4756). See Fig. 1 for key to foramina. Foramina The ratio of length of the incisive foram- ina to diastemal length ranges from .42 to .45. The lateral margins of the foramina are intersected behind the middle by the premaxillary-maxillary suture, which runs posterolaterally away from them. The posterior palatine foramina are wholly within the palatine bones. The larger anterior pair is close behind the maxillary-palatine suture and medial to the posterior halves of the first molars. The smaller posterior pair is more laterally situated than the anterior and is medial to the posterior halves of the second molars. The maxilla ends behind the cheek teeth in a blunt point. There is a slight posterior maxillary notch between it and the ptery- goid extension of the palatine. In front view the infraorbital foramen is elliptical; the major axis is inclined so that the top of the foramen is farther lateral than the bottom. The axis in P. copei measures 3.0 mm; in P. delicatus, 4.0 mm. In lateral view the foramen is approxi- mately vertical. The anterior alveolar fora- men, which is in the curve formed by the orbital wall and floor, is just posterior to the infraorbital, and is directed antero- medially. In front of the infraorbital there is a small foramen, probably for the nasal branches of the infraorbital arter)' and nerve. This foramen is more pronounced and more ventral in P. delicatus dian in P. copei. The lachrymal region is preserved onh- in the type specimen of P. copei, AMNH 4755. The nasolachiymal foramen is dorsal 376 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 ^PJ op eih spl aa til 1 cm cc tnlf fo pp7 in Figure 3. Paramys delicatus (USNM 23556). See Fig. 1 for key to foramina. to and not far above the infraorbital, and it is below the lachrymal flange of the zy- goma. A channel for the lachrymal duct descends the face of the lachrymal bone to the foramen. Sutures around the foramen are not visible. The sphenopalatine foramen is dorsal to the junction of the second and third molars. The maxilla and orbital process of the palatine make up its borders; the frontal may reach it dorsally, but this is not clear. The orbitosphenoid is excluded from the margin. Wood (1962:15, fig. 3A) figures the sutures incorrectly and shows the fora- men surrounded by the maxilla. The ethmoid foramen is dorsal and posterior to the third molar. It is within the frontal bone and is overhung by a lip from it. In the type specimen of P. copei, the frontal- orbitosphenoid suture reaches it posteriorly; in P. delicatus, AMNH 12506, the suture does not. The optic foramen, which is within the orbitosphenoid, is nearly 1.0 mm in diameter. It is dorsal and considerably posterior to the third molar. In P. copei it is closer to the sphenoidal fissure than in P. delicatus. The dorsal palatine foramen, which is in the maxillary-palatine suture, is immedi- ately ventral to the sphenopalatine, and both are within a single depression. Three specimens, AMNH 4755, 12506, and 55675, show this condition clearly. The same occurs in Thisbemys corrugatus, AMNH 94008 (for which there is no locality data; this is the only detail known of the foram- ina of Thisbemys, so I include it here). Minute nutritive foramina are present, as in all rodents, in the floor of the orbit above the roots of the cheek teeth. Cranial Foramina • Wahlert 377 The sphenoidal fissure at its opening is separated from the cranial cavity by a wall of bone; it is situated well behind the cheek teeth. In P. delicaius a slight ridge sets off the dorsal portion as a distinct channel. The alisphenoid canal joins the sphenoidal fissure laterally. The prominent sphenofrontal foramen is in the orbitosphenoid-alisphenoid suture near its junction with the frontal. In P. ilclicatus, USNM 23556, a conspicuous channel leads gradually downward and forward from the foramen. Wood (1962: 15, fig. 3A) has labeled a puncture in the bone as the sphenofrontal foramen; actu- ally, it is indicated in his figure by a dark area 2 mm behind and 3 mm above the point he has labeled. The masticatory and buccinator foramina are separate, the distance between them ranging from 1.0 to 3.0 mm. They face anterodorsally and anteriorly, respectively, and are a minimum of 3 mm from the foramen ovale. A minute foramen occurs between them in P. delicaius but not in P. copei; this was possibly for a branch that spht off the masseteric nerve before it emerged from the masticatory foramen. The buccinator foramen is farther anterior with respect to the masticatory in P. deli- catus. Wood (1962:15, fig. 3) has inter- preted these foramina differently and, I believe, incorrectly. The pterygoid region of P. copei is bounded medially by a flange and laterally by a faint ridge that is sufficient to enclose a foramen ovale accessorius. Medial to the foramen there is an oval depression. Within it are four foramina (Fig. 4). The posterior one leads from the braincase and is clearly the foramen ovale. The medial foramen leads into two canals; one, anteriorly di- rected, is the alisphenoid; the other, medi- ally directed, is the transverse canal. The anterior and lateral foramina lead to the buccinator and masseteric nerve canals respectively. I am in agreement with Black (196Sa:291, fig. 18) as regards their inter- pretation. In P. delicaius the lateral ptery- cc fac she mlf fo asc*trc / C^l Figure 4. Auditory and pterygoid regions of Paramys copei (AMNH 4756). Labeled outline drawing (hamular process hypothetical), and shaded drawing of the same. See Fig. 1 for key to foramina. goid ridge is somewhat weaker. A single large foramen, the foramen ovale, is visible in ventral view; in one specimen, AMNH 12506, a small foramen, probably the trans- \'erse canal, opens in its anteromedial wall. The t\anpanic bones are absent in all specimens of Paramys, and the middle lac- erate foramen is completely exposed. In P. delicaius it seems to be a single, irregular opening. In P. copei a stmt of bone trans- forms the medial portion into a separate, oval-shaped foramen. Wood (1962:42) interprets this latter as the entire middle lacerate foramen and states that the larger lateral part is apparently the foramen ovale. Since the specimen has a readily identifi- 378 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 able foramen ovale in the usual position, Wood's designation cannot be correct. Black (1968a: 291, fig. 18) has identified the foramina as I do; the smaller medial one he identifies as a part of the carotid canal. The carotid canal begins at the anterior end of the jugular foramen, it was probably open ventrally, and presumably transmitted a branch of the internal carotid artery. The canal may emerge from between the basi- occipital and the petrous portion of the periotic and enter the cranium through the medial portion of the middle lacerate fora- men, but this cannot be determined with certainty. The jugular foramen, as in all rodents, is lenticular in shape and is situ- ated between the periotic and basioccip- ital. The hypoglossal foramen is single in P. copei and P. clelicatiis. The posterior of two hypoglossal foramina shown by Wood (1962:15, fig. 3E) is a break in the bone. The postglenoid foramen, which is be- hind the glenoid fossa below the zygomatic root, is in the squamosal bone. Its major axis is smaller in P. copei than in P. cleli- catus, about 1.5 and 2.5 to 3.5 mm respec- tively. Temporal foramina are concentrated in the squamoso-parietal suture. Antero- dorsal to the postglenoid is a single fora- men in the suture. In some specimens it is accompanied by a second opening either above in the parietal (AMNH 4756, right side; AMNH 12506, left side) or below in the squamosal (USNM 23556, left side). A smaller foramen, also situated in the suture, is about halfway between these foramina and the back of the parietal. A mastoid foramen is present in the mastoid-occipital suture. It is well above the level of the top of the foramen mag- num. The foramina within the auditory region are exposed because the tympanic was not attached and has been lost. The least distorted periotic is preserved in a specimen of P. copei, AMNH 4756. The major features of the auditoiy region are shown in Figure 4. Wood (1962:43, fig. 14A) and Black (1968a:291, fig. 18) figure the same portion of this specimen. The most complete description of a paramyid periotic is given for P. delicatus bv Wood (1962:15, fig. 3B and C; page 18)! A lateral shelf of the periotic begins at the middle lacerate foramen and continues posteriorly for two-thirds of the length of the petrous portion. Behind it the mastoid portion broadens, curves medially to the posterior end of the jugular foramen, and ascends the occipital surface; most of this region is exposed outside the middle ear. Lateral to the fossa for the stapedius muscle there is a protuberance of the mastoid that is not situated so far posteriorly as the mastoid process in later rodents. Medial to the lateral shelf a broad chan- nel, which narrows posteriorly, runs from the middle lacerate foramen to the fossa for the stapedius muscle. In the absence of a tympanic the stylomastoid foramen is simply a groove lateral to the fossa on the medial surface of the mastoid protuberance and not a foramen as indicated in Wood's figure. The anterior part of the channel is presumably the area of origin of the tensor tympani muscle. In the middle portion are two posteriorly facing foramina, which are just internal to the shelf. The anterior one appears to be the foramen through which the stapedial artery left the middle ear; the posterior one, the foramen through which the facial nerve entered the middle ear. The medial portion of the auditory region is occupied by the promontorium. A faint channel, which marks the course of the stapedial artery, i-uns from the fenestra vestibuli to the anterior end of the jugular foramen. This portion of the channel cor- responds in position to the indentation for the stapedial foramen in the bulla of Reithroparamys (Fig. 5). The fenestra cochleae is in the posterior surface of the promontorium. I do not see, as Wood did (1962:18), evidence for the position of the auditory bulla. He states that the ridge paralleling Cranial Foramina • Wahlert 379 the median margin of the petrosal and over- hanging the petrosal-basioceipital suture (in AMNH 12506) seems to have served for bracing tlie median wall of the bulla. But the particular specimen he described is distorted; the petrosal has been tipped and the basioccipital crushed so that this ridge, which originally abutted against the basi- occipital, now stands away from it. The ridge in its proper position could not have braced the bulla. Wood also states that the depression be- tween the mastoid region and the lateral shelf of the periotic "... seems to have h(>ld the meatal tube of the bulla" (p. 18). It is more likely, however, that the meatus was lower down, as in Reithroporamys (Fig. 5) and Sciuravus (Fig. 7) and that the de- pression contained the dorsal portion of the tympanic. Leptotomus Specimens examined: Leptotomus hridgeremis: Twin Buttes Member, Bridger Formation: AMNH 12507 t. L. costilJoi: Huerfano Formation: AMNH 55110 s, 55111 (type) s. L. parvus: Twin Buttes Member, Brid- ger Fonnation: AMNH 12519 (type) p, 93030 p. Foramina Although two of these specimens are complete skulls, they are so fractured and crushed that very little information can be gotten from them. The two partial palates of L. parvus show that the posterior palatine foramina are wholly within the palatine bont^s, close behind the maxillary-ioalatine suture. The large pair is medial to the anterior part of the second molars. These are connected, each by a canal through the bone, to their respective dorsal palatine foramen. The latter is situated in the maxillary-palatine suture immediately ventral to the spheno- palatine foramen and above the anterior- most part of the third molar. In lateral view the infraorbital foramen is vertical; its exact shape and disposition cannot be determined. The sphenoidal fissure at its opening is separated from the cranial cavity by a wall of bone. It is well behind the last molar. Details of the region are visible in the fragiiKMitary specimen of L. hrid<^crensis. A slight ridge sets off the dorsal portion as a distinct channel. The alisphenoid joins the fissure laterally. A foramen in the medial wall of the alisphenoid canal is probably the entrance to the transverse canal; it would be completely hidden in an unbroken specimen. The exposed channel through the bone to the sphenofrontal fora- men is large and runs anteroventrally to a position that was probably very close to the top of the sphenoidal fissure. The pterygoid region is partially pre- served in AMNH 55110. The foramen ovale is large, and the lateral pterygoid flange bridges it ventrally to form a foramen ovale accessorius. The carotid canal appears to be like that of Paramys; it was probably open ventrally with the lateral lip of the basioccipital shielding the artery. Whether it carried the artery, or just the inferior petrosal sinus, however, cannot be determined. The postglenoid foramen is in the squa- mosal under the root of the zygoma. Its major axis measures about 1.8 mm. The auditory region is poorly preserved, but important details can be seen in the type of L. costilloi. The channel for the stapedial artery crosses the promontorium laterally to the fenestra vestibuli as in Paramys. At a point about a third of the way along its course another channel about half as wide diverges anterolaterally. Within a short distance this channel sub- divides. The diverging branch runs antero- medially across the promontorium. This bifurcating channel, I believe, marks the course of the promontory artery. 380 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 ms 1 cm Figure 5. Reithroparamys delicatissimus (AMNH 12561) See Fig. 1 for key to foramina. Reithroparamys Specimens examined: Reithroparamys delicatissimus (Fig. 5): Blacks Fork Member, Bridger Forma- tion: AMNH 12561 (type) npc. Foramina The ratio of length of the incisive foram- ina to diastemal length is .48. The lateral margins of the foramina are intersected very near the back by the premaxillary- maxillary suture, which runs posterolater- ally away from them. A single pair of posterior palatine foram- ina is present within the palatine bones. It is situated far laterally, almost on the maxillary-palatine suture, and is medial to the anterior ends of the second molars. The maxilla ends behind the cheek teeth in a blunt point. There is a slight posterior maxillary notch between it and the ptery- goid extension of the palatine. In front view, the infraorbital foramen is elliptical. The major axis, which measures 3.5 mm, is inclined so that the top of the foramen is farther lateral than the bottom. In side view the foramen is approximately vertical. The anterior alveolar foramen, which is in the curve made by the orbital floor and wall, is just behind the infra- orbital foramen. The wall of the orbit and the alisphenoid bone are missing. Enough of the nasolach- rymal canal is present to show that the nasolachrymal foramen was dorsal and close to the infraorbital. The pterygoid region (Fig. 5) is mostly missing. Several details can be made out, however. The external pterygoid flange, homologous to the lateral ridge in Paramys, is substantial and seems to enclose a fora- men ovale accessorius. Medial to the flange, the back of the foramen ovale is preserved; a channel, which is most likely the entrance to the alisphenoid and transverse canals, leads anteromedially from the foramen ovale. The middle lacerate foramen is completely covered by the auditory bulla. Just anterior to the bulla and lateral to the styloid process are two elongate foramina. A channel extends posteriorly under the bulla from the medial one. This suggests that the foramen may have been an aperture for a branch of the internal carotid artery, possibly the promontorial. The Eustachian canal passes over it. The lateral foramen may have transmitted a meningeal vessel. The carotid canal appears to begin at the Cranial Foramina • Wahlcrt 381 anterior c>nd of tlie jugular foramen and does not have a distinct entrance. A shc>U of the periotic is exposed anterohiteral to the jugular foramen at the point where the bulla is indented. The stapedial foramen is in this indentation and between the tym- panic and periotic. The hypoglossal fora- men is double on both sides. The larger foramen opens ventrally and faces antero- latc^ralh'; its rim continues out toward the ius from .18 to .21. The lateral margins of the foramina are intersected near the back by the pre- maxillary-maxillary suture, which runs posterolaterally away from them. The pair of larger posterior palatine foramina is in die maxillary-palatine suture and is medial to the middle region of the first molars. The smaller posterior pair, entirely within the palatine, is in line with die larger pair and medial to the ant(>rior halves of the second molars. In I. oweni, USNM 17161, there are two minute foram- ina situated more laterally in the palatin(\ The maxilla ends behind the cheek teeth in a distinct point that is best seen in /. horribilis. There is a post(>rior maxillary notch between it and the pterygoid exten- sion of the palatine. In front view, the infraorbital foramen is elliptical, and the major axis is inclined so that the top of the foramen is farther lateral than the bottom. The axis in 7. Jiorribilis is 3.3 mm long; in 7. oweni, 4.4 mm; in 7. petersoni, 4.5 mm. In side view the foramen is approximately vertical. The lachrymal region in these specimens is either missing or damaged. Wood (1962: 189) reports that the nasolachrymal fora- men is between the lachrymal and maxil- lary bones in the medial wall of the orbit. Wood (1962:189-190) states, "The sphe- nopalatine foramen sometimes lies on the frontal -maxillary suture and sometimes in the maxilla as in Faramijs. It is a little farther to the rear, just behind M'^ instead of just in front of it." His interpretation of its position in Paramys is incorrect, as noted above, and his placement of it in Ischyro- tomus also seems erroneous. Wood ( 1962: 207, fig. 71), in his figure of USNM 17160, shows the foramen within the maxilla and dorsal to the back of the second molar. Its position cannot be determined in the other specimens. The ethmoid foramen is above and between the sphenopalatine and optic foramina. A slight lip of bone overhangs it. Sutures in this region are indeterminate. The optic foramen, which is about 1.0 mm in diameter, is dorsal and considerably 382 BiiUetin Museum of Comparative Zoology, Vol. 146, No. 8 asc If fo / cm Figure 6. Auditory and pterygoid regions of Ischyroto- mus oweni (USNM 17161). See Fig. 1 for key to foramina. posterior to the third molar, and it is near the sphenoidal fissure. The dorsal palatine foramen was not seen; the region in which it would occur is fractured in every specimen. The sphenoi- dal fissure at its opening is separated from the cranial cavity by a wall of bone. A low ridge sets off its dorsal portion as a distinct channel. Tlie alisphenoid canal unites with the fissure. The sphenofrontal foramen is dorsal to the sphenoidal fissure and postero- dorsal to the optic foramen; a conspicuous channel leads downward and forward from it. Sutures in tliis area are indistinct in all specimens. Available specimens are too ci-ushed in the alisphenoid region to reveal whether the masticatory and buccinator foramina were separate or united. The pterygoid region of Ischijrotomus (Fig. 6) is bounded medially by a flange and laterally by a ridge; it is not developed into a fossa and is occupied almost entirely by a depression in which there are two openings. Posterolaterally the foramen ovale opens from the cranium; the lateral ridge is interrupted alongside it, and the beginning of a foramen ovale accessorius is suggested by the hook-like termination of the ridge. Anterior and medial to the foramen ovale is the second opening; it is deep within the angle formed where the lateral ridge and internal pterygoid flange meet. The alisphenoid canal runs anteriorly from it, the transverse canal medially. The dorsal portion of the alisphenoid canal is slightly damaged, but one small foramen, which probably transmitted the buccinator nerve, is clearly visible in its wall. The middle lacerate foramen is distorted by crushing in all specimens. The carotid canal begins at the anterior end of the jugular foramen. The hypoglos- sal foramen is either single or double; when double, the apertures open into a single pit. The postglenoid foramen is wholly within the squamosal bone. The major axis measures 1.5 mm in 7. horribiUs and 2.7 mm in 7. oweni. Temporal foramina are in the vicinity of the squamoso-parietal suture. The largest foramen is dorsal to the post- glenoid and above the zygomatic root; in some specimens it is entirely within the squamosal; in others on the suture. There is a small foramen anterior to it and another posterior in the parietal. The occipital sur- face is damaged, and sutures near the mas- toid foramen cannot be distinguished. The major features of the auditory region (Fig. 6) are essentially as in Paramys, but there are differences in detail. The mastoid portion of the periotic has a descending process lateral and posterior to the fossa for the stapedius muscle. This mastoid process is essentially modern in aspect. The foramina in the petrosal for the stapedial artery and for the facial nerve bear the same relationship to each other as in Paramys. In venti^al view, however, they are hidden under a single shelf which runs anterolaterally from a point on a level with the front of the fenestra vestibuli to a point overhung by the lateral shelf of the peri- otic. There is a distinct channel for the stapedial artery which crosses the promon- torium. It is broadest at the fenestra vesti- buli and narrows somewhat near the anterior end of the jugular foramen. This portion corresponds in position to the in- Cranial Foramina • Walilcrt 383 dentation for tlic stapedial foramen in ihc seems to be separated from the cranial bulla of Reithropommys (Fig. 5) and cavity by a wall of bone. The sphenofrontal Sciuravus (Fig. 7). foramen is visible on the right side of this speciracui just dorsal to the sphenoidal fis- Pseudotomus sure. I do not see a channel leading from it. The masticatory and buccinator foramina Specimens examined: ^j.e separatt> and over 4.0 mm anterior to Pseudotomus hiam: PBlacks Fork Mem- ^}^^, foramen ovale. A broad channel leads ber, Bridger Formation: AMNII 5025 dorsally from the masticatoiy foramen. The (type) nptc. buccinator foramen opens anteriorly; it is Foramina directly under the middle of the mastica- tory and less than 1.0 mm away from it. A portion of the external margin of the 7^^ pterygoid portion of Pseiidotomm right incisive foramen is present. Its curva- j^ similar to that of Ischyrotomus. The ture suggests that the foramen was rela- foramen ovale has only a hint of a lateral tively short, as in Ischyrotomus. pterygoid ridge alongside it; there is no At the back of the maxilla, near the suggestion of a foramen ovale accessorius. middle of the palate, a slight channel leads j]^q anterior portion of the pterygoid de- posterodorsally into what was probably the pression leads into the alisphenoid and larger of tlie posterior palatine foramina. It transverse canals. A posterior projection was evidently medial to the middle of the fj-Q^i the anterior margin of the middle first molar and in the maxillaiy-palatine lacerate foramen indicates that the foramen suture. was partially differentiated into medial and The infraorbital foramen is broad and lateral portions. The posterior margin is elliptical. The major axis, which measures j^ot preserved. 3.5 mm, is inclined so that in front view the xhe bullae are missing. The left periotic top of the foramen is farther lateral than ^nd the anterior end of the right are gone, the bottom, and in side view the top is ^nd the portion of the basioccipital that slightly farther anterior than the bottom, nomially abuts the periotic is exposed. The The anterior alveolar foramen, which is in ventral surface of the basioccipital extends the curve made by the orbital floor and laterally as a flange that would have over- wall, is just behind the infraorbital. The lapped the anterior extremity of the peri- nasolachrymal foramen is dorsal to and not otic. Dorsal to the flange on the lateral far above the infraorbital and is below the surface of the basioccipital, tliere is a posterior protuberance of the lachrymal channel that turns up toward the cranium bone. A wide channel descends the surface j^^t behind the middle lacerate foramen; of the lachrymal and bends anteriorly into possibly this is the carotid canal, or the the foramen. The maxilla appears to form course of the inferior petrosal sinus. The the ventral margin of the foramen. area of the basioccipital contained within Both orbits are considerably damaged the curve of the channel is sculptured and and the fragments of bone displaced; was most likely the place where the periotic sphenopalatine and dorsal palatine foram- attached. The posterior part of the basi- ina cannot be seen. The anterior part of occipital is missing. the ethmoid foramen is preserved on the The postglenoid foramen is within the right side; a lip from the frontal overhangs squamosal bone. Temporal foramina are it. The optic foramen, about 1.0 mm in present in or near the s(j[uamoso-parietal diameter, seems to have been dorsal and suture, but their number and exact posi- considerably posterior to the last molar. tions are indeterminate. The mastoid fora- The sphenoidal fissure at its opening men is above the level of the top of the 384 Bulletin Museinn of Comparative Zoology, Vol. 146, No. S foramen magnum in the mastoid-occipital suture. Manitsha Specimens examined : Manitsha tanka: Chadron Formation^: AMNH 39081 (type) np. Foramina The ratio of length of the incisive foram- ina to diastemal length is .23. The lateral margins of the foramina are not intersected by the premaxillary-maxillary suture, and it crosses the diastema behind them. The maxilla ends behind the cheek teeth in a point appressed to the palatine; there is no posterior maxillary notch. The infraorbital foramen seems small relative to the skull size. In side view it is approximately vertical. A single hypoglos- sal foramen is preserved on the left side. Discussion of the Paramyidae The paramyid rodents fomi a unified group with respect to cranial foramina. There are some differences between genera and species, among which changes in the pterygoid region and various patterns of arterial channels in the auditory region are the most striking. The ratio of length of the incisive foram- ina to diastemal length is high in Paramys and Reithroparamys, .42 to .48, and low in Ischyrotomiis, Pseudotomus, and Manitsha, .18 to .23. The lateral margins of the foramina are intersected behind the mid- point, but not at the very back, by the premaxillary-maxillary suture. In Manitsha the suture crosses the diastema behind the foramina and does not run into their mar- gins. ^ The American Museum catalogue incorrectly gives the horizon and locality for this specimen as Lower Brule, North Point of Slim Buttes, and this misinformation has been perpetuated in the litera- ture. The correct data, suppHed by M. F. Skinner (personal communication), are as follows: Chadron Formation, west side of Reva Pass, Hard- ing County, South Dakota. The posterior palatine foramina are wholly within the palatine bone in Paranujs, Leptotomus, and Reithroparamys. In Ischyrotomiis they are in the maxillary- palatine suture. A posterior maxillary notch is present in all except Manitsha. In Paramys, Thisbemys, and Leptotomus the dorsal palatine foramen is associated with, but separate from, the sphenopalatine foramen; its position is uncertain in the other genera owing to crushing in the orbital region. The sphenofrontal foramen is present in Paramys, Leptotomus, Ischyro- tomtis, and Pseudotomus. Other skulls were too damaged for it to be found. The presence of this foramen indicates that the ophthalmic artery was a branch of either the stapedial or the internal carotid artery. Masticatory and buccinator foramina are separate from each other and not especially close to the foramen ovale. I do not expect that Ischyrotomiis will prove to be an ex- ception when adequate material is found. Foramina in the vicinity of the foramen ovale differ among genera and even among species. The pattern found in Paramys copei could be that from which later ar- rangements were derived. In this species the foramen ovale, masseteric nerve canal, buccinator nerve canal, and alisphenoid and transverse canals open into a single depression. In Ischyrotomus and Pseudo- tomus entrances to the masseteric and buc- cinator nerve canals are hidden, and the depression is differentiated into two parts, one for the foramen ovale and another for the alisphenoid and transverse canals. The transverse canal was hidden in Leptotomus and a foramen ovale accessorius may have been present. The only available specimen of Reithroparamys appears to have been similar to Ischyrotomus. In Paramys deli- catus the alisphenoid canal is hidden, and the foramen ovale is the only conspicuous opening in the region; the transverse canal is small but visible in one specimen and hidden in the other. The middle lacerate foramen, when present and undistorted, appears to be Cranial Foramina • WaJdert 385 ^ msc ba spf op ^i^ 7 cm T^-sc ^u spn OiSC Figure 7. Sciuravus nitidus (reconstructed from USNM 17683, 18100, and 22477). See Fig. 1 for key to foramina. divided into hvo parts, the small medial one possibly for passage of the internal carotid artery. In Reithroparamijs, only this small aperture is visible; the t^'mpanic bulla covers the middle lacerate foramen if it is present. In the auditory region of Paramys and Ischyrotoinus there is a channel for the stapedial artery which crosses the promon- torium, and a stapedial foramen is present in Reithroparamys. In Paramys and Ischyrotoinus there is no channel indicating the presence of the promontorial arter\', whereas in Leptotomus this channel is clearly marked. The hypoglossal foramen is single in Paramys, single or double in Ischyrotomus, and double in Reithro- paramys. A rudimentary post-alar fissure is present in Reithroparamys and absent in the other genera. SCIURAVIDAE Specimens examined : Sciuravus nitidus (Fig. 7): Blacks Fork Member, Bridger Formation: AM Nil 12531 n, 12551 nptc, 13101 npoc; USNM 17683 c, 17697 np, 17700 np, 18023 np, 18100 s, 22477 s; CM 9683 np; YPM 13458 p. Foramina Accurate measurement of the incisive foramina is possible in three specimens. The ratios of their lengths to diastcmal 386 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 lengths are .41, .45, and .47. The lateral margins of the foramina are intersected near the back by the premaxillary-maxillary suture, which runs posterolaterally away from them. The posterior palatine foramina are within the palatine bones. The large an- terior pair is close behind the maxillary- palatine suture and medial to the junction of the first and second molars in some specimens, medial to the second molars in others. The smaller posterior pair is some- what more lateral in position than the anterior and is medial to the junction of the second and third molars. The maxilla ends behind the cheek teeth in a blunt point. There is a slight posterior maxillary notch between it and the pterygoid extension of the palatine. In front view, the infraorbital foramen is elliptical. The major axis is inclined so that the top of the foramen is farther lateral than the bottom. The axis ranges in five specimens from 1.7 to 2.6 mm. In side view the foramen is approximately vertical. The anterior alveolar foramen is just behind the infraorbital foramen in the curve made by the orbital wall and floor, and plunges anteromedially. The structvu-e of the lachrymal region, although not entirely preserved in any one specimen, can be determined for the most part. The nasolachrymal foramen is well above and slightly posterior to the infra- orbital. It is directly below the posterior protuberance of the lachrymal bone and may be surrounded by that bone, but sutures are not clear. A short channel leads into the nasolachrymal foramen and continues anteroventrally as a canal. The canal, exposed in AMNH 12531, passes in- ternal to the infraorbital foramen and turns medially a short distance in front of it. The sphenopalatine foramen is dorsal to the anterior half of the second molar. It seems to be bounded posteriorly by a long orbital process of the palatine, and on the other sides by the maxilla. The frontal may be barely excluded from its margin; the orbitosphenoid is completely excluded. The minute ethmoid foramen is in the frontal above and about halfway between the sphenopalatine and optic foramina. The orbitosphenoid does not seem to reach it. It is overhung by a slight lip from the frontal. The optic foramen is not preserved clearly in any specimen, but seems, at least in USNM 18100, to have been within the orbitosphenoid. It is posterodorsal to the third molar and near the sphenoidal fissure. The small dorsal palatine foramen is in the floor of the orbit posterior and slightly lateral to the sphenopalatine. The suture between the palatine and maxilla dips into it. The entire course of the canal descend- ing from it can be traced in two specimens, USNM 18100 and YPM 13458 (better seen in the latter). For a short distance the canal runs between maxilla and palatine; then it emerges and continues anteroven- trally on the internal surface of the palatine as a channel open into the choanae; finally it turns anteriorly through the posterior palatine foramen. The available specimens are too damaged to show whether the sphenoidal fissure at its opening is separated from the cranium. It is situated well behind the cheek teeth. The sphenofrontal foramen, seen in one specimen, USNM 18100, is in the orbito- sphenoid-alisphenoid suture just below the point at which the suture meets the frontal. A short channel leads anteroventrally from it. The masticatory and buccinator foramina are clearly preserved in only one specimen, USNM 18100. They are close together near the foramen ovale; channels from them lead upward and forward respectively. The pterygoid region is a relatively flat triangular surface bounded medially by a flange and laterally by a ridge; it is not developed into a fossa. There is an elon- gated depression medial and parallel to the posterior half of the lateral ridge. The foramen ovale opens from the cranial cavity into the posterior part of this depression. The alisphenoid canal runs forward from Cranial Foramina • Wahleit 387 the depression, and a lateral canal branches off from it to lead toward the masticatory and buccinator foramina. Medial to the alisphcnoid canal is the opening into the transverse canal. Details of this region are clear in two specimens, USNM 18100 and 22477. Dawson (1961:10, plate II) figured a distorted specimen and did not restore structures to their original positions; her sphenopalatine canal is my alisphcnoid canal. In specimens lacking the auditory bulla, the middle lacerate foramen is exposed. Its exact shape cannot be determined from the material a\'ailable, but the medial portion is partiall)' separated from tlie large lateral part. The medial portion is in the basi- sphenoid-basioccipital suture and pierces the side of the cranial floor. A channel leads posteriorly from the aperture across the anterior shelf of the periotic onto the promontorium. At the anterior end of the jugular foramen there is no space between the periotic and basioccipital for a carotid canal, and there is no separate entrance to a carotid canal elsewhere. The hypoglossal foramen is single. The postglenoid foramen is within the squamosal bone. Temporal foramina can be seen clearly in USNM 18100. The largest is in the squamoso-parietal suture about halfway between the back of the zygomatic root and the posterior margin of the skull, A second small foramen is immediately be- hind it in the suture, and there is a small foramen in the parietal anterodorsal to these. In most specimens the auditory bullae are missing; the periotic (Fig. 8) is clearly displayed in three specimens, USNM 18100 and 22477, and AMNH 13101. The major features are similar to those described for Paramys, but there are differences in relative proportions. The fenestra v(>stibuli, the fenestra cochleae, and the fossa for the stapedius muscle, indicated by a depres- sion, are as in Paramys. A channel leads anteriorly from the stylomastoid foramen to a single foramen that is slightly anterior y,^, sly f" {ac^slc /' .5 cm Figure 8. Auditory region of Sciuravus nitidus (re- stored from USNM 17683). See Fig. 1 for key to foramina. to the fenestra vestibuli; the facial nerve and the stapedial artery evidently shared this one opening. Channels showing the courses of blood vessels are present on the surface of the promontorium. The main channel for the internal cartoid artciy, pos- sibly the promontory branch, begins just anterior to the jugular foramen; midway across the promontorium it turns antero- dorsally and runs to the medial portion of the middle lacerate foramen. A channel for the stapedial artery curves postero- dorsally from the point where the internal carotid turns anteriorly, and it leads to the fenestra \'estibuli. The bulla is preserved on one specimen, 388 Bulletin Museum of Comparative Zoology, Vol. 146, No. 8 USNM 22477. It has been moved out of place, and no markings on the periotic indicate precisely where it was situated. I did not notice a stapedial foramen, but one must have been present in the margin of the bulla, as indicated in Figure 7, to per- mit passage of the internal carotid artery. It is not possible to see in the specimen whether the bulla completely covered the middle lacerate foramen; measurements suggest that it did not, and I have so shown it in the figure. Discussion of the Sciuravidae The ratio of length of the incisive foram- ina to diastemal length is high, as in Paramys and Reithropammys, and is much higher than tliose of Ischyrotomus and Manitsha. The posterior palatine foramina are within the palatine bone, again as in Paramys and Reithroparamys. A posterior maxillary notch is present, as in most paramyids. The orbital process of the palatine reaches the back of the sphenopalatine foramen, whereas the orbitosphenoid does not. This arrangement occurs in Paramys. The sphenopalatine and optic foramina are farther forward relative to the cheek teeth than those of paramyids. In Sciuravus the dorsal palatine foramen clearly is separated from the sphenopala- tine, a condition perhaps foreshadowed in paramyids. The sphenofrontal foramen is in the orbitosphenoid-alisphenoid suture, as in paramyids. Masticatory and buccinator foramina are present and considerably closer to the foramen ovale than they are in paramyids. The arrangement of foramina in the pterygoid region is similar only to that of Ischyrotomus and Pseudotomiis. Separation of the medial part of the middle lacerate foramen to receive a branch of the internal carotid artery occurs in paramyids. Sciuravus, however, does not have a carotid canal between the periotic and basioccipital. Instead, the internal carotid artery entered the middle ear and crossed the promontorium in a shallow channel before entering the cranial cavity. The carotid circulation in Leptotomus may be the same, though the carotid canal is also present. There is only a single aperture in the petrosal for the stapedial artery and the facial nerve to exit from the middle ear, whereas in paramyids a pair of openings is visible. The postglenoid and temporal foramina are of about equal size, as in paramyids. ISCHYROMYIDAE Specimens examined: I have seen and measured so many speci- mens (approximately 65) that a complete list would be excessive. The specimens re- corded below are only those that are nearly complete or are cited in the text. Ischyromys douglassi: Chadron Forma- tion equivalent: CM 1123 pot; 10966 potc. I. typus (Fig. 9): Orella Member, Brule Formation: AMNH 694 s; FMNH, P 12747 poc; MCZ 18979 potc; YPM 12521 s; PU 11383 s. Brule Formation: USNM 15929 npc; 15933 s. 7. sp: Chadron Formation equivalent: CM 24129 c. Orella Member, Brule Formation: AMNH 38865 s; CM 9463 npoc. Brule Formation: USNM 16953 s; 175352 tc; 175354 npo; CM 9755 pt. Titanotheriomys veterior: Chadron For- mation equivalent: CM 9809 p; 10660 npot. POrella Member, Brule Forma- tion: MCZ 17202 s. T. tcyomingensis: Chadron Formation equivalent: AMNH 14579 (type) npt. T. sp.: Chadron Formation equivalent: CM 8924 npt. Foramina The ratio of length of the incisive foram- ina to diastemal length, measured in seven- teen ischyromyids, ranges from .21 to .30 with a cluster of nine around .24 and a cluster of eight around .28. The three specimens of Titanotheriomys in which this region is preserved fall at the low end of the range. The lateral margins of the Cranial Foramina • Wahlcrt 389 spf do eth sly / fo*asc ■paf foa I cm cc rnn Jo Ire pom ppl in Figure 9. Ischyromys typus (AMNH 694). See Fig. 1 for l 0 X 01 n u •-t c ■H c e l4 N "0 -H W o lO g 0 o cr (N n 0 B (N c 4) «l n 0) 5.S O O a. 1 •H O u 1 £ -a >, o ■f 1 ■f 10 "0 rH O' C + 1 1 1 ^5 o •K vO V 4) ^ k4 rH 3 <0 n iH 1 5 r- in O 4J VI CO 0 ^ •rt iH S • « %ii'^ n Oi iJ ♦ 1 § o z M i o >! a H Q 2 z M W o O (*> Q U 2^ w feg O U 2 >< "i § s n s M S H X M b ^ w U 10 W M b j s < M Eh to u (/) H u H H M to u ^2 ri8 M z z 10 z Z M O Q >< J5 < i ,1 i S O o •i 2 (0 -1 o o M h M 0. •o 01 > u 0) •0 v> 01 X) I 0 01 0> 0) c J3 0 0 c II II + 1 516 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 for successive land surfaces, although in some areas paleosols appear to be present (G. D. Johnson, personal communication). A laterally continuous beach sand trans- gressed over the deltaic flats, but ap- parently nowhere formed preserved beach bars or ridges. Predominantly lacustrine conditions followed, with influxes of tuffa- ceous material from distributary mouths and spreads of shell debris over level sub- aqueous surfaces. The lack of extensive erosion on the mudflats with the coming of lacustrine conditions indicates the character of the transgression. Although the mudclasts and CaCOa nodules incorporated in the sand are no doubt derived from the mudflat (as are the fossil bones), the mudcracks on the surface have not been eroded away. This can only mean a very low gradient shore- line, low wave energy, and probably a relatively rapid transgression. Otherwise, it is difficult to explain why the increased energy level which carried the sand would not have formed beach ridges and eroded beach fronts, destroying the upper surface of the mudflats. The Cretaceous Wealden Lake environ- ment in the Anglo-Paris Basin provides some close analogues for the transgressive deposits of the Koobi Fora Fm., and par- ticularly those of 103-0256. P. Allen (1959) reports graded sheets of pebbly sand that were spread extensively over deltaic de- posits as the Wealden Lake rose. One of these, the "Top Ashdown Pebble Bed," is a graded unit with pebbly sands fining upward to sands and silts. It is only 10-20 cm thick, and truncates all underlying structures and sediments. The base is erosional, and the components of the bed are derived from underlying deposits (P, Allen, 1959:292). This is directly compa- rable in most characteristics to the 103-0256 transgressive sand, but differs in that 103-02.56 does not appear to be derived from the underlying beds, except for the mudclasts and carbonate nodules. The base is less erosional than in the Wealden trans- gressive sheets. The sand in 103-0256 was evidently redistributed from former beach and distributary mouth deposits and carried shoreward by the advancing lake. The fossil bones derived from the trans- gressive sand and the mudflats deposits are concentrated on the slope below a strike ridge created by the westward tilted, resistant sand. They are highly mineralized, although the pore spaces of many of the fossils are not filled with cement of any kind, a unique characteristic of this as- semblage. There is evidence for mixing of bones with varying degrees of predeposi- torial weathering. Some retain fresh, un- cracked and unflaked surfaces while others are weathered and have cracked or worn surfaces preserved rmder their sandstone matrix cover. The quartz equivalents for the fossils, estimated according to their densities when fresh (Table 4) range from 1.0->20 mm. This is a very different size range than that of the quartz sand which forms the matrix of the fossils (<. 1-1.0 mm). The distributary sands associated with tlie mudflats contain grains up to 5 mm in diameter, yet this size range is absent from the transgressive sand and apparently was not present on the deltaic flats. If the bones were derived from the distributaries, it is reasonable to expect them to be associated with sand larger than 1.0 mm. It is possible to con- clude that most of the bones were probably not brought into the area by fluvial pro- cesses, but were derived from a death assemblage that lay upon the deltaic flats. The presence of many fresh, unabraded bone surfaces further supports a locally de- rived fossil deposit. The bones were prob- ably redistributed by the transgression, but final burial was evidently rapid and abrasion minimal. Locality 130-0201: Delta Margin Vertebrate fossils were sampled from a relatively large stratigraphic thickness (7.0 m) of tilted and faulted sediments. A variety of lithologies occur, and overall East Rudolf Palkoecology • BcUnnsmcycr 517 <2;rain sizes range from <. 1-6.0 mm. The dominant litliologies are evenly stratified sandy silts, silt}^ elays and medium-grained Flagstone sands. The sands are generally elean and rieh in biotite. Coarser, more poorl\- sortcxl sediment oeeurs in laterally restrieted lenses. This locality lies within the marginal deltaic facies of the Lower Mb. of the Koobi Fora Fm. The units sampled for fossil vertebrates appear to be on an actively aggrading margin of the deltaic complex. The sedimentary characteristics listed in Table 5 agree in many respects with Butzer's (1971a:79) description of the modern Omo interdistributary basins (la- goonal mudflats and marsh), including the presence of limonitic mottling in the silts and clays. The more evenly bedded and extensive silts and sands may belong to the prodeltaic zone as well. The lack of evi- dence for surface exposure and root-bio- turbation suggests generally subaqueous conditions, with water depths greater than the maximum tolerated by aquatic vegeta- tion (about 1-2 m). The poorly sorted gravelly sands are re- sti'icted to lenses that represent channels. Pebbles up to 6 mm in diameter occur in these lenses as floating grains in a coarse sand matrix. Mudclasts are also present. The combined evidence suggests at least periodic currents over 100 cm/sec, and possibly flood deposition of the kind lead- ing to the very poor sorting and large floating grains (Pettijohn, 1957:254-255). (Vertebrate bone fragments and teeth are often extremely abundant in these gravelly sands, and include a high proportion of nonaquatic forms, in contrast to the aquatic assemblages derived from laterally associ- ated lithologies.) Cross-stratification is often well-developed in the medium- to fine-grained sandstones. These include small-scale structures com- parable to "Kappa" and "Nu" cross-strati- fication that indicate linguoid ripples ( Fig. 18). The well-sorted medium to coarse sands show planar foresets, and in some cases the cross-stratification suggests beach or barrier bar deposition comparable to that reported for recent barrier environments (Davies et «/., 1971). Current directions for the various forms of cross-stratification are highly variable^. The bed forms and grain sizes indicate water movement in the lower flow regime. The deltaic margin interpretation of 130- 0201 agrees well with the lacustrine and deltaic models of Visher (1965) for sedi- ment types and bedding characteristics. The transgressive sand-pebble sheets of the Wealden Lake and 103-0256 are absent or poorly developed. Instead, the delta of 130-0201 appears to have been continu- ously aggrading into a subsiding basin, with occasional periods when sediment accumulation overtook subsidence and shal- low water features (root casts, sand and gravel lenses) developed. Hydraulic equivalents for the bones range up to 50 mm, which is much larger than the maximum size of other associated particles. However, when large aquatic animals are eliminated (e.g., hippopotamus and crocodile), the mammalian remains have an estimated maximum hydraulic equivalence of 20 mm and most are less than 10 mm. This is closer to the matiix grain size in the channel lenses. The bones that are close to being hydraulically equiva- lent to their matrix grains also show more evidence of abrasion and weathering. These may have been carried to the delta margin during periods of high discharge (i.e., floods), and therefore may be derived from a variety of upstream source areas. Locality 105-0208: Delta Margin and Lagoon. The sediments are predominantly silty sands, poorly sorted and ripple-laminated with abundant mica. These form a recog- nizable 2-3 m thick unit over much of 105, bounded above and below by finer units of silty clays. Abundant \'ertebrate bone occurs in association with the silty sands but is rare in the silty clays. The silty sands 518 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 Cross-Stratification typical of the sediments sampled for vertebrate fossils. (Localities in parentheses) Beta-cross-stratification (102-0201, 103-0267) Mu-cross-stratification (102-0201) Kappa-Cross-Stratification Nu-cross-stratification (130-0201, 105-0208) (130-0201, 105-0208, 103-0267) Onikron-cross-stratifi cation (102-0201) Pi -cross-stratification (102-0201) (From J. Allen, 1963) Figure 18. East Rudolf Paleoecology • Bchrcnsmcycr 519 arc characterized by liorizontal, cvcmly bedded cosets of small-scale "Kappa" and "Nu" cross-stratification (Fig. 18). The individual beds are between 2 and 5 cm in thickness. Contorted bedding and un- even lenses of silt are present, but the bedding shows little evidence of dis- turbance from bioturbation. Bedding struc- tures indicate aggradation from the advance of successive ripple fronts, such as might be expected in a prodeltaic, lagoonal en\'ironment. Coarser sands are interbedded in dis- continuous sheets and lenses. In one lens, the overall characteristics suggest a barrier or beach bar. This sand overlies the mud- cracked surface of a thin lens of silty clay, and incorporates clay clasts in its lower 10 cm. The sand body is elongate and extends for over 100 m before pincliing out. Large tubular structures resembling root casts are abundant. Upward the bed becomes better sorted and has well-developed low angle planar cross-bedding that closely resembles the cross-stratification reported for barrier and beach environments ( e.g., Davies et al., 1971). The more tabular, thinner sand bodies interbedded in the siltv sands are com- monly cross-stratified, with single sets of planar and concave-upward laminae. Shal- low troughs are also common. Ripple formation at variable current velocities and depths is indicated, as in 130-0201. The sands pinch out into discontinuous nodular layers. Root casts are common, in associ- ation with the sheet sands, but mudcracks at the lower bedding contact are rare. The sands occasionally preserve a variety of fresh-water mollusks that are unbroken and locally autochthonous, including BeUamya, Cleopatra, Melanokles, Pila and Pseudo- hovaria. The invertebrate fauna indicates "prodeltaic or even marshy" conditions (D. Van Damme, personal communication). There are no poorly sorted, coarse-grained channel lenses within this part of the Area 105 section, in contrast to 130-0201. The overall sedimentary characteristics suggest a lagoonal (>nvironment, with sand and silt pr(nided from nearby distributary mouths. This compares well with delta margin con- ditions in actively aggrading sectors of the Omo Delta (Rutzer, 1971a:75). Beach ridges and barrier bars formed at the lake- ward side of the lagoonal complex and occasionally transgressed shoreward over lagoonal sediments. The water in the lagoon probably varied in depth with shal- lower phiises represented by coarser sand lenses with root casts indicating the spread of shoreline vegetation. The area may have been periodically (perhaps seasonally) sub- aerial, although most characteristics indi- cate overall shallow subaqueous conditions. Localities 10.5-0208 and 130-0201 are closely comparable in stratigraphic position within the Koobi Fora Fm. Both lie near the top of the Lower Member; 105-0208 is about 8 m below the KBS Tuff, and 130- 0201 between 10 and 15 m below the tuff. 130-0201 is probably the older of the two. The localities are about 15 km apart, and represent related depositional situations on the margins of the prograding delta system. Rones of aquatic and nonaquatic animals are abundant and well preserved, and often consist of associated skeletal parts. Bone surfaces are generally fresh, with only occasional evidence of predepositional weathering and abrasion. Most of the bones of nonaquatic animals are fragmented, with spiral and saw-tooth fractures indicating predepositional breakage. The largest bones are of hippo, and these reach hydraulic equivalents of 15-30 mm, well outside the sediment range for the coarser sands. Most of the other bone frag- ments and teeth are between 1.0 and 20 mm in quartz equivalent sizes. This overlaps the size range for other sediment in 105- 0208, but most of the bones, and particularly the teeth, exceed 2 mm in e(iuivalent size. Larger sediment grains occur in laterally associated facies to the east and northeast, but are absent in 105-0208. The combined cN'idence points to a local source of bones from the lagoon and shoreline environ- 520 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 ments, with perhaps a small component from the distributaries, including some floating carcasses. Bones were probably redistributed and buried during the mi- gration of beach sands, a process similar in some respects to that proposed for 103- 0256. Localitij 105-1311: Channel Complex. The fossil-bearing unit is up to 4.5 m thick and overlies an erosional surface with up to 12 m of relief. Coarse gravels, includ- ing CaCO.s nodules reworked from the underlying beds, arc concentrated primarily near the base of the unit. The lithology is consistently a coarse sand with lenses of gravel. As shown in Figure 16, the main body of the sand is linear, with current directions indicating a west to southwest bend. To the south, the sands and gravels intertongue with silts and silty clays repre- senting levee and floodplain deposition lateral to the channel. Well-developed large scale cross-stratifi- cation is present throughout the sand unit. Troughs are the most common form and are between 20-50 cm in diameter. Gravel lenses are present at the bases of many of them. These compare well in morphology and size with "Pi" cross-stratification (Fig. 18) and with the cross-strata occurring in ephemeral streams in Central Australia (Williams, 1971). Such stratification is formed by downstream-migrating ripples of varying size in the lower flow regime (Allen, 1968:110; Williams, 1971:37). All the above characteristics indicate that the 105-1311 sand body is a fluvial channel. The upward fining of the sands is typical of point bar formations (J. Allen, 1965:140), and it is likely that much of the sand was deposited in the point bar formed by the lateral migration of the channel bend. Root casts are abundant in the sands and laterally related silty clays, but are less common in the coarse gravels near the base of the unit. This indicates vegetation lateral to the active channels, and it is likely that a gal- lery forest existed along the channel. The geologic evidence does not reveal the extent of this forest, or whether the water flow was permanent or ephemeral. The fossil bones in 105-1311 form dis- tinct groups according to surface texture. In one group, bones are highly rounded and polished and are less than 5 cm in diameter. They can accurately be described as "bone pebbles." These compare well with the second-cycle bones of Rief (1971), which were mineralized prior to final trans- port and burial. Although it is possible for bones to be thoroughly mineralized in relatively short periods of time (e.g., 5000 years for bones from Lake Rudolf Holocene deposits), the degree of rounding of the bone pebbles in 105-1311 indicates long- tenn abrasion. It is more likely that they were derived from earlier fossiliferous sedi- ments associated with Miocene volcanics to the east than from floodplain deposits associated with the 105-1311 channel. The second group includes a wide range of sizes of relatively well-preserved teeth and bones. These are generally fragmental, and show signs of abrasion in their rounded edges, broken processes and exposed tra- beculae. A third group consists of only a few specimens, including whole skulls col- lected outside the sampling areas, which show little or no weathering or abrasion. The latter two assemblages are composed of bones that had not been mineralized prior to transport, and that had undergone variable degrees of surface weathering and abrasion prior to burial. These can be referred to as "first cycle" bones. The largest bone fragments are hydrauli- cally equivalent to quartz particles up to 40 mm, and most of the teeth fall in the 5-25 mm range. This is well within the particle size range of the associated gravels, which range up to 60 mm in maximum diameter. A very different taphonomic situ- ation exists in 105-1311 compared with the three localities discussed previously, which have bones that exceed the associated quartz particles in hydraulically equivalent grain sizes. It is clear that the bone as- East Hitdolf Paleoecology • Behrensmeyer 521 scmblagc in 105-1311 is much more likely to reflect the processes that ha\'e affected the associated sediment, i.e., abrasion and sorting through hydraulic transport. The same forces tliat moved sediment through the channel could also have moved the bones. A large proportion of these are probably deri\'ed from upstream sources, with a more local component derived from the imdercutting and reworking of previous floodplain deposits by the laterally migrat- ing channel. Both of these assemblages should consist of isolated teeth and the more durable parts such as ends of limb bones, all showing some degree of abrasion. The third component, consisting of the best-preserved material, would come from bones left in the immediate vicinity of the channel and rapidly buried. The bone assemblage of 105-1311 is thus a mixture of autochthonous and allochthonous ma- terial, and most of the bones show the effects of being in a fluvial system. Since the general environment of deposition is fluvial, the bones should belong to animals found in tiie floodplain or channel habi- tats, as opposed to the deltaic or lacustrine habitats. Locality 102-0201: Channel. The sequence is tilted some 15-20° west both in Area 102 and its continuation in Area 103. Current directions indicated in the sand are dominantly NNE to SSW, so that the strike of the beds is roughly parallel to the current. The deposits of 102-0201 overlie a scoured surface on silty clays with paleosol development, and they are followed by widespread sheet sands with stromatolites and shell debris. The stromatolites indicate shallow-water la- custrine conditions (S. Awramik, personal communication). 102-0201 represents a brief period of channel cutting between two longer lacustrine and deltaic deposi- tional phases. The dominant lithology is a coarse sand with gravel near the base, fining upward to medium and fine-grained sand and finally to silt. There is no obvious trend to\\^ard downstream fining in the 3 km segment examined. Large-scale lenses of gravel up to 1.5 m thick are common in the lower 3 m of the unit. The upper 2 m have only occasional small gravel lenses and dispersed pebbles. Mudclasts and carbonate nodule clasts, which are abundant near the base of the unit, appear to be derived from the underlying silty clays. Otherwise the gravel is composed of mixed quartz, feldspar, volcanic material such as welded tuff and pieces of silicified wood, all well- rounded. There are a few polished bone pebbles and occasional large polished bone fragments indicating a source of previously mineralized material. Cross-stratification includes planar fore- sets 10-25 cm in height, and a variety of trough cosets. Many cosets of the planar cross-beds are comparable to "Beta"-type stratification (Fig. 18). In some cases the cross-strata are more upwardly concave than planar, comparing well with "Mu" and "Omikron" stratification (Fig. 18), Allen (1963:110) attributes the formation of the latter types of cross-strata to mi- grating asymmetrical ripples. "Beta" cross- strata result from the downstream migration of single, straight edged ripple trains over a planar eroded surface (J. Allen, 1963: 102 ) . It seems that both of these conditions of ripple bedding, plus intermediates, took part in the formation of 102-0201. The troughs are generally large-scale ( 10-50 cm across) and compare with Allen's "Pi" or "Nu" types of cross-stratification (Fig. 18). These are attributed to the migration of large-scale asymmetrical ripples with curved crests and projecting lobes or tongues (J. Allen, 1963:110)." All of the above structures can be formed by flowing water in the lower flow regime. The evidence is conclusively in favor of a channel origin for the 102-0201 sand. The gravel concentrations near the base repre- sent channel bars and channel lag deposits. In one case, where the coarse material in- cludes an unusual amount of bone, bedding 522 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 structures and local upward fining suggest point bar fomration. Root casts are more common in the finer sands of the upper part of the unit than in the gravels. Verte- brate fossils show a sharp upward decrease in abundance and were clearly concen- trated along with coarse sediment near the base of the channel. The stratigraphic context of 102-0201 indicates much closer proximity to the lake than for 105-1311. In fact, 102-0201 can be regarded as a channel or complex of chan- nels incised into a temporarily inactive delta. Evidently, base level was lowered due to either tectonic or climatic processes. The period of cut and fill separating two deltaic-lacustrine units may reflect one of the local tectonic events which affected this part of the Koobi Fora Fm. during its de- position (G. D. Johnson, personal com- munication ) . The channel-cutting and gravels of 102- 0201 may be the downstream counterpart of 105-1311. Both lie near the base of the Upper Member of the Koobi Fora Forma- tion, and are in the MetridiocJioerus faunal zone (Fig. 14). The composition of the gravels is similar, and some evidence for an extensive erosion surface analogous to that in Area 105 has been found in the vicinity of Area 103. If 102-0201 is the deltaic- distributary counterpart of 105-1311, then it is probably slightly earlier in time. After an erosional phase, the areas closer to base level (i.e., 102) would begin to aggrade earlier than more upland areas such as 105. The fossil vertebrate material is variable in surface texture and overall preservation. Bones of aquatic and semiaquatic forms show minimal abrasion and are often com- plete. Other vertebrates are represented by teeth, limb parts, etc., usually broken and weathered. This indicates probable transport and a subaerial source (i.e., chan- nel banks) for the bones of nonaquatic animals. A few relatively complete parts, such as a complete rhinoceros jaw, indicate closer sources and less transport. The bone fragments are occasionally over 50 mm (e.g., the rhinoceros jaw) in hydraulically equivalent quartz sizes. How- ever, most are equivalent to grains less than 20 mm, and thus are similar to the size range of the gravels. As in 10.5-1311, most of the bone in 102-0201 has probably been subjected to winnowing and abrasion dur- ing transport. The close association be- tween bones and gravels in 102-0201 im- plies similar concentrating processes. This may be an example of Langbein and Leo- pold's "kinematic wave" effect (1968), where large particles tend to concentrate other large particles and form gravel bars. The sediment particles, including bones, are a mixture of allochthonous and autoch- thonous material. The more complete skeletal parts, the mudclasts and the armored mudballs, are examples of locally derived material from the channel banks or channel bed. The gravels, including the polished bone fragments, have been trans- ported from upstream sources. The largest proportion of bones and teeth may have either local or distant sources, and prob- ably represent animals which inhabited channel and floodplain environments as well as the temporarily dry and emergent deltaic plain. Locality 103-0267: Distributary complex. The fossil-bearing horizon is exposed in widely separated areas covering over four square kilometers. The dominant lithology is a poorly sorted gravelly sand. The sands are of variable thickness and occasionally cut several meters into the underlying beds. Coarse sediment fines upward and inter- tongues with silts and silty clays near the top of the unit. The 103-0267 sands and gravels overlie the Koobi Fora Tuff, which is predomi- nantly lacustrine in origin and is capped by a widespread, oolitic carbonate sand with stromatolites. The deposits of 103-0267 are followed by lacustrine silts and shell beds. Thus, the channeling and sand deposition represent a brief period of subaerial ex- posure and erosion similar to that of 102- East Rudolf P.vleoecology • Behrensmeyer 523 0201. About 50 m of continuous section separates the two units. G. D. Johnson (personal communication) has suggested that a tectonic event mav be responsible for 10.3-0267 as well as 102-0201. The upper part of 10.3-0267 is occasion- ally characterized by a discontinuous hori- zon of CaCO.T concentration with abundant root casts. The root casts are truncated by the overlying sediment. CaCOs layers are formed of linked, irregular nodules which become more massi\e upward. This la\er appears to bear a primar>- relationship to the associated sediments; i.e., it fomied at the time when the top of 103-0267 was a land surface. The carbonate la\er is thus tentativeh- identified as a caliche. It is comparable in sti-ucture and form to caliches of the .\merican Soutliwest (Reeves, 1970; Aristarain, 1962; Bretz and Horberg. 1949). Although the processes leading to caliche fomiation are not well known, seasonal upward and downward percolation of ground water is usually indi- cated b\' such carbonate concentrations in soil horizons (Reeves, 1970: 353). Cross-stratification is more widely vari- able in scale than in the 10.5-1311 or 102- 0201 channels. The largest sets are up to 20 m across and are broadly concave up- ward. They compare with "Pi" cross-strati- fication (Fig. 18) and "festoon" bedding of the mega-ripple zone (Msher, 1965:47). A variet)- of smaller scale cross-stratifications are also present, including "Beta" and "Xu" t\'pes (Fig. 18). Troughs are well developed in sandy gravels near the base of the unit, while the festoon bedding occurs near the middle in coarse sands with gravel lenses. Characteristics of 10.3-0267 suggest a distributar\' complex, with some redistri- bution of sediment by shoreline processes. Current directions are highl\- variable, from NW to S. The deposits represent laterally extensive channel cut and fill with sub- sequent aggradation over emergent deltaic flats. The large-scale cross-strata indicatc> distributary channels with flow depths of several meters. This contrasts with the channels in 10.5-1311 and 102-0201, which lack cross-stratification of comparable scale and probabK- carried shallower flows. The bones of 10.'3-0267 are concentrated in the lower 3 m, and are usually associated with pebbles of about 1 cm in diameter. Large-scale gravel and bone concentrations such as in 102-0201 are absent and bones are more or less evenh' dispersed o\er tlie area co\ered b\- the deposit. There is a mLx- tiire of bone surface textures indicating \arious kinds of weathering and abrasion before burial. Parts of aquatic animals are generally the best preserved. Second cycle "bone pebbles" are present, as in the channel deposits of 102-0201 and 10.5-1311. Grain size equivalents for the bones range up to 30 mm in diameter, but most fall between .5-15 mm. Since grain sizes in the gravels are up to 30 mm, the bones are within the o\erall sediment size range. Many have been transported, and the as- semblage includes both autochthonous and allochtlionous bones, as in the 102-0201 and 105-1311 channels. Locality 8+6-0104: Floodplain. This unit is composed of lithofacies unique to the upper part of the Koobi Fora Fm., occurring only in the Ileret Mb. and in the Upper Mb. in Areas 130 and 131 (Fig. 12). The dominant lithology is a lidit-colored tuffaceous silt. The environ- ment of deposition evidently extended over a wide area, and the silts are exposed in Areas 6 and S, which are some 2.5 km apart. The unit is stratigraphicalh' marked by the "middle tuff complex," which inchides locally discontinuous lenses of reworked \olcanic ash and pumice. The silts are remarkabl\- consistent in textine and appearance. They are inter- bedded at regular inter\'als with zones of silt>- clays. These show \ertical prismatic structure and cla>- concentrations suggest- ing paleosol development. Zones of CaCO:< nodules occin- within the silts and at con- tacts of silt>- clays on silts or sand>- silts. The nodule horizons are often laterallv continu- 524 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 ous and formed of elongate or flattened, irregular carbonate concentrations. Inter- nally these are composed of fine sand float- ing in structureless micrite. They vary in size from 2-15 cm maximum diameter. Smaller nodules of CaCOa are dispersed throughout the clays and silts. The sedi- ments themselves have very little dispersed carbonate, and do not react to the HCL test. Fragments of nodules are incorporated in sand lenses representing small channels interbedded in the silts. In some cases the nodule horizons are truncated by later beds. The combined evidence leads to the con- clusion that the carbonate concentrations formed during the deposition of 8+6-0104. The nodule horizons can best be ex- plained as incipient caliches or carbonate concentrations formed in the "B" soil zones of successive subaerial deposits. Lobova (1967:290-299) describes the formation of similar carbonate concentrations in desert soils of the USSR. He suggests they are formed by biogenic carbonate concentrated in water percolating downward from the surface which later evaporates, leaving CaCOs precipitates. These nodule horizons commonly form at depths of 20-60 cm be- low the surface. The presence of such horizons in primary association with the sediments of 8+6-0104 indicates seasonal fluctuations of water content in soils with a local (biogenic?) source of CO;r and a source of Ca^^ (clays?). The absence of extensive, thicker caliches is perhaps due to the steady aggradation of the floodplain, with continued burial of former land sur- faces. Root casts are abundant throughout the unit. They are usually less than 1 cm in diameter and are formed of CaCOs similar to that found in the nodule horizons. CaCOa-filled root casts are also found in the desert soils of the USSR, and are used as evidence of biogenic formation of carbonate concentrations (Lobova, 1967: 290). Some of the root casts are truncated by the channel scour-and-fill structures. The silts are riddled with tubes which may be burrows rather than root holes. These are usually 1.5 mm in diameter and have distinctive clay rims. One well-developed channel can be traced NNW across the exposures in Area 8. It is approximately 40 m across and is filled with medium- to coarse-grained sand plus pumice cobbles up to 10 cm in diameter. Some of the silt beds (and per- haps the Area 8 lens of "middle tuff) represent levee and overbank deposits from this channel. Other channels occur within the silts and clays. Most are small scale, with variable current directions. The chan- nel sands are often well sorted and ce- mented with CaCOs, and in some cases the cemented sands weather out as rounded, resistant blocks and nodules. It would be difficult to assign 8+6-0104 to any environment other than a floodplain. In general, the characteristics fit Allen's (1965) concept of vertically accreting flood- basin deposits. The whole complex of small channels and silt deposits may represent a zone intermediate between deltaic fan and floodplain, similar to that 1 to 2 km east of the margin of the present-day Tulu Bor Delta at Ileret. In Area 6, the "middle tuff" fonns a widespread, mudcracked surface indicating deposition in a pond or lagoon, with later desiccation. It is possible that more deltaic conditions existed farther to the west of Area 8 in Area 6. The bones of 8+6-0104 are generally very well preserved and often covered with a CaCOs crust. Some show surface weather- ing and cracking, and there are abundant isolated teeth. Associated skeletal parts of terrestrial mammals are also fairly common. This evidence suggests variable degrees of surface weathering and rates of burial. The bones in 8+6-0104 are associated with much smaller grain sizes than in the channels. Hydraulic equivalents of most of the bones fall well above the 1 mm maxi- mum grain size of the silts and sandy silts in which they occur. If individual bones had been carried in the channels and spread over the floodplain during floods, then they East Rudolf Paleoecology • Bcliiciisnwycr 525 should be found in association with grains If the composition of tlic bone assem- closcr to their hydrauHc e(iuivalents, i.e., blages is hnkcxl to sedimentary processes, coarser sand and gravel. Sediment of this then the channel assemblages should be size is available in channels lateral to the more like other channel assemblages than silt deposits. Since it is not found with the like floodplain or deltaic assemblages. Th(^ bones, and since these show a gcMieral lack characteristics that should be similar within of abrasion, most of the bones are probably similar deposits include the degree of bone- autochthonous to the floodplain environ- sorting and the degree of weathering and ment. The presence of associated skeletal abrasion. In the extreme case, the de- parts may indicate carcasses buried in situ positional processes could sort and partially or floated in during the floods. Most of the destroy a given thanatocoenose so as to bones probably were buried by the periodic obscure all of the original ecological infor- influxes of floodstage silts. The trapping mation in the assemblage, effect of floodplain vegetation may have Thus, the first step in recovering eco- been influential in anchoring the bones logical information from East Rudolf as- until they could be buried. Some of the semblages is to isolate those cases where lighter elements may have been dispersed the effects of depositional processes are by these floods, but most of the thanato- minimal. The evidence presented so far coenose remained in place as a lag deposit against extensive alteration of a thanato- to be covered, or destroyed by later coenose by sedimentary processes includes: wea leimt,. ^^ Bones with fresh, unabraded surfaces 2) Complete bones, sk-ulls with teeth and Discussion and Conclusions ^^^.^^^^ structures intact The seven localities can be grouped into 3) Associated skeletal parts (indicating three broad categories on the basis of simi- l^ck of reworking) larities in lithofacies: On these criteria, the assemblages of 1) Delta: 103-0256, 130-0201, 105-0208, Localities 103-0256, 105-0208 and 8+6-0104 /■iQn_9Qgy\ have been least affected by depositional 2) Channel: 102-0201, 105-1311, (103- processes, and retain a maxiinum amount of 0'?67') paleoecologic information. The other loeali- 3) Floodplain: 8+6-0104 ties have assemblages with mixed histories, and ecological information may be more These groupings are similar in lithology, difficult to isolate. bedding structures, and lateral facies Fossil assemblages from the channel relationships. The deltaic localities are environments (including 103-0267) are more diverse in these characteristics than similar in that they all bear evidence for the channels, with 103-0256 representing a bone abrasion and include mixed autoeh- transgressive beach, 130-0201 distributaries thonous and allochthonous material. The and a delta margin, and 105-0208 a beach lacustrine-deltaic environments are less and lagoon complex. 103-0267 can also be similar among themsehes. with Locality regarded as deltaic, since it represents a 130-0201 combining the characteristics of distributary complex rather than a single, transported and nontransported assem- well-defined channel. However, its lithol- blages, while the others appear primarily ogy and sedimentary structures are more untransported. In general, however, it ap- like those of the channels. Henc(% it is pears that some aspects of the bone as- intermediate between the deltaic and chan- semblages are similar in similar lithologies, nel groupings, and is included parentheti- and thus reflect the processes operating in cally in both. the different sedimentary environments. 526 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 SORTING IN BONE ASSEMBLAGES OF THE KOOBI FORA FORMATION The primary object of this section is to estabhsh the relative numbers of different skeletal parts in the seven bone assemblages and to discuss their taphonomic implica- tions. Different skeletal parts have very different potentials for dispersal, as dis- cussed in the section on bones as sedi- mentary particles. Depositional processes operating on bones should affect the ratios of skeletal parts, particularly those that have widely different densities, such as teeth and vertebrae or phalanges. Assem- blages that have a concentration of elements with similar dispersal potential indicate sorting of the original components of the thanatocoenose. Assemblages with a mix- ture of heavy and light, large and small bones indicate either less alteration of the thanatocoenose before burial, or a mixture of bones with different taphonomic histories. Most skeletons are incomplete when their parts become sedimentary particles, due primarily to destruction by carnivores. The initial assemblage, after carnivore activity (such as in East Africa today), consists of teeth, skulls, horn cores, vertebrae and limb ends, with more parts surviving for large animals than small. This results in an assemblage of bones with a wide range of sizes and densities, which will be subject to sorting in transport situations. The bones and teeth also have different survival po- tentials in most situations, with the former being more readily destroyed by tapho- nomic processes than the latter. Sampling of Bone Assemblages Most of the fossil vertebrates of the East Rudolf deposits occur in surface lag con- centrations due to the removal of surround- ing sediment. In general, movement of fossils away from their source rocks is minimal, and they remain in clear associ- ation with particular sediments. Conditions of preservation and recent erosion are such that even delicate fossils usually remain reasonably intact, once exposed, and com- pact objects such as teeth may last for long periods of time ( 100+ years? ) on the sur- face. This provides a large amount of accessible material for collection. Bones from the seven localities described in the previous section were collected using the following procedure: Grid squares of 10 X 10 m were laid out over the chosen area of outcrop. The first square in each locality was positioned using an arbitrary spot on an aerial photograph or simply by selecting a local landmark (e.g., a tree or conspicuous outcrop), without specific reference to the degree of surface bone concentration. Subsequent squares were measured off from the first, with a mini- mum of 20 m between squares. On hori- zontal sti'ata, the squares were laid out on an orthogonal 30 X 30 m grid. On dipping strata, the squares were positioned along the strike of the units being sampled. The grid system was adjusted, where necessary, to avoid patches of recent sediment and vegetation. The selection of squares was not adjusted to sample particularly attrac- tive patches of bone fragments, in order to prevent subjective biasing of the bone samples. Collecting was done by system- atically traversing a square first east-west, then north-south (for a square oriented NSEW). All the surface bone larger than 5 cm (mamixum length) was collected in addition to those smaller bones that could be identified to class (Fish, Mammal, Rep- tile, Bird). During the first field season all samples were removed for identification and study. During the second season, after workers were familiarized with the verte- brate taxa and skeletal parts, it was pos- sible to do most identification in the field. This greatly simplified the logistics of the sampling, and enabled workers to leave the field with a card for each square recording taxa and skeletal elements plus geological data. This was a welcome alternative to carrying out 50-60 lbs. of fossil bone frag- ments after each day of collecting. East Rudolf Paleoecology • Behrensmeyer 527 Table 6. SAMPLING LOCALITY 130-0201 105-0208 103-0267 103-0256 102-0201 105-1311 8+6-0104 Stratigraphic data and sample size of the seven fossil sampling localities. Sample squares are 10 x 10 meters, representin(; 100 m^ each. # OF SAMPLE SQUARES 21 20 20 27 34 25 66 STRATIGRAPHIC INTERVAL SAMPLED (IN METERS) 7.0 2.5 3.0 .75 5.0 3.0 4.5 BASIC LITHOLOGY Sand, silt, and clay Sand, silt, and clay Sand and gravel Sand Sand and gravel Sand and gravel Silt GENERAL DEPOSITIONAL ENVIRONMENT Delta margin Delta margin and lagoon Distributary- beach complex Transgression over deltaic mudflats Channel Channel Floodplain KOOBI FORA FM. STRATIGRAPHIC FAUNAE UNIT UNIT Mesochoerus Mesochoerus Metridiochoerus Metridiochoerus Metridiochoerus Metridiochoerus Loxodonta Lower Mb. , Koobi Fora Fm. Lower Mb. , Koobi Fora Fm. Upper Mb. , Koobi Fora Fm. Upper Mb. , Koobi Fora Fm. Upper Mb. , Koobi Fora Fm. Upper Mb. , Koobi Fora "^m. Ileret Mb., Koobi Fora Fm. Maps of each locality showing the positioning of the sample squares are given in Figures 15 and 16, and the number of squares collected in each locality is given in Table 6. The major problems encountered in the sampling were: 1) choosing localities that showed a clear relationship between the surface bones and the sedimentary units, 2) obtaining comparable samples from each locality that adequately repre- sented the bone assemblages. Choosing the Sample Areas The primary goal was to collect an assemblage of bones that represented the material buried in a well-defined sedi- mentary deposit. In selecting the sampling localities, the following guidelines were established: 1) A locality was chosen on beds, or a series of beds, representing deposition in one of three broad environmental categories: channel, floodplain or delta. 2) The topographic situation was such that contamination of the fossil con- centrations with material from other horizons was minimal. Efforts were made, for example, to sample beds on drainage divides rather than in val- leys. 3 ) Vegetation and recent sediment in the area were minimal. 4) Previous collecting in the area was minimal, or collection sites were marked and the removed fossils re- corded. 5) The locality was extensive enough so that a representative sample of the fossil assemblage could be collected. Fortunately, the East Rudolf region pro- vided many areas that satisfactorily met all these requirements. Since stratigraphic series of environmentally related beds rather than single beds were used, the chances of contamination from different series of beds representing different de- positional environments was greatly re- duced. In the course of sampling, the actual bone-producing beds were often indicated by matrix adhering to fossils, and some of the samples could be assigned to particular horizons. Such evidence further supported the association of bones with the enxiron- mental units of interest. The advantage of sampling different lithologies that are genetically related (e.g., sands, silts, and clays, all deposited in deltaic conditions) is that this will give a more general picture of the faunal and skeletal elements preser\'ed in a rather broadly defined environment. This con- 528 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 trasts with sampling a particular bed ( as in some quarry deposits ) , which is more likely to be the result of very local or special conditions. The sampling method described above allows coverage of extensive areas (square kilometers) of outcrops represent- ing single, broadly defined sedimentary environments. This permits sampling on a scale more comparable to the habitat sizes of many East African vertebrates (on the order of square kilometers to thousands of square kilometers). Sampling by widely spaced squares should establish faunal and bone abundances that represent broad-scale differences between sedimentary environ- ments and the habitats associated with them. Moreover, sampling through several meters of sedimentary strata representing extended periods of time should reveal more general pictures of bone and sediment associations than assemblages representing single events. Sample Size Surface bones are so abundant in the sampling localities that even a few 10 X 10 meter squares provided large numbers of fragments, and over 9,000 were collected in the total sample from 213 squares. More than 7,000 {787c) of these were identifi- able as to skeletal part or vertebrate group or both. Very few of the sample squares lacked fossil material, even though they were laid out without regard to fossil dis- tribution. The abundance of fossil material was suf- ficient to provide an average of 34 identifi- able pieces per square and to give a good representation of the most common parts and animals. Field collecting was aimed at obtaining the largest possible compar- ative samples from all the localities. Since the surface concentration of bone varied from locality to locality, the number of squares collected in each varied as well. Thus, it was necessary to collect over 60 squares for 8+6-0104, which had a low surface concentration, but only 20 for 105- 0201. At least 20 squares (=2000 m-) were collected in each locality. Method of Representing Fossil Abundance It is possible to represent the relative abundance of different bones in more than one way. For instance, within each locality the total number of fragments identifiable as vertebrae can be compared with the total number of tooth fragments. Percentage representations of these totals can be com- pared between localities. Alternatively, the total number of squares with vertebrae can be compared with the total number of squares with teeth, etc. For reasons de- scribed below, the second method of repre- senting relative abundance is used in all the following analyses of the fossil assemblages. Difficulties in using total numbers of parts for comparative purposes include the following: 1) One tooth, for instance, can weather on the surface into dozens of frag- ments which are still identifiable as teeth, but a vertebra may only pro- duce a few fragments that can defi- nitely be identified as vertebrae. In both cases the numbers of broken fragments, if totaled, would count for more than the whole elements, and give erroneous data on the relative numbers of these elements. This prob- lem is particularly pertinent to a frag- mented surface sample, and is almost impossible to correct for by attempt- ing to calculate the "minimum num- bers" of fragments per bone in the manner of Shotwell (1955). 2) A single skeleton, if disassociated prior to burial or during recent erosion, may be counted as several individuals of the same animal group, while the whole skeleton would be counted as one individual. This can lead to errors in representing the actual abundance of different ani- mals. Shotw ell's method of using mini- East Rudolf Paleoecology • Bchrensmeyer 529 muni numbers of indi\'iduals^ lielped to resolve this probl{>m for his quarry samples (1955). In the East Rudolf surface assemblage*;, with many verte- brate groups represi'uted by a wide range of identifiable bone fragments, the minimum numbers method was not feasible. The more satisfactory method of repre- senting bone abundance for the East Rudolf localities is to use the number of squares with a particular skeletal part. This is done as follows: if one vertebra, or several, or dozens of pieces of the same one, occur in a sample square, this is counted as 1 oc- currence. If one tooth of the same taxon occurs in each of 5 squares, this is counted as 5 occurrences. The number of occur- rences of each bone can be converted into a "square frequency" by dividing by the total number of squares in each locality. Thus, 5 occurrences out of a sample of 20 squares gives a frequency of .25 or 25%, This method has a number of advantages \\'hich make it a valid measure of bone abundance in the broadly defined sedi- mentary units of interest for this study: 1) It gives a measure of the dispersed abundance of the different bones in space and time, which should be a result of the overall conditions of each sedimentary environment. 2) The problems encountered in using fragment totals are essentially elimi- nated, since using occurrences in squares will greatly reduce the effects of differential identifiability and frag- mentation of the surface bones. Also, since the squares are widely spaced, the probability of sampling parts of the same bone or even of the same animal more than once is very low. A comparison of the two measures of abundance, by fragment number and by ^ The relative abundance of different taxa is represented by the number of the most common similar skeletal part (e.g., left femora) of each taxon (Shotwell, 1955:331). squares, illustrates the advantages of the latter method. In Figure 19, the frequency of vertebrae in each locality is given ac- cording to total numbers of fragments identifiable as vertebra, and bv the fre- ciuency in terms of scjuares with vertebrae. Numbers of vertebral fragments that are high relative to the square frequencies, as in 8+6-0104, imply localized concentra- tions. In fact, for 8+6-0104 the large num- ber of vertebrae results from two associated partial skeletons of bovids. In contrast, a high square frequency and a low fragment number shows a widely dispersed sample of isolated vertebrae, as in 103-0256 and 103- 0267, where only one or two vertebrae occur per square. The representation of the dispersed abundance is more useful in comparing bone assemblages that result from interrelated processes in channel, floodplain or deltaic environments. The "square frequency" of bones ( = the number of squares with a particular bone or taxon divided by the total number of squares per locality) will thus be used in the following sections. Characteristics of the Bone Assemblages During the collecting of the bone sample, and prior to numerical analysis, it was ap- parent that some parts, such as teeth, were more abundant in some localities than others. However, most of the differences in bone proportions among the localities became apparent only after relative abun- dances were tabulated in the laboratory. The bone sample contains abundant skeletal fragments from mammals, reptiles and fish, and a few from birds. Analysis of bone frequencies is restricted mainly to the mammals, which form the largest and most diverse component of the sample. Fre- quencies of the skeletal parts are given in Table 7. Discussion of the method of identification, which can influence tlie ap- parent abundance of parts, will precede analysis of the data. 530 BuUetin Museum of Comparative Zoology, Vol. 146, No. 10 120 100 I/) I 80 i- • »*- o X) o 60 40 20 rl.OO .80 ,60 .40 s- X3 c te part (e.g., ends of limb bones, vertebral centra, whole phalanges, etc.). Significance of the Frequency Data The data given in Table 7 show that most of the skeletal parts are represented in each locality. Some have consistently high frequencies, such as teeth; some low, such as patellae, and some are variable. The lower frequencies indicate occurrence in only a few squares out of the total for each locality. Both high and low frequencies are of interest in comparing the samples. To assess the statistical significance of the frequencies, one must ask, "How repre- sentative of the actual bone assemblage in each locality are the 'square frequencies'?" In some respects the problem is comparable to establishing binomial sampling limits for accurately detecting character frequencies in any given population (Simpson et ah, 1960:199). In such cases, tables are avail- able for relating actual frequencies to ob- served frequencies using various sample sizes. For example, a character with 40% frequency in the actual population could vary from 12-35 occurrences in a sample of 60, with a probability of only .001 that fewer than 12 or greater than 35 occur- rences would be observed. The binomial sampling limits for square frequencies can be calculated using the Harvard Tables (1955). For a sample of 34 squares, a frequency of .32 (11 squares) could represent a possible range of actual frequencies between .17 and .48, with a probability of only p = .05 that the actual frequencies in the bone assemblage would fall outside of this range. The sampling error indicated by simple binomial prob- ability is potentially rather large. However, it can be assumed that the square fre- quencies are more closely representative of the true bone fre(|uencies because: 1), each sample square consists of a 10 X 10 m area, which greatly increases the probability of finding a particular bone if it is present in the assemblage and 2), many square's in- clude more than one bone of a particular kind, and the actual frequency is higher in these cases than representation by square frequency would indicate. Therefore, the square frequencies will l)e treatcxl as repr(^- sentative frequencies for the following data analysis. The bone abundances, as repre- sented by these frequencies, should be comparable from locality to locality. The statistical significance of specific differ- ences or similarities between localities was tested using Chi-Square analysis. Comparisons of Overall Bone Concentrations The relative concentration of identifiable bones varies greatly in the sample squares of the seven localities. Overall bone abun- dance can be conveniently expressed by dividing the cumulative total of bone oc- currences in squares by the number of squares in each locality. These figures are given for identifiable mammal and reptile parts in Table 7. Locality 8+6-0104 has the lowest concentration and 105-0208 the highest. The three channel assemblages are no more concentrated than the lacustrine- deltaic ones for mammals, but are slightly less prolific in terms of reptiles. There does not appear to be any consistent correlation between sediment grain sizes and identifi- able bone abundance in the deposits sampled. The localities with more bones per square do not appear to have more of any par- ticular elements. Rather, they show an increase in the frequencies of all skeletal parts. This implies better conditions for preserving bones of all kinds, regardless of size and density, and argues against ac- cumulation due to sel(>cti\'e proees.ses of sorting (which would tend to concentrate bones of similar sizes or densities or both ) . 532 Bulletin Museum of Co7nparative Zoology, Vol. 146, No. 10 Table 7. The square frequencies of reptile and mam- mal SKELETAL PARTS IN THE SEVEN SAMPLE LOCALITIES. FRE- QUENCIES ARE CALCULATED AS THE NUMBER OF SQUARES WITH A PARTICULAR ELEMENT DWIDED BY THE TOTAL NUMBER OF SQUARES IX EACH LOCALITY. ThE FREQUENCIES OF ASSOCIATED PARTIAL SKELETONS AND JUVENILE BONES ARE CALCULATED IN THE SAME MANNER. MaMMAL AND REPTILE BONES ARE COM- bined in the second listing to include those which could not be definitely assigned to one or the other class. This shows the relatively high proportion of rib and diaphysis fragments in the total bone sample. DELTA — CHANNEL— FLOOD- REPTILE PLAIN 130- 0201 105- 0208 103- 0267 103- 0256 102- 0201 105- 1311 8+6- 0104 Tooth .86 .85 .50 .33 .26 .68 .12 Skull/jaw .10 .05 .30 .07 .15 .08 .00 Vertebra .19 .05 .15 .19 .18 .00 .02 Limb .14 .05 .05 .15 .09 .00 .00 Scute .24 .40 .50 .30 .24 .40 .02 Phalanx .10 .15 .00 .04 .00 .04 .00 Carapace/ plastron .24 1.00 .70 .81 .24 .32 .12 # occurrences 1.9 2.5 2.2 1.9 1.1 1.1 .3 per square (average) MAMMAL AND REPTILE Tooth .95 1.00 .80 .59 .76 1.00 .67 Rib .76 .90 .85 .63 .53 .64 .32 Pelvis .14 .25 .25 .04 .06 .16 .05 Diaphysis .57 1.00 .95 .89 .71 .92 .55 Phalanx .52 .55 .35 .30 .15 .32 .12 Vertebra .57 .85 .55 .59 .29 .48 .15 Relative Abundance of Skeletal Parts The frequency data in Table 7 can be analyzed : 1 ) , in terms of the most common bones in each locality and 2), in terms of the correlations between localities caused by similar proportions of different mam- malian bones. Teeth are the most common mammalian element in all localities except 103-0256. Otherwise, the patterns of fre- quency are variable, with some indication that vertebrae and phalanges concentrate in the deltaic environments. In order to clarify possible correlations between lo- calities, two numerical analyses were used: a multiple regression analysis, which gives correlation coefficients for locality to lo- cality comparisons, and a Q-Mode Factor Analysis, which shows groupings of the localities in terms of skeletal parts. Correlations Based on Bone Abundance Figure 20 shows a correlation matrix re- sulting from multiple regression treatment of skeletal part frequencies in the squares. The correlation is "Pearson's product moment correlation" which assumes con- East Rudolf Paleoecology • Behrensmeyer 533 I Table nr 1 ■ 7 (CONT.) T A p U A FLOOD- DLL in tHAniNtL ' MAMMAL PLAIN 130- 0201 105- 0208 103- 0267 103- 0256 102- 0201 105- 1311 8+6- 0104 Tooth .67 .85 .70 .56 .62 1.00 .52 Jaw part .24 .10 .10 .04 .21 .08 .09 Maxilla .05 .00 .00 .00 .00 .00 .02 Cranial part .05 .25 .15 .07 .12 .08 .08 Horn core .19 .25 .35 .15 .18 .36 .03 Vertebra .48 .75 .50 .59 .15 .36 .15 Sacrum .00 .00 .05 .04 .00 .00 .00 Scapula .14 .45 .15 .15 .12 .20 .09 Pelvis .10 .20 .20 .04 .03 .12 .05 Humerus .19 .50 .30 .11 .06 .20 .14 Radius/ulna .14 .40 .20 .11 .15 .20 .15 Femur .14 .40 .05 .07 .18 .16 .08 Tibia .10 .30 .25 .07 .06 .28 .14 Patella .05 .05 .00 .00 .00 .04 .00 Metapodial .10 .40 .40 .22 .18 .32 .14 Astragalus .10 .20 .25 .11 .03 .08 .14 Calcaneum .10 .15 .15 .15 .00 .12 .08 Podial .10 .45 .10 .19 .24 .20 .18 Phalanx .48 .65 .35 .26 .15 .28 .12 Total # squares 21 20 20 27 34 25 66 # occurrences 3.4 6.4 1.2 2.9 2.4 4.1 2.2 per square (average) Associated parts Juveniles .04 .14 .20 .30 .05 .05 .04 .04 .00 .06 .00 .12 .06 .00 % hippo bones 18^; 16% 21% 6% 12% 1% 2% Total # squares for all localities: 213 Average occurrences per square: 690/213 =3.2 tinuous data and normal bivariate distri- butions. Both conditions are satisfactorily met by the squares data. Correlations are based on the five most common elements: teeth, vertebrae, phalanges, scapulae and radii/ulnae. An obvious feature of all the correlations is that they are high (> .5). This shows a basic similarit)' in the ratios of the five skeletal elements in all the sample assem- blages, although these elements vary greatly in size and densit)'. Therefore, the differences in the sedimentary environ- ments were not enough to alter the basic similarity of the thanatocoenoses sampled in each deposit. This similarity is probably produced by those bones most likely to survive carnivore activity and become sedi- mentary particles. Many of the correlations shown in Figure 20 are significantly different, in spite of the overall similarity. The highest and lowest coefficients differ significantly, with a probability of <.05 according to the "z test" 534 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 DELTA CHANNEL FLOOD- PLAIN 130- 0201 1 .000 1 .000 1 .000 1.000 1 .000 1 .000 105- 0208 .958 103- 0256 .851 .961 103- 0267 .950 .951 .915 102- 0201 .715 .629 .566 .816 105- 1311 .807 .751 .697 .897 .985 8+6- 0104 .716 .641 .595 .834 .995 .983 1.000 LOCAL- ► ITIES 130- 0201 105- 0208 103- 0256 103- 0267 102- 0201 105- 1311 8+6- 0104 V DELTA ■^ '^ CHANNEL^ FLOOD- PLAIN Figure 20. Correlation coefficients (Pearson's product moment correlation) between sampling localities accord- ing to the proportions of the five most common skeletal parts: teeth, vertebrae, phalanges, radii/ulnae and scapulae. Highest correlations show strong similarities between channel and floodplain environments in terms of the proportions of different skeletal parts. for significance (Simpson et ah, 1960:246). Other coefficients are indicative of trends even when their differences are not within the acceptable limits of significance (pC05). The coefficients show that tlie channel assemblages, 105-1311 and 102-0201, are closely correlated with each other and with the floodplain, 8+6-0104. The deltaic as- semblages have relatively low correlations with the floodplain, variable degrees of correlation with the channels, and high correlations among themselves. Thus the proportions of the five different bones are similar in similar sedimentary environ- ments, showing the effects of processes operating within these environments. Some of the close interenvironmental correlations, such as between the floodplain and channel assemblages, and between 130-0201 and 103-0267, suggest processes that are com- mon to more than one sedimentary situ- ation. These can be further clarified by examining which bones are influential in causing the interlocality correlations. Factor Analysis of the Bone Assemblages Factor analysis was used to indicate which skeletal parts cause similarities or differences among the seven bone assem- blages. The Q-Mode Factor Analysis, "CABFAC," was run on the frecjuency data from all of the mammalian skeletal parts. A solution of three varimax factors (axes placed within the data array) explains 97% of the total variance in the assemblages. The projection of the data for each locality on these axes is plotted on the triangle diagram shown in Figure 21. The diagram shows graphically how the three factors group (cluster) the bone assemblages. The three factors consist of 1) vertebrae East Rudolf Paleoecology • Behrensmeyer 535 METAPODIALS. TIBIAE. ETC. (Voorhies Group II ) ® DELTA CHANNEL AND FLOODPLAIN VERTEBRAE AND PHALANGES (Voorhies Group l) TEETH (Voorhies Group III) Figure 21. Triangle diagram showing the results of a three-factor analysis of the frequency data for all mam- malian bones. The factors correlate with Voorhies' dispersal groups, showing a relatively high proportion of Group I (most easily dispersed) in the deltaic assemblages and Group III (lag) in the channel and floodplain assemblages. and phalanges, 2) teeth, 3) hmb parts such as tibiae, metapodials, and astragaH. The triangle diagram shows a clear separation of assemblages on the basis of Factors 1 and 2. The three deltaic localities have a high proportion of vertebrae and phalanges, while the channels and the floodplain have high proportions of teeth. 103-0267 falls between the two groupings, and is some- what anomalous in its lack of similarity to the channel assemblages. Localities 103- 0267 and 8+6-0104 both show that there is no strict correlation between tooth con- centrations and coarse-grained sediment. 103-0267 is a coarse-grained deposit lack- ing a high proportion of teeth; 8+6-0104 is fine-grained, but is characterized by a high tooth concentration. It is clear that the high correlation coefficients between assemblages from similar sedimentary environments are due to tlie proportions of teeth, vertebrae and 536 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 Table 8. The relative frequencies of skeletal parts in a single skeleton, the average of bovid, SUID, EQUID and HIPPO SKELETAL PROPORTIONS. UNDERLINED PARTS ARE THOSE WHICH ARE MOST COMTvION OR MOST CONSISTENTLY PRESENT IN THE FOSSIL ASSEMBLAGES. No. in No. of each part/total No. in No. of each part/total average no. of parts in average average no. of parts in average skeleton skeleton (156) skeleton skeleton (156) Teeth 38 .26 Radii 'Ulnae 2 .01 Jaw 1 .01 Femora 2 .01 Maxilla 1 .01 Tibiae 2 .01 Cranium 1 .01 Patellae 2 .01 Horn Cores 2 .01 Metapodia 4 .03 Vertebrae 28 .18 Astragali 2 .01 Sacrum 1 .01 Calcanea 2 .01 Scapulae 2 .01 Podials 22 .14 Pelvis 1 .01 Phalanges* 42 .27 Humeri 2 .01 TOTAL 156 * Including metapodials of suid and hippopotamus. phalanges. These two groups of skeletal parts have very different properties of density and destructibility. The deltaic environments preserve more of the easily transported and destructible elements, the vertebrae and phalanges. The channel and floodplain environments presei"ve more of the denser and durable parts, primarily teeth. The experimental data on bone transport discussed in the section on bones as sedimentary particles can be used to interpret these differences in the fossil assemblages. Comparisons with Voorhies Groups The Voorhies Groups consist of bones with very different dispersal potentials. For animals from suid- to equid-size, phalanges and vertebrae are included in Group I, limb parts in Group II to III and teeth in Group II-III. Group I is most easily transported, Group III least easily transported and Group II intermediate, in currents up to 150 cm/sec, given Voorhies' ( 1969 ) experi- mental conditions. The three factors shown in Figure 21 are closely comparable to the three Voorhies Groups. Group I is more typical of the deltaic assemblages and Group II of the channel and floodplain assemblages. This provides evidence that transport sorting may be an important process in creating differences between the bone assemblages, i.e., the lag group is left behind in the channels while the transportable group is carried out to the deltaic and lacustrine deposits. The loss of Group I in the flood- plain may result from winnowing of the lighter elements during floods, if the cur- rent velocities on the floodplain exceed 10-20(?) cm/sec. Single Skeleton Comparisons Comparisons of the bone frequency data with the percentages of different bones in a single, whole skeleton show how the as- semblages have been altered from their original states. If all bones had been pre- served together, then the correlations be- tween the proportions of different parts in the sample assemblages and a single skele- ton should be high. Average proportions of parts in a single skeleton were calculated, combining the most common mammal groups in the fossil assemblages. These consist of bovids, hippos, suids and equids. Frequencies of the different parts are given in Table 8. Figure 22 shows the comparison of fossil and single skeleton bone frequencies for East Rudolf Paleoecolocy • Bchrcnameyer 537 SQUARE FREQUENCY no ^ a> o o o 00 o .'"^ ..'V .-^^""^ ■f'/^ ■ '/ / >4 4 . 'v i\ •■ -TOOTH — VERTEBRA- — SCAPULA •-RADIUS/ULNA -PHALANX' SQUARE FREQUENCY ro o 4:^ o C7> O cx> O \^ ^ * a \ ^ / m \ /' 1 t y\ 1 / \ / y ^ y \,-^^ y ^ -^ ^^^ -^ X ^< ^^^^ / ^ rf^ r 1 ■/ i 1 / ■ 1 / •. 1 L i» i? i ^^ V ■ \ \ ^^^ ^V- \ o Vll » 4 FLOODPLAIN 00 + O Sl CHANNEL o _.. < _.. CL (S 3 » T U3 0> — ' C 00 -■• o ^ CL -h (B cr (B 3- O <-!• -•• < o •a -1. 3 •a Q. o • DELTAIC LOCALITIES o I oj I tn I o ' o o 00 Figure 22. The square frequencies of the five most common mammalian skeletal elements in each locality compared with the proportions of the same elements in a single, average skeleton. The localities separated by factor analysis (Fig. 21) are distinct in their degree of alteration from single skeleton proportions. the 5 most common or most consistently occurring parts. The assemblages fall into two obvious groups: 103-0256, 130-0201, 10.5-0208 and 103-0267 are closely cor- related with the single skeleton, and 105- 1311, 102-0201 and 8+6-0104 are not. It appears that the lacustrine-deltaic environments, plus the 103-0267 channel- beach complex, preserve skeletal parts with a minimum of change from the original proportions. This implies the absence of processes that would sort the bones accord- ing to size, density or destructibility. In contrast, the channels, 102-0201 and 105- 1311, and the floodplain, 8+6-0104, pre- serve altered assemblages with a high pro- portion of the heavier and more durable parts and a much lower proportion of the lighter and more destructible elements. Discussion of Evidence for Transport Sorting The combination of evidence from the comparisons of bone assemblages with Voorhies Groups and single skeletons leads to important conclusions regarding the histories of the bones in each locality. In the deltaic deposits, bones from all Voorhies Groups are present in proportions similar to those of an average single skeleton. Therefore, the major component of Group I in these deposits is probably not trans- ported from elsewhere (i.e., the channels). If it were, then it has combined with lag assemblages to closely approximate the proportions in one skeleton. A better inter- pretation for the deltaic assemblages is that they have not been sorted. The relatively 538 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 fresh, unabraded surface textures of many of the bones, plus their lack of hydraulic equivalence with matrix grain sizes, further supports this interpretation. The high pro- portion of Voorhies Group I in the deltaic assemblages is a product of nonselective taphonomic processes rather than selective ones. In the 105-1311 and 102-0201 channels, the concentration of teeth is the result of sorting by fluvial processes. This sorting combines the lag concentration of teeth because of their greater density and because of their greater durability in transport situations. In addition, teeth are probably concentrated from floodplain deposits as the channel migrates laterally, eroding its banks. Other parts derived from re- excavated skeletons w^ould not be likely to survive erosion unless already mineralized. The 103-0267 distributary-beach complex combines the sedimentary characteristics of the other channels with a bone assemblage similar to the deltaic ones. The assemblage shows lack of selective sorting, and ap- parently a large lag component of teeth was not a product of the fluvial processes operating in 103-0267. Why this should be so is as yet unexplained. The floodplain assemblage shows selec- tive preservation of teeth in a fine-grained sedimentary context. As mentioned on p. 536, this indicates the removal of lighter elements from an untransported thana- tocoenose. Such removal could result from winnowing out of the light parts or from surface weathering and preferential de- struction of vertebrae and phalanges relative to teeth. The relative importance of these two processes can be determined by future experimental work on the critical entrainment velocities of vertebrae and phalanges, and by observation of thana- tocoenoses on modern floodplains. The low frequency of horn cores in 8+6-0104 may provide a clue indicating selective destruc- tion by weathering, since in modern situ- ations horn cores are often destroyed by womis that feed on their organic constit- uents (H. B. S. Cooke, personal communi- cation) (Plate 3). The extent to which sedimentary pro- cesses have altered the bone assemblages is clear from the examination of bone fre- quencies. The deltaic assemblages are least altered, and probably represent autochtho- nous accumulations in sedimentary environ- ments where the potential for rapid burial, without re-excavation, is high. The flood- plain assemblage is also autochthonous but has been altered by taphonomical processes so that it resembles the channel assem- blages. These show the most extensive alteration of bone ratios due to sedimentary processes. 103-0267 is intermediate in the degree to which taphonomic processes have affected the bone assemblage. The most useful localities for paleo- ecologic information are thus established as the deltaic and floodplain environments. The channels will also prove useful, since the factors contributing to their bone as- semblages are known. They will include a mixture of animals from the vicinity of a fluvial system, in contrast to deltaic de- posits, which preserve animals that fre- quented lake margin habitats. Additional Aspects of the Bone Assemblages More infomiation regarding taphonomic history can be drawn from bone character- istics unrelated to relative abundance. These include the occurrences of associated skeletal parts and the ratios of proximal and distal ends of limb bones. Associated parts of skeletons are rare in the East Rudolf deposits in general. The frequencies of these in the sample localities are included in Table 7. The channels 105- 1311 and 102-0201 have none, while the floodplain and deltaic localities, including 103-0267, have at least one. Most of these consist of associated vertebrae, with more complete partial skeletons occurring in 105- 0208 and 8+6-0104. The associated skeletal parts may result East Rudolf Paleoecology • Behrensmeyer 538 Table 9. Totals of proximal (P) and distal (D) limb ends IN THE fossil ASSEMBLAGES FROM EACH SAMPLE LOCALITY. Ut -lA — CHA NNEL- FLOOD- PLAIN 130- 0201 105- 0208 103- 0267 103- 0256 102- 0201 105- 1311 8+6- 0104 TOTAL Humerus P 2 2 2 0 3 2 2 13 D 3 8 1 3 2 2 7 26 Radius/ ulna P D 1 1 n 1 3 2 1 1 2 1 1 1 10 4 29 11 Femur P 1 4 1 0 2 1 3 12 D 4 5 1 2 5 3 2 22 Tibia P 1 4 1 1 1 5 5 18 D 2 1 2 3 4 3 6 21 Metapodial P 4 1 4 5 0 3 6 23 D 1 6 6 2 1 2 8 26 Total P 9 22 11 7 8 12 26 95 D 11 21 12 11 13 11 27 106 from carcasses buried at the site of death or from carcasses transported by flotation. There are very few criteria that could be used to distinguish between these two possible taphonomic histories. However, the associated parts do indicate a minimum of reworking of the bone assemblages after initial burial. This agrees well with other evidence for lack of reworking of the delta margin and floodplain assemblages. The absence of associated parts in the channels is consistent with the abraded surface tex- tures of the bones as an indication for ex- tensive reworking of sedimentary particles in the channel environments. Most of the limb bones in the samples are represented by one end or the other. It is of interest to determine whether some ends are more common than others, as an indication of preferential sorting or de- struction prior to burial. The numbers of proximal and distal ends of the major limb bones are listed in Table 9. For all localities and all limbs combined, the totals of 95 proximal and 106 distal are very close to a 1:1 ratio. This might be interpreted as indicating that no more proximal than distal ends are preserved, or vice versa. When the totals for each limb in all localities are examined, however, the relative frequencies of proximal and distal ends prove to be quite variable. There are nearly twice as many distal as proximal ends of humeri and femora, and many more proximal ends of radii/ulnae than distal. From the density measures of proximal and distal recent bones given in Appendix 2, it is apparent that the denser end is more commonly preserved in humeri and radii/ulnae, while the lighter end is more common in femora. A model for differential preservation because of transport sorting or more rapid weathering of low density ends does not fit this evidence. The differences in frequency of the proximal and distal ends can best be ex- plained by carnivore activity. In animals killed by carnivores or scavenged after death, the limbs are usually pulled off the carcass at the proximal articulation ( humerus/scapula and femur/pelvis joints) (Muller, 1957:256-258). Proximal ends of the humerus and femur would be subjected to stress and later exposed for gnawing. In contrast, the elbow and knee joints are more likely to remain held together by 540 BuUetiu Museum of Comparative Zoology, Vol. 146, No. 10 ligaments and survive the l)one-cnisliing activity directed at more nourisliing parts such as marrow-filled diaphyses. This ex- plains the relatively high proportion of distal humeri and femora. The dispropor- tionate number of proximal "radii/ulnae" actually consist primarily of olecranon processes from the ulnae. These are liga- ment-covered, lack a marrow cavity, and thus would survive better than the distal ends of radii. The pattern of proximal and distal limb element frequencies can be regarded as good evidence for carnivore activity in fos- sil assemblages in general. Such evidence has also been used by Voorhies (1969:20) to indicate carnivore activity prior to the final burial of the Pliocene Verdigre Quarry bone assemblage. For the East Rudolf localities, the evidence for carnivore ac- tivity can be detected in spite of differences in the taphonomic histories of the bone assemblages in the different depositional environments. The Reptilian Assemblages Reptilian parts form less consistent fre- quency patterns than those of mammals. The most common elements, as shown in Table 7 are crocodilian teeth and chelonian shell parts. The relative numbers of crocodil- ian parts are very similar from locality to locality. There is no indication of any in- crease in similarity between assemblages from similar sedimentary environments. This is probably a result of the universal availability of crocodile bones in the aquatic (generally depositional) environ- ments where crocodiles live. The low fre- quency of crocodiles in the floodplain environment, which crocodiles do not usu- ally frequent, emphasizes this point. The chelonian shell parts are variable in oc- currence, and are slightly more abundant in the deltaic deposits including 103-0267. This suggests some correlation between the more aquatic sedimentary environments and the chelonian occurrences. Conclusions Concerning the Bone Assemblages The evidence given in the preceding two major sections of this study brings out a number of taphonomically important fac- tors that can be combined to support a definite history for any given vertebrate fossil assemblage comparable to those oc- curring in the Koobi Fora Fm.: 1 ) The correlation of bone assemblages with dispersal groups from Voorhies' flume study 2) The correlation of bone assemblages with the proportions of a single skele- ton 3) The comparison of hydraulic equiva- lents of bones with grain sizes in the associated sediments 4) The completeness of bones, and sur- face characteristics that indicate pres- ence or absence of weathering or abrasion prior to burial 5) Presence or absence of articulated or associated skeletal parts 6) Ratios of proximal and distal ends of limb bones that deviate from 1:1 All of these factors provide a basis for interpreting the East Rudolf data. The bone frequencies for each locality thus can be used to determine the taphonomic histories of the vertebrate assemblages. The following points can be made: 1) The different sedimentary environ- ments of East Rudolf show a general similarity in the compositions of their mammalian bone assemblages. The same bones are present in all environ- ments, and none of the assemblages consist exclusively of one of the three Voorhies Dispersal Groups. 2) Evidence for a certain degree of sorting and redistribution of bones is present in the different sedimentary environments. Significant differences in relative numbers of different bones are shown by the concentrations of teeth in the channels (105-1311 and East Rudolf Paleoecology • Behrcnsmcyer 541 102-0201) plus the floodplain (8+6- 0104), and by the concentrations of \'ertebrae and phalanges in the deltaic localities ( 130-020l/ 105-0208, 103- 0256, 103-0267 ) . These can be corre- lated with the sorting effects of tapho- nomic processes in the channels and on the floodplain, and the absence of sorting on the delta margins. 3) Consideration of the bone frecjuencies in the light of Voorhies Groups, hydraulic equivalence and single- skeleton comparisons shows that au- tochthonous and allochthonous assem- blages of fragmental vertebrate bones can be distinguished. The deltaic and floodplain localities consist of basic- ally autochthonous vertebrate fossils, while the channels contain a mixture of allochthonous and autochthonous assemblages. FAUNAL ASSEMBLAGES OF THE KOOBI FORA FORMATION Taphonomic analysis has shown that all of the sample fossil assemblages can be considered autochthonous in the broadly defined deltaic and fluvial environments. It is now possible to examine the faunal compositions of the seven bone assemblages and to relate these to the different sedi- mentaiy environments. Comparisons can be made from environment to environment which should indicate ti'ue paleoecologic differences or similarities in the faunas. In the following discussion, several aspects of the paleoecology of the Koobi Fora Fm. and its vertebrates will be given particular attention. These include the differences in numbers of aquatic and nonacjuatic x'ertebrates, the relatixe frequency of dif- ferent terrestrial mammals in the different environments, and the patterns of occur- rence of mammalian groups that havc^ close counterparts in modern ecosystems. The fauna from the square sample as a whole includes 14 out of the 20 major vertebrate groups listed by Maglio (1972: 380-381) for the Koobi Fora Fm. The sample assemblages also include most of the genera of bovids, suids, equids and hippos. The carnivores listed by Maglio (1972:380-381) are the most poorly repre- sented groups in the samples used for this study. Method of Identification The fossil collections consist of material that can be identified at a number of dif- ferent taxonomic levels. Major groupings of vertebrates used for faimal comparisons among the sample localities were desig- nated so that each member of a group has approximately equal numbers of identifi- able parts. In practice, for example, this amounted to teeth, skull parts, limb ends and foot parts for mammals. The mammals listed below could be identified e(}ually well using any of these parts. Consideration of this factor was necessary to prevent undue biasing of the square frequency for a form with substantially more or less identifiable parts. The fossil assemblages can be divided into faunal groups, cor- responding roughly to several taxonomic Categories, as follows: 1) Class: Mammal, Reptile, Bird, Fish. Identifications were based primarily on the morphology of the bone fragments. Bone micro-structure was useful as a distinguishing character for very small fragments. In some cases parts of pelves, scapulae, ribs and diaphyses could not be certainly assigned either to mammal or reptile, and such parts are not in- cluded in any of the totals. 2) Groups of Reptiles and Mammals. Mammals Reptiles Elephant Suid Crocodyliis Deinothere Eciuid Etitliccodou Hippopotamus Primate Trionychid Rhinoceros Carnixore Peloni• o UJ o .60 LU on u. UJ or ^ .40 a CO .20 East Rudolf P.\leoecology • Behrensmcyer 549 useful measure of the eff(X'ts of taphonomic processes (weathering and transport) on the fossil assemblages. Figure 24 shows the frequencies of the four families in terms of all identifiable elements and in terms of teeth only. Where the lines divc>rge, a large proportion of the sample consists of parts other than teeth. Representation of hippos and bovids is similar in all localities except 103-02.56 and .20 LU .10 Mesochoerus y \ Metridiochoerus/ Notochoerus J- 130- 0201 105- 0208 102- 0201 105- 1311 DELTA CHANNEL 8+6 0104 ♦ FLOODPLAIN Figure 25. A comparison of tlie square frequencies of the two suid groups, Mesochoerus and Notochoerus/ Metridiochoerus. Mesochoerus is considered here to be a more closed habitat (bush) form due to its relation- ship to the modern Hylochoerus (Giant Forest Hog), and to its low-crowned molars, which appear to be adapted for relatively soft vegetation. Notochoerus/ Metridiochoerus suids are more closely related to the modern Phac- ochoerus (Warthog) and have high-crowned molars adapted for abrasive vegetation. These suids may have been more open habitat (grassland) forms. Localities 103-0256 and 103-0267 are omitted due to the low fre- quencies of suids identifiable to genus (Table 12). East Rudolf Paleoecology • Behrensmeyer 551 Table 13. Generalized ecological characteristics of Recent bovid groups that are commox in THE East Rudolf fossil assemblages. ( Modified from R. Estes, ms. in press. ) Water projcimity IIal)itat Near Far Food h.tbits Social habits Tiagelaphini Dense Bush X (X) Browsers small groups Reduncinae Woodlands, Floodplains X Grazers small groups Alcelaphinae Open grasslands (X) X Grazers large herds sistent with predictions based on tooth morpliology and recent analogues. The channel assemblages might be expected to sample the more open-country habitats, particularly if the gallery forests fring- ing the channels are not very extensive. Deltaic environments, if comparable to the most vegetated areas of the recent Omo Delta, would have more forested habitats. The paleoecologic evidence associ- ates Mesochoerus with deltaic, potentially more densely vegetated environments and Notochoerus/Metrkliochoerns with fluvial, mixed- to open-habitat environments. The third molars of the two suid groups are different in size, and there is a possi- bilitv that the smaller Mesochoerus teeth have been sorted out of the channel de- posits. Mesochoerus third molars are between about 40 and 60 mm long and 20 mm in height, while Notochoenis/MetrkUo- choerus third molars are from about 50 to 75 mm in length and 40 to 60 mm in height. However, the hydraulic equivalents of the teeth in both channels fall within the 10-25 mm range, which is near the median for the total range of other teeth in the deposits as well as the associated sediment. One cannot logically assume that sorting would sepa- rate the pig teeth but nothing else with similar size differences. Therefore, sorting can be eliminated, and ecological factors provide the best explanation for the sepa- ration of the two suids in the fossil assem- blages. EQuros Although the equid sample is poor and consists mainly of teeth (Figure 24) there is some suggestion of habitat separation of Eqtms and Hipparion in the samples. Eqitus is most abundant in 105-1311 and 102-0201, and Hipparion in 130-0201 and 105-0208. There is a time separation be- tween these two groups of samples, but Eqtms is known to occur elsewhere in East Rudolf at the same level as Hipparion. The correlation of Eqtms with the channel envi- ronments and Hipparion with lake margins is comparable to the pattern of occurrences of N otochoertis /Metridiochoerus and Meso- choerus. Hipparion is preserved in associ- ation with the environment most likely to have been densely vegetated, and Eqims is found in the deposits more likely to have sampled open country, savanna forms. BoviDS Three bovid groups are abundant enough in the sample assemblages for detailed analysis. These include the Alcelaphinae (hartebeest, etc.), Tragelaphini (kudus, elands, etc.) and Reduncinae (bush buck, waterbuck, etc. ) . Recent members of these groups are well known in terms of habitat preference (Bigalkc, 1972; Estes, in press; Dorst and Dandelot, 1970). Ecological characters are listed in Table 13. Fre- quencies of bovid tribes in the fossil as- semblages are given in Table 14. All localities combined, alcelaphines and reduncines are nearly equal in abundance, while Tragelaphines are less common. The high frecjuencics of the smaller alcelaphines in 103-0256 and of both large and small in 105-1311 are significantly larger than the frequencies of the other groups in these localities, with p ^ .05 (Chi-square tests). 552 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 Table 14. Frequencies of fossil bovids and the small hippopotamus in the East Rudolf assemblages. Delta Channel Floodplain 130-0201 105-02 08 103-0267 103-0256 10; J-0201 105 -1311 8+6-0104 Tragelaphini .24 .10 .20 .00 .12 .16 .02 Reduncinae .24 .25 .15 .11 .15 .12 .15 Alcelaphinae ( Damaliscus-size ) .14 .15 .05 .33 .06 .40 .12 Alcelaphinae ( Megalotragus ) .00 .00 .15 .04 .00 .48 .03 Hippopotamus .05 .20 .10 .26 .09 .00 .06 sp. nov. Reduncines and tragelaphines are of similar frequency except in 103-0256 and 8+6- 0104, which have a high proportion of reduncines and few tragelaphines. These differences are probably related to eco- logical factors since it is difficult to imagine any other processes which could preferentially sort the tribes. (They are of approximately equal body size.) All are represented by multiple skeletal parts where they are abundant. The patterns of occurrence are not as well defined as for the suids, but bush forms ( reduncines ) are generally more common in the deltaic environments while alcelaphines are as- sociated with the potentially more open environments sampled in the 105-1311 channel and in the 103-0256 mudflats. The alcelaphine Megalotragus is a large, extinct form known only from teeth in the sample assemblages. It has an unusually high frequency in the 105-1311 channel. This is significantly different from its oc- currences in the other localities and can- not be explained except by ecological fac- tors. Megalotragus is associated with the grassland suids and equid, and may well have been an open-country form itself. Its absence in 102-0201 is somewhat puzzling, if it is typically preserved in fluvial deposits. However, another large bovid (probably Pelorovis) is represented in 102-0201 with a frequency of .18. The data may indicate a rather finely resolved habitat separation between the two forms which can only be clarified by additional sampling. Hippos The habitat of the extinct small hippo, H. sp. nov., can be generally inferred from its frequency in the bone assemblages (Table 15). It occurs in all localities ex- cept 105-1311 and is most abundant in 103-0256. Nearly all the hippo remains in 103-0256 belong to this form, and a variety of skeletal parts exist in the sample squares together with teeth. It is generally more abundant in the deltaic environments, including 103-0267, and is associated with both bush and open country animals. It is definitely autochthonous in the deltaic mudflats environment of 103-0256. From this evidence, it can be concluded that H. sp. nov. was probably a lake margin fonn, preferring deltaic flats with mixed bush and grassland environments. It may have been less aquatic than the larger hippos, but this can only be validly inferred from morpho- logical data, not from the taphonomic evi- dence now available. A large extinct hippo, peculiar to the East Rudolf Plio-Pleistocene, cannot be definitely identified in the fossil assem- blages from the squares. Ecological data on this hippo would be interesting since somehow three or more forms of Hippo- potamus managed to coexist at East Rudolf. East Rudolf Paleoecology • Behrensmeyer 553 SEMI-AQUATIC TO AQUATIC (hippo, Crocodylus Euthecodon) TERRESTRIAL (bovid, suid, equid) PRIMARILY AQUATIC (Euthecodon. trionichid, pelomedusid) Figure 26. Triangle diagram showing the results of a three factor analysis of the frequency data from all groups of mammals and reptiles. The deltaic localities are spread between two "aquatic" factors, and the channel and floodplain localities are distributed closer to the "terrestrial" factor. At least one of these is not present in the entire Omo seqvience, suggesting significant ecological differences between the two regions, at least as far as the hippos were concerned. We might assume that these differences were expressed in the utilization of broadlv different environments. If so, further careful sampling of the sedimentary e\'idence should reveal more about the ecology of the different hippos. Conclusions Regarding the Faunal Assemblages Much of the information provided b\' the faunal frequencies is summarized in the triangular diagram in Figure 26. This shows the results of a CAi3FAC Q-Mod(> Factor analysis for three varimax axes (which explains 97'/' of the variance). The data consist of the square frequencies for all the animal groups given in Table 11. The three factors can be direct!)' r(>lated to the acjuatic or nonacjuatic affinities of the various animals. Factor 1 includes the terrestrial forms and the other two factors include aquatic and semiaciuatic forms with affinities for channel or deltaic-lacustrine habitats. These separate the sample^ locali- ties into three groups depending on their components of ac^uatic animals. Thus, the evidence at various taxonomic 554 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 levels indicates faunal differences between the localities which agree with environ- mental interpretations based on geological data. These are real and meaningful ecological differences between environ- ments; differences expressed in the square frequencies of the faunas and supported by the geologic and taphonomic characters of the sediments and their bone components. The important points brought out by the faunal data include: 1) The relative numbers of the different vertebrate classes in the fossil assem- blages is more dependent on their original numbers and proximity of habi- tat to a sedimentary environment, than on their aquatic or nonaquatic habits. 2 ) Relative numbers of animals can indi- cate whether they were aquatic or nonaquatic in nonaquatic sedimentary environments, but not in aquatic ones. 3) The more terrestrial sedimentary en- vironments preserve a greater diversity of terrestrial animals. However, the more aquatic environments may preserve an equivalent or greater number of bones from terrestrial animals, representing fewer groups. 4) For terrestrial families larger than a baboon or a small antelope, the frequen- cies expressed in the sample assemblages should be roughly proportional to their original numbers. 5 ) The more abundant mammals (bovids and hippos) are generally represented by more kinds of skeletal parts of different sorting potential, indicating autochtho- nous accumulations of bones. This is consistent with the generally autochtho- nous nature of the assemblages on the deltaic and floodplain environments. 6) The different sedimentary environ- ments clearly preserve different ratios of some animals with different habitat preferences. Terrestrial animals which prefer grassland habitats are found in greater abundance in fluvial deposits, while bush or forest mammals occur in greater abundance in the deltaic deposits. This indicates deltas with denser vege- tation than the gallery forests which fringed the channels while the sample bones were accumulating. PALEOECOLOGY OF THE VERTEBRATE ASSEMBLAGES OF THE KOOBI FORA FORMATION Much independent but cross-supporting evidence provides a basis for interpreting the paleoecology of the East Rudolf fossil assemblages. These lines of evidence in- clude: 1) Geologic evidence. Characteristics of the overall sedimentary environments and the processes operating within them (Fig. 27). 2) Taphonomic evidence, a) The extent and type of sorting in the bone assem- blages, interpreted with the aid of theoretical considerations from experi- mental evidence for bone disposal, b) Relationships of the hydraulic equiva- lences of the bones and of the associated matrix sediment, c) Characteristics of the bone fragments, interpreted accord- ing to observations on weathering, frac- turing and abrasion of modern bones. 3) Faunal evidence. Interpretations based on the faunal composition of the fossil assemblages, using the ecology of modern analogues to Plio-Pleistocene animals. Figure 27. Block diagrams showing reconstructions of East Rudolf sedimentary environments. Circles show the interpretation of the general sedimentary environment of each fossil sampling locality, indicated by locality numbers. The representation is schematic; the localities do not occur on the same time planes or closely adja- cent to each other as might be construed from the diagrams. 102-0201 is more closely associated with an emergent delta than can be shown on the diagram. 103-0267 includes a more extensive complex of distributary mouth and beach environments than is indicated by the encircled area. East Rudolf Paleoecology • Behrensmeyer 555 8+6-0101 FLUVIAL SYSTEM 103-0256 130-0201 105-0208 DELTA 556 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 1.00 .80 o o or S. s. CO .60 .40 .20" .00 Increase in Aquatic component I a- I ^ § I I Oa Aaaac 130-0201 103-0267 103-0256 Delta Margin Distributary and Beach Complex Deltaic Mudflats 8+6-0104 Floodplain Figure 28. A comparison of the square frequencies of four vertebrate groups with aquatic, semiaquatic and ter- restrial habits in four different depositional environments. These environments range from primarily aquatic to primarily terrestrial in terms of their geologic characteristics. The frequencies of the aquatic and semiaquatic animals increase as the depositional environments become more aquatic (floodplain to delta margin). However, the frequencies of terrestrial forms remain essentially constant in all environments. East Rudolf Paleoecology • Bchrensmeyer 557 Other lilies ot exidence can be important in paleoecologie interpretations but are not at present available for the East Rudolf assemblages. These include botanical and gcochemical data, which can reveal im- portant factors about vegetation, climate, and salinity of the lake. Continuing re- search should eventually provide such data. Ecological Comparisons of the Samples The overall similarities and differences among localities show that sedimentary en- \'ironments can be characterized according to distinct taphocoenoses and biocoenoses. At East Rudolf, three broadly defined sedimentary environments are represented: delta, channel and floodplain. The faunas are all basically autochthonous in each of these environments and reveal meaningful ecological differences among them. Aquatic and Terrestrial Faunas The more aquatic sedimentary environ- ments as detennined from geologic evi- dence have an increased rejjresentation of a({uatic animals but show no decrease in the absolute number of terrestrial animals. This is demonstrated in Figure 28 by the increase in the frequencies of crocodilians and hippos relative to bovids and equids. I The absolute frequency of bovids and eciuids does not change significantly from environment to environment, even though these range from floodplain to delta margin. The pattern of aquatic and terrestrial occurrences can be represented for faunas from each locality as shown in Figure 29. The ratio of terrestrial animals increases as environments become more terrestrial, at the expense of the acjuatic forms. The ratio of the semiaquatic hippo, which spends approximately half its time in and half out of the water (Dorst and Dandelot, 1970: 172), changes little from aquatic to non- aquatic environments. These patterns are the result of geologic and taphonomic processes which should have similar effects on fossil assemblages other than these East Rudolf examples. The crucial variables appear to be: 1) the total volume of bones available from acjuatic and nonaquatic animals and 2) the pro\imit\- of an animal's habitat to an actively aggrading sedimen- tary environment. The habits of a fossil vertebrate cannot be inferred from its abun- dance in an a(iuatic sedimentary environ- ment unless this can be compared with more terrestrial environments from about the same time. Open and Closed Habitat Mammalian Faunas The mammalian assemblages provide evidence for two terrestrial faunas with preferences for open (grassland) or closed (bush) habitats. In order to establish these ecological differences, the habitats of the fossil mammals must be inferred from morphologic evidence plus analogy to re- lated living forms. For the paleoecologie interpretation of the East Rudolf fossil assemblages, Mesochoerus, reduncines and tragelaphines are used to represent the closed habitat fauna, and DamaJiscus-size alcelaphines, Notochoems/MetridiocJwerus suids and Equus represent the open habitat fauna. The evidence for relating these mammals to the different ecologic situ- ations has been discussed previously. The ecological separation of such groups accord- ing to habitat preference for grassland or bush environments is a common feature of recent East African ecosvstems (Lamprey, 1963; Harris, 1970; Estes,' 1973). The relative percentages of closed and open habitat forms in the seven fossil localities are shown in Figure 30. All localities include both, but the deltaic environments in general include more closed habitat forms and the channels more open habitat forms. The deltaic mudflats (103-0256) have an open habitat fauna, in agreement with geologic e^'idenee for an extensive, unforested delta margin environ- ment. The patterns of faunal occurrence indicate that the deltaic, chaimel and flood- 558 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 % 10 o 80 u TERRESTRIAL VERTEBRATES Environments Increasingly Terrestrial ^ Figure 29. The ratios of aquatic, semiaquatic and terrestrial vertebrate groups represented in the East Rudolf fossil assemblages. The sample localities are arranged from the more aquatic to least aquatic depositional en- ^'ironments on the basis of geologic interpretations. Aquatic animals include crocodilians and chelonians except Geochelone; semiaquatic includes only hippopotamus, and all other groups are considered to be terrestrial. Abundance is calculated as the % of the cumulative square frequencies for each locality. plain sedimentary environments sample both closed and open habitats, but that closed habitats were more abundant on the deltas. A comparison between 130-0201, representing a deltaic fauna, and 105-1311, representing a nondeltaic fauna, shows the most distinct ecological difference among any of the localities (Fig. 31). Comparisons of Koobi Fora Formation Faunas and Recent Terrestrial Faunas Bovids, suids and equids are the most abundant large mammals in the fossil as- semblages and also in most of the recent undisturbed East African ecosystems (e.g., Foster, 1967; Sheppe and Osborn, 1971). The ratios of these mammals in the fossil East Rudolf Paleoecology • Bchrcnsmcijcr 559 TOO ^ u o u 80 60 i- 1 *J S 30 cC^^" -^ o \ ^ X/ u combination of geologic, taphonomic and faunal evidence, 564 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 it will be possible to compare fossil verte- brate communities throughout East Africa and to reconstruct changes in chronofaunas through much of the latter part of the Cenozoic. SUMMARY This study has developed methods for deriving paleoecologic information from fossil assemblages of fragmented vertebrate bones subjected to various geologic pro- cesses before burial. These methods have been applied to paleoecologic interpreta- tion of the Plio-Pleistocene bone deposits of East Rudolf, Kenya. The conclusions relate to vertebrate assemblages in general as well as to the assemblages of East Rudolf and the Lake Rudolf Basin in particular. General Conclusions Taphonomy 1) The amount of fragmented bone buried in any given sedimentary environ- ment will depend on the rate of sedimenta- tion and the amount of bone originally put into that environment. The important fac- tors which control bone input are: a) verte- brate abundance, b) carnivore activity, c) the proximity of bones to depositional en- vironments, d ) the rates of surface weather- ing of bones, and e) the dispersal potential of bones. The composition of the resulting fossil assemblage will also in part depend on diagenetic factors. 2) Carnivore activity will have a major effect on the composition of a thanato- coenose. Intense mammalian carnivore ac- tivity results in fewer bones of small animals and increased fragmentation of bones of large animals. The evolution of bone-crushing dentitions in mammals has changed the character of Cenozoic tapho- coenoses compared with those of the Mesozoic, when reptilian carnivores lacked the capacity for bone mastication. 3) Bones are disarticulated and acquire characters of surface weathering in months to years if exposed on a land surface. Hydrodynamic transport will tend to leave features of rounding and abrasion on bones. Therefore, well-preserved bones with frag- ile parts intact and surfaces unflaked or uncracked record conditions of rapid burial without subsequent re-excavation. 4) Bones vary greatly in density, size and shape and are sensitive to hydro- dynamic sorting. Disarticulated thanato- coenoses include bones with a wide range of dispersal potentials. This will result in the formation of dispersal groups if the bones are subjected to normal or flood- stage current velocities ( 10-150+ cm/sec. ) . The dispersal groups will move at different rates from the point of origin. If bones with a wide range of dispersal potentials are found in sedimentary^ association, this indi- cates that the assemblage is not a product of selective transport sorting of the original thanatocoenose. 5) Mammal bones immersed in water for 5 minutes have densities from < 1.0 to 2.0, and teeth have densities between 1.7 and 2.3. Reptile and fish bones are be- tween 1.3 to 2.3 density. Bones are gen- erally hydraulically equivalent to quartz particles of smaller nominal diameter. Cur- rents should transport bones together with quartz particles that are roughly equivalent hydraulically. Therefore, sedimentary as- sociations of quartz grains and bones of a much larger hydraulic equivalence (e.g., a hippopotamus skull in a siltstone) may indi- cate other modes of origin for the bone- sediment association. These include in situ death, flotation of carcasses, or predator/ scavenger transport of bone. 6) Theoretical considerations indicate that velocities of 80 to 200+ cm/sec. must be achieved near the bottom of a flow in order to move bones of moderate density (~ 1.5) and size (100+ cc). Therefore, most disarticulated, water-logged parts of large vertebrates are unlikely to move far from their point of origin except in special trans- port situations such as floods in channels. East Rudolf Paleoecology • Behrensmcycr 565 Paleoecology i) Ecological characteristics ol fos- sil vertebrates can be dc^fined using a combination of geologic and taphononiic evidence, independent of ecological inter- pretations based on \'ertebrate morphology or the adaptation of li\'ing analogues. Such evidence can link habitat preferences with preservation in particular sedimentary en- vironments. This correlation can be inferred solely from the geologic context and the taphonomy of a gi\'en bone assemblage. Such e\idence can then be combined with morphological and recent-counterpart data to support paleoecologic interpretations. 2 ) Fragmented bone assemblages can be used with confidence for paleoecologic interpretations if they: a) consist of bones with a wide range of dispersal potentials, b) are not hydraulically equivalent to associated sediment and c) retain fresh, unweathered or unabraded surfaces. As- semblages with these attributes can be interpreted as generally autochthonous to their environment of deposition. Most of the animals represented in such an assemblage were preserved in the general context of their original habitats. 3) Aquatic environments of deposition can preserve variable amounts of bone from aquatic and terrestrial vertebrates, de- pending on the relative bone input from each ecological group. Bone assemblages of terrestrial and aquatic animals in aquatic deposits (e.g., channel, delta margin) may differ only in the better preservation of the latter, not in their greater abundance. 4) Terrestrial environments of deposition (e.g., floodplains) preserve a high pro- portion of terrestrial vertebrates along with a few aquatic ones. Semiaquatic vertebrates tend to occur in both terrestrial and acjuatic deposits, with better representation in aquatic environments. 5) The bone input from groups of large terrestrial vertebrates into fragmented, autochthonous taphocoenoses should gen- erally reflect their relative numbers in the original ecosystem. The fossil abundances can be used to approximate relative num- bers of different \ertebrate groups in a gixen en\'ironment. This provides a basis for reconstructing paleo-communities and comparing them through time. 6) Vertebrate communities at different time horizons or in different regions may differ in their response to broad-scale environmental change in ways that can be detected in paleoecologic studies. These responses include rapid morphological evo- lution, shifts in the relative numbers of animals suited to particular habitats, and a general decline in species diversity ac- companied by the extinction of forms in all the available habitats. Conclusions for the Vertebrate Assemblages of tfie Koobi Fora Formation, East Rudolf 1) Fossil-bearing deposits reveal sedi- mentation and bone preservation in at least three major depositional environmc^nts: delta margin, channel and floodplain. 2) The three depositional environments show a basic similarity in their representa- tion of different skeletal parts, with teeth the most abundant component. However, the relative numbers of certain skeletal elements differ in ways that reflect the dif- ferent processes operating in the three environments. Teeth are relatively more abundant in the channel deposits and in the floodplain, while vertebrae and phalanges are more abundant in the delta margin deposits. This can be related to the concentration of heavy, durable parts in the channels through sorting and re- working of bones, and to the absence of such processes in the delta margin environ- ments. The taphononiic characters of the floodplain assemblage indicate preferential removal of the lighter elements without transport or reworking of thc> associated heavier l:)()nes. 3) The sum of taphonomic and geologic evidence shows that the delta margin and floodplain bone assemblages are autoch- thonous with respect to the overall sedi- 566 Bulletin Museum of Comparative Zoologtj, Vol. 146, No. 10 mentary environment. Channels contain a mixture of allochthonous and autochtho- nous bones and show the most evidence for taphonomic alteration of the original than- atocoenose. 4) The East Rudolf faunas include aquatic, semiaquatic and terrestrial verte- brates that vary in abundance according to sedimentary environment. Analogies with recent East African ecosystems indicate that the relative fossil abundance of ter- restrial mammalian families probably re- flects their abundance in the original ecosystem. Bovids, suids and equids are the most common groups in the fossil as- semblages and in most recent undisturbed East African faunas. 5) Two terrestrial faunas can be defined for the East Rudolf assemblages, based on ecological analogies between recent and fos- sil mammals. The open habitat fauna in- cludes alcelaphines, Metridiochoerus/Noto- choerus suids and Eqiius. The closed habitat fauna is characterized by Meso- choerus, reduncines and tragelaphines. There is overlap of these faunas in all of the sample assemblages. However, delta margin deposits generally preserve a greater proportion of closed habitat forms, and the channels preserve more open habitat forms. 7) The paleoecologic results for East Rudolf show that it is possible to define ecological groups of terrestrial vertebrates from surface samples of fragmental bone assemblages. Similar sampling of fossil assemblages at different time horizons can provide a basis for establishing East African chronofaunas and for reconstructing their interaction with environmental changes through time, ACKNOWLEDGMENTS The interdisciplinary nature of this work led to a great deal of productive and en- joyable interaction with researchers in a broad range of disciplines, including Geol- ogy, Paleontology, Zoology, Anthropology, and Ecology. An interdisciplinary approach inevitably generates a large number of well-deserved acknowledgments. I attempt here to express my gratitude to those people who have made particular contributions to this study, but I wish to preface this with a simple and very sincere note of thanks to all who provided help and encouragement during the course of the project. I would like to give special thanks to Bryan Patterson, Richard E. Leakey, Glynn L. Isaac and Vincent J. MagHo. Without their unfailing encouragement, help and ideas, this work would not have been pos- sible. Gratitude is due to Parish A. Jenkins, Jr., Raymond Siever, Robert T. Bakker, Daniel C. Fisher, Peter Dodson and H. B. S. Cooke for many useful com- ments and suggestions during the prepa- ration of the manuscript. Discussions with Andrew Hill, Richard D. Estes, Stephen J. Gould, Jack Sepkoski, F. B. Van Houten, G. Jepsen, John Fleagle, Allen Greer, T. Hopson and Stanley Awramik resulted in many additional ideas and references. Dirk van Damme (Geologish Institute, Ghent, Belgium) also deserves thanks for identifi- cation of the invertebrate fossils. I have greatly appreciated the exchange of geo- logical information with co-workers on the East Rudolf Expedition, including Bruce Bowen and Carl Vondra of Iowa State Uni- versity, Gary Johnson of Dartmouth College and Ian Findlater of Birkbeck College, University of London. Field work ( consist- ing of long hours of sample collecting in 10 X 10 meter squares) was accompHshed through the assistance and stoicism of Susan Abell, Penny Bowen, John Bart- helme, John W. Harris, John M. Harris, John Onyango-Abuje, Jonathan Karoma, John Kimingitch, Fred Lucas, Dinah Grader, Diane Gifford, Dan Stiles, Paul Abell, Kelly Stewart and Andrew Hill. John Barthelme and John Harris deserve special thanks for their help in laboratory analysis and identification of the fossil material. Barbara Lawrence and Charles Mack of the Department of Mammalogy, Museum of Comparative Zoology ( Harvard ) were very I East Rudolf Paleoecology • Bchrvnsmexjer 567 h(>lptul in providing recent skeletal material used for this stud>-. Photographic reproduc- tion of the figures and plates was done by John Lupo (Biological Laboratories, Har- \'ard) and additional assistance was pro- vided by Al Coleman (MCZ Laboratories, Harvard). Typing was the joint effort of Karen Mason, Maureen Sepkoski and Agnes Martin. Their work is gratefully acknowl- edged with special thanks to Agnes Martin for taping the final draft. Finally, among the many friends who have provided mis- cellaneous assistance in time of need, I would particularly like to thank Vickie Rowntree, Catherine Badgley, A. Gordon Brown and Elizabeth Whitehouse. This i;tudy was done as a Ph.D. Disser- tation in the Department of Geological Sciences, Harvard University, and was completed in June of 1973. The work re- ceived financial support through grants by the National Science Foundation (Grant No. 28607a) and the National Geographic Society to the East Rudolf Research Project. REFERENCES AuEL, O. 1912. Grinidzuge der Palaeobiologie der Wirbeltiere. Stuttgart- E. Schweizer- bart'sche Verlagsbuchhandliing Niigele und Dr. Sproesser. 470 pp. Allen, J. R. L. 1963. The classification of cross-stratified units, with notes on their origin. Sedimentol., 2: 93-114. . 1965. A review of the origin and characteristics of recent alluvial sediments. Sedimentol., 5: 89-191. . 1970. Physical Processes of Sedimenta- tion. London: George Allen and Unwin, Ltd. 248 pp. Allen, P. 1959. The Wealden environment: Anglo-Paris Basin. Roy. Soc. London, Phil Trans. (B), 242(692): 283-346. Ansell, W. H. F. 1971. Part 15: Order Artio- dactyla. In: Meester, J. and 11. \V. Setzer, (eds.). The Mammals of Africa. An Identifi- cation Manual. Washington, D.C: Smith- sonian Institution Press. Aristarain, L. F. 1962. Caliche deposits of New Mexico. Ph.D. Dissertation. Harvard Universit>-, Department of Geological Sci- ences. Baker, B. H., and J. Wohlenberg. 1971. Struc- ture and evolution of the Kenya Rift Valley. Nature, 229: .538-542. Beadle, 1^. C. 1932. The waters of some East .\frican lakes in relation to their fauna anci flora. J. Linn. Soc. (Zool.), 38: 135-136. BEimKNSMEYEH, A. K. 1974. Late Cenozoic sedimentation in the Lake Rudolf Basin, Kenya. Annals Geol. Soc. Egypt, IV: 287-306. . In press. The habitat of Plio-Pleistocene hominids in East Africa; taphonomic and mi- cro.stratigraphic evidence. In C. Joly (ed. ), African Hominidae of the Plio-Pleistocene: Evidence, Prolilems and Strategies. New York: Duckworth, Inc. Berggren, W. a., and J. Van Couvering. 1973. Late Neogene chronostratigraphy, biostratig- raphy, biochronology and paleoclimatology. Woods Hole Oceonogr. Inst. Tech. Rep., WHOI-73-40. 334 pp. Berry, L. G., and B. Mason. 1959. Mineralogy. San Francisco: W. H. Freeman and Co. 630 pp. Bigalke, F. C. 1972. The contemporary mam- malian fauna of Africa. In Keast, A., B. Glass, F. C. Erk, (eds.). Evolution, Mammals and Southern Continents. Albany, New York: State University of New York Press. 141-194. Bishop, W. W. 1968. The evolution of fossil environments in East Africa. Trans. Leicester Lit. and Phil Soc, 62: 22-44. BowEN, B. E., and C. F. Vondra. 1973. Strati- graphical relationships of the Plio-Pleistocene deposits. East Rudolf, Kenya. Nature, 242: 391-393. Brain, C. K. 1967a. Bone weatliering and the problem of bone pseudo-tools. S. Afr. J. Sci., 63(3): 97-99. . 1967b. Hottentot food remains and their bearing on the interpretation of fossil bone assemblages. Sci. Pap. Namib Des. Res. Sta., No. 32. Bretz, J. H., AND L. HoRBERG. 1949. Caliche in southeastern New Mexico., J. Geol. 57: 491-511. BrIGGS, L. I., D. S. McCULLOCH, AND F. MOSER. 1962. The hydraulic shape of sand particles. J. Sediment. Pet., 32(4): 645-656. Brock, A., and G. Isa.a.c. 1974. Paleomagnetic stratigraphy and chronology of hominid bear- ing sediments east of Lake Rudolf, Ken\a. Nature, 247: 344-348. BuTZER, K. W. 1971a. Recent History of an Ethiopian Delta. Res. Pap. No. 136, Dept. Geogr. Chicago: The Uni\ersit\- of Chicago Press. 184 pp. . 1971b. The Lower Onio Basin- geol- ogy, fauna and hominids of Plio-Pleistocene formations. Naturwiss., 58: 7—16. BuTZER, K. W., G. L. Isaac, J. L. Ric:iiardson, 568 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 AXD C. Washboukn-Kamau. 1972. Radio- carbon dating of East African lake levels. Science, 175(4027): 1069-1076. Clark, J., J. R. Beerbow^r, and K. K. Kietzke. 1967. Oligocene sedimentation, stratigraphy and paleoecology and paleoclimatology in the Big Badlands of South Dakota. Fieldiana, Geol., 5. 158 pp. Cooke, H. B. S., and V. J. Maglio, 1972. Plio- Pleistocene stratigraphy in East Africa in relation to proboscidean and suid evolution. In Bishop, W., and J. Miller, (eds.). Cali- bration of Hominoid Evolution. New York: Scottish Academic Press. 303-329. CoTT, H. B. 1961. Scientific results of an in- quiry into the ecology and economic status of the Nile Crocodile (Crocodylus niloticiis) in Uganda and Northern Rhodesia. Trans. Zool. Soc. Lond., 29(4): 211-356. CuRKEY, J. 1970. Animal Skeletons. London: Edward Arnold, Publishers. Davies, D. K., F. G. Ethridge, and R. R. Berg. 1971. Recognition of barrier environments. Bull. Am. Assoc. Pet. Geol., 55(4): 550-565. Dodson, P. 1971. Sedimentology and taphonomy of the Oldnian Formation (Campanian), Dinosaur Provincial Park, Alberta (Canada). Paleogeogr., Paleoclimatol., Paleoecol. 10: 21-74. . 1974. The significance of small bones in paleoecological interpretation. Contrib. to Geol., Univ. of Wyoming. Spec. Pap. No. 2 Laramie, Wyoming: Univ. Wyoming Press. DoRST, J., AND P. Dandelot. 1970. Larger Mammals of Africa. London: Collins Press. 287 pp. DowsETT, R. J. 1966. Wet season game popu- lations and biomass in the Ngoma area of the Kafue National Park. The Puku (Occas. Pap. Dept. Game and Fish, Zambia), No. 4: 135-145. Efremov, J. A. 1940. Taphonomy: A new branch of Paleontology. Panam. Geol., 74: 81-93. . 1953. Taphonomie et annales geologi- ques. Ann. du Cent. d'Etud. et de Doc. Paleontol., No. 4. 164 pp. Estes, R. D. 1967. Predators and scavengers. Nat. Hist., 75(2, 3): 20-29, 38-47. . (1973), in press. Social organization of the African Bovidae. I.U.C.N., Symposium on Ungulate Behavior and Management. Field, C. R., and R. M. Laws. 1970. The dis- tribution of the larger herbivores in the Queen Elizabith National Park, Uganda. J. Appl. Ecol., 7(2): 273-294. Fitch, F. J., and J. A. Miller. 1970. Radio- isotopic age determinations of Lake Rudolf artefact site. Nature, 226(5242): 226-228. Foster, J. B. 1967. Nairobi National Park Game Census, 1966. East Afr. Wildl. J., 5: 112-120. Guggisberg, C. a. W. 1972. Crocodiles. New- ton Abbot, England: David & Charles, Pub- lishers. 195 pp. Harris, L. D. 1970. Some structural and functional attributes of a semi-arid East African ecosystem. Ph.D. Dissertation, Michi- gan State University. Harvard Tables of the Cutmulative Binomial Probability Distribution. 1955. Cam- bridge, Massachusetts: Harvard University Press. 503 pp. Haynes, V. 1968. Radiocarbon: analysis of in- organic carbon of fossil bone and enamel. Science, 161: 687-688. de Heinzelin, J., F. H. Brown and F. C. Howell. 1971. Pliocene/Pleistocene for- mations in the Lower Omo Basin, southern Ethiopia. Quaternaria, 13: 247-268. Isaac, G. L. 1967. To\\'ards the interpretation of occupation debris: some experiments and observations. The Kroeber Anthropol. Soc. Pap. 37: 31-57. Isaac, G. L., R. E. F. Leakey and A. K. Behrens- MEYER. 1971. Archeological traces of early hominid activities, east of Lake Rudolf, Kenya, Science, 173: 1129-1134. Johnson, L. C. 1965. Morphological analysis in Pathology. In Frost, H. M. (ed. ). Bone Biodynamics. (Henry Ford Hospital Int. Symp. ). Boston: Little, Brown and Co. 543-654. KOxNiZESKi, R. L. 1957. Paleoecology of the middle Pliocene Deer Lodge Local Fauna, western Montana. Bull. Geol. Soc. Am. 68: 131-150. Kruuk, Hans. 1972. The Spotted Hyaena. Chicago: The University of Chicago Press. 335 pp. KuRTEN, B. 1953. On the variation and popu- lation dynamics of fossil and recent mammal populations. Acta Zool. Fenn., 76: 5-118. Lamprey, H. F. 1963. Ecological separation of the large mammal species in the Tarangire Game Reserve, Tanganivika. East Afr. Wildl. J., 1: 63-92. Langbein, W. B., and L. B. Leopold, 1968. River channel bars and dunes — theory of kinematic waves. U. S. Geol. Surv. Prof. Pap. No. 422-L: 1-20. Leakey, R. E. F. 1973. Fiuther evidence of Lower Pleistocene hominids from East Rudolf, North Kenya, 1972. Nature, 242: 170-173. Leopold, L. B., M. G. Wolman and J. P. Miller. 1964. Fluvial Processes in Geomorphology. San Francisco: W. H. Freeman and Co. 522 pp. East Rudolf Paleoecology • Belirensmeijer 569 Leopold, L. B., W. VV. E^t^IETT and R. M. Myrick. 1966. Channel and hillslope pro- cesses in a semiaiid area. New Mexico. V. S. Geol. Surv. Prof. Pap. No. 352-G: 193-253. LOBOVA, E. V. 1967. Soils of the Desert Zone of the USSR Jenisak'iii: Israel Program for Scientific Translations. 405 pp. LovERiDGE, A. 1941. Revision of the African terrapin of the family Pelomedusidae. Bull. Mus. Comp. Zool., 88(6): 462-524. LovERiDGE, A., AND E. E. WiLLiAMs. 1957. Re- vision of the African tortoises and turtles of the suborder Cryptodira. Bull. Mus. Comp. Zool, 115(6): 163-557. Maglio, V. J. 1972. Vertebrate faunas and chronology of hominid-bearing sediments east of Lake Rudolf, Kenya. Nature, 239: 379- 385. McKee, E. D., E. J- Crosby and H. L. Berryhill, Jr. 1967. Flood deposits. Bijou Creek, Colorado, June, 1965. 1. Sediment. Pet., 37 (3): 829-851. Millar, C. E., L. M. Turk and H. D. Foth. 1966. Fundamentals of Soil Science. New York: John Wiley & Sons. 491 pp. Muller, a. H. 1957. Lehrbuch der Palao- zoologie. Vol. 1: Allgemeine Grundlagen. Jena: Gustav Fischer Verlag. 322 pp. Olson, E. C. 1952. The evolution of a Permian vertebrate chrono-fauna. Evolution, 6(2): 181-196. . 1958. Fauna of the Vale and Chosa: 14. Fieldiana, Geol., 10(32): 397-448. Patterson, B., A. K. Behrensme\'er, and W. D. Sill. 1970. Geology and fauna of a new Pliocene localitv in north-western Kenya. Na- ture, 212(5062): 918-921. Payne, J. E. 1965. Summer carrion study of the baby pig, Siis scrofa. Ecology, 46(5): 592-602. Pettijohn, F. J. 1957. Sedimentary Rocks. New York: Harper & Row, Publishers. 718 pp. Pettijohn, F. J., P. E. Potter and R. Siever. 1972. Sand and Sandstone. New York: Springer-Verlag. 618 pp. Reeves, C. C. 1970. Origin, classification and geologic history of caliche on the southern high plains, Texas and eastern New Mexico. J. Geol., 78: 352-362. Reif, Wolf-Ernst. 1971. Zur Genese des Mus- chelkalk-Keuper-Grenzbonebeds in Siidwest- deutschland. Neues lahrb. Geol. Palaontol. Abh., 139(3)- 369-404. Rittenhouse, G. 1943. Transportation and de- position of heavy minerals. Bull. Geol. Soc. Am., 54(12): 1725-1780. Sadek-Kooros, H. 1966. Jaguar Cave: an early man site in the Beaverhead Mountains of Idaho. Ph.D. Dis.sertation. Harvard Uni- \ersit\-. Department of Anthropology. Sanders, H. L. 1968. Marine benthic diversity: a comparative study. Am. Nat., 102(925): 243-282. Sciiafer, W. 1972. Ecologj' and Palaeoecology of Marine Environments. Craig, G. (ed.), I. Oertel (transl.). Chicago: The University of Chicago Press. 568 pp. ScHALLER, G. B. 1972. The Serengeti Lion. Chicago: The University of Chicago Press. 480 pp. Selous, F. C. 1908. African Nature Notes and Reminiscences. London: MacMillan and Co. 356 pp. Shapiro, A. II. 1961. Shape and Flow: The Fluid Dynamics of Drag. New York: Double- day & Co., Inc., Anchor Books. 186 pp. Sheppe, W., and T. Osborne. 1971. Patterns of use of a floodplain by Zambian mammals. Ecol. Monogi-., 41(3): 181-205. Shotwell, J. A. 1955. An approach to the paleoecology of mammals. Ecology, 36(2): 327-337. . 1963. The Juntura Basin: studies in earth history and paleoecology. Trans. Am. Phil. Soc, (N. S.), 53(1): 3-77. Simpson, G. G., A. Roe, and R. C. Lewontin. 1960. Quantitative Zoology. New York: Harcourt, Brace & World, Inc. 440 pp. Stewart, D. R. M. 1963. Wildlife census. Lake Rudolf. East Afr. Wildl. J., 1: 121. Van Lawick-Goodall, H. and J. 1971. Inno- cent Killers. Boston: Houghton Mifflin Co. 222 pp. Visher, G. S. 1965. Use of vertical profile in environmental reconstruction. Bull. Am. Assoc. Pet. Geol., 49(1): 41-61. VooRHiES, M. 1969. Taphonomy and population dynamics of an early Pliocene \'ertebrate fauna, Knox County, Nebraska. Contrib. to Geol. Univ. of Wyoming. Spec. Pap. No. 2. Laramie, Wyoming: Uni\'. Wyoming Press. 69 pp. Walsh, J. and R. G. Dodson. 1969. Geology of Northern Turkana. Rep. No. 82, Geol. Surv. Kenya. 42 pp. Weigelt, J. 1927. Rezente Wirbeltierleichen und ihre paliiobiologische Bedcutung. Leip- zig: Verlag \on Max Weg. 208 pp. White, T. 1955. Obser\'ations on the butcher- ing techniques of some aboriginal peoples, numbers 7, 8, and 9. Am. Antiq., 21(2): 170-178. Williams, G. E. 1971. Flood deposits of the sand-bed ephemeral streams of Central Aus- tralia. Sedimentology, 17: 1—40. 570 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 APPENDIX 1 Measurkments of densities, volumes and wet weights of modern bones. Densities are calcu- lated FOR THE BONES AFTER THEIR PORE SPACES WERE FILLED WITH WATER. DENSITIES Skeletal part Ovis ( sheep ) MCZ 1939 Redunca ( reedbuck ) MCZ 14917 Hylochoenis ( forest hog) MCZ 27851 Damaliscus (topi) MCZ 15724 Equus ( zebra ) MCZ 5003 Hippo- potamus MCZ 5020 HUMERUS 1.53 1.40 1.55 1.51 1.77 1.74 RADIUS 1 ULNA j 1.64 1.58 1.16 1.66 1.72 1.41 1.45 1.69 FEMUR 1.45 1.36 1.41 1.37 1.36 1.79 TIBIA 1.46 1.23 1.54 1.71 1.45 1.55 METATARSAL 1.44 1.33 1.07 1.62 1.68 1.50 METACARPAL 1.35 1.45 1.15 1.51 1.52 1.34 ASTRAGALUS 1.68 1.81 1.28 1.66 1.16 1.46 CALCANEUM 1.62 1.37 1.22 1.50 1.00 1.52 PODIAL #1 1.43 1.13 1.30 1.46 1.23 1.31 #2 — 1.25 1.20 1.48 1.01 1.46 PHALANX #1 1.45 1.34 — 1.40 1.00 1.37 #2 1.60 — 1.36''» 1.34 1.02 1.29"> #3 1.06 — 1.02" > 1.01'" 1.05<" 1.07' '> TEETH M 2.19 — — 2.23 2.08 2.00 PM — — — 2.24 1.97 1.97 C — — 1.53 — — 1.83 I - — 1.53 1.88 _ 1.74 RIB #1 1.11 1.54 1.41 1.36 1.22 1.63 #2 — 1.43 1.20 1.08 1.84 VERTEBRA ATLAS 1.24 .78 1.56 1.43 1.28 1.64 AXIS 1.07 .94 1.41 1.33 1.24 1.87 CERVICAL 1.04 1.13 — 1.11 .98 1.82 THORACIC 1.06 1.05 1.21 1.30 1.11 1.26 LUMBAR .89 1.23 1.23 1.13 .99 1.36 SACRUM 1.11 .92 1.18 1.07 _ _ PATELLA 1.07 1.07 1.01 1.30 .64 1.24 PELVIS 1.19 1.17 — STERNUM .97 — _ _ ^ _ SKULL 1.42 1.39 __ __ JAW (1/2) 1.43 1.74 _ 1.58 _ SCAPULA 1.65 1.88 _ 1.53 VERT. CENT. #1 .98 .75 1.60 1.00 1.06 1.29 #2 1.09 — 1.40 _ 1.00 ULNA, PROX. — .90 _ 1.21 SESAMOID #1 — — _ _ _ 1.46 #2 — — — _ — . 1.18 HUM. PROX. 1.26 1.34 1.42 1..32 1.63 1.55 DIST. 1.75 1.96 1.69 1.81 1.83 1.96 R/U. PROX. 1.64 1.47'"^ 1.65 1.96"^' 1.29 1.74 DIST. 1.59 1.72'«' 1.67 1.52"" 1.50 1.63 FEM. PROX. 1.47 1.44 1.50 1.58 1.33 1.83 DIST. 1.42 1.30 1.29 1.21 1.45 1.64 TIB. PROX. 1.32 1.20 1.27 1.43 1.27 1.30 DIST. 1.64 1.28 1.96 2.30 1.77 1.91 MT. PROX. 1.31 _ 1.48 1.49 DIST. 1.56 — _ 1.55 1.36 MC. PROX. 1.38 _ _ 1.33 1.40 _ DIST. 1.25 — _ 1.37 1.40 SCAPULA (GLENOID) 1.30 1.48 1.58 1.29 1.32 R = Radius only. t = Terminal Phalanx East Rudolf Paleoecology • Behrensmeyer 571 VOLUMES ( Cubic Centimeters ) Skeletal part Ovis ( sheep ) MCZ 1939 Rediinca ( recdbvick ) MCZ 14917 Hijlochoenis ( forest hot; ) MCZ 27851 Damaliacus (topi) Mc;z 15724 Equits (zebra) MCZ 5003 Hippo- potamus MCZ 5020 HUMERUS 53.5 67.0 404 225 310 2542 RADIUS ) ULNA j 39.6 39.0 11.0 232 148 39.0 303 1700 FEMUR 65.0 116 383 296 635 3000 TIBIA 56.0 128 186 246 411 1852 METATARSAL 21.0 46.0 23.0 117 140 144 METACARPAL 20.5 40.0 25.5 116 176 174 ASTRAGALUS 4.1 7.2 27.5 20.4 63 296 CALCANEUM 5.5 14.1 44.5 36.0 87 352 PODIAL #1 2.8 6.4 12.2 16.2 10.4 150 #2 — 2.0 8.8 5.6 14.8 94 PHALANX #1 2.9 4.5 12.0 17.9 48 78 #2 1.0 4.8 7.4'" 8.1 — 17.8'" #3 1.7 8.5'" 7.9'" 20.0'" 10.7'" TEETH M 1.7 — — 3.6 25.4 73.0 PM — _ — — _ 17.6 C _ _ 42.2 _ — 290 I _ — — 0.8 — 130 RIB #1 10.0 5.2 55.0 14.0 26.5 229 #2 — 7.7 31.8 33.0 25.0 — VERTEBRA ATLAS 25.0 24.4 75.0 63.0 139 866 AXIS 30.5 19.8 56.0 67.0 155 500 CERVICAL 24.0 18.4 — 62.0 170 450 THORACIC 14.0 8.4 57.0 22.8 64.0 480 LUMBAR 15.6 16.6 45.0 40.0 49.0 480 SACRUM 30.0 33.0 165 125 — — PATELLA 2.9 5.2 28.0 20.0 45.0 244 PELVIS 107 64.0 — — _ _ STERNUM 3.2 — — — — — SKULL 209 124 — — — — JAW (1/2) 39.5 23.0 — 119 — — SCAPULA 26.0 20.5 — 110 — — VERT. CENT. #1 5.1 6.0 25.0 20.0 31.0 233 #2 4.7 _ 25.0 30.0 28.0 — ULNA, PROX. _ 7.8 _ 20.0 _ SESAMOID #1 — "^ — _ _ 7.0 #2 — — — — — 10.6 HUM. PROX. 32.5 35.0 220 129 168 1420 DIST. 23.4 24.0 184 94.0 150 1122 R/U. PROX. 19.5 21.0"^^ 117 65.0"" 170 822 DIST. 20.1 18.0""' 115 84.0"" 147 878 FEM. PROX. 32.0 55.0 179 128 325 1466 DIST. 33.0 61.0 204 168 299 1634 TIB. PROX. 31.0 66.0 113 154 235 1100 DIST. 25.0 62.0 73.0 96.0 168 750 MT. PROX. 11.5 — _ 61.0 68.0 — DIST. 9.7 _ 58.0 68.0 _ MC. PROX. 10.0 — _ 60.0 82.0 _ DIST. 12.0 _ _ 58.0 90.0 _ SCAPULA (GLENOID) 16.6 13.0 97.0 65.0 100 - 572 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 WET WEIGHTS (Grams) Skeletal part Ovis ( sheep ) MCZ 1939 Redunca ( reedbuck ) MCZ 14917 Hylochoenis ( forest hog ) MCZ 27851 Damaliscus ( topi ) MCZ 15724 Equus ( zebra ) MCZ 5003 Hippo- potamus MCZ 5020 HUMERUS 82.1 93.5 626 340 550 4413 RADIUS ) ULNA { 64.9 61.6 12.8 385 255 55 440 2869 FEMUR 94.5 158 539 407 866 5364 TIBIA 82.1 158 286 420 597 2873 METATARSAL 30.3 70.8 24.7 190 244 216 METACARPAL 27.7 62.3 29.3 171 176 234 ASTRAGALUS 6.9 13.2 35.1 33.9 71 432 CALCANEUM 8.9 19.9 54.3 54.0 88 536 PODIAL #1 4.0 7.2 15.8 23.7 12.8 196 #2 — 2.5 10.6 8.3 15.0 137 PHALANX #1 4.2 6.3 — 21.9 49 107 #2 1.6 7.1 12.2'" 11.4 47 23" > #3 1.8 — 10.2"' 7.5"> 21"> 11.5'" TEETH M 4.4 — — 6.7 68.7 146 PM — — — 11.4 *'35.0* 35 C — — 64.5 — 530 I - — — 1.5 _ 226 RIB#1 11.1 8.0 77.8 18.7 34.2 374 #2 — 11.0 38.2 35.8 46.0 VERTEBRA ATLAS 30.9 19.0 117 90.0 178 1418 AXIS 32.6 18.7 78.7 89.0 193 934 CERVICAL 24.9 20.7 _ 69.0 166 818 THORACIC 14.8 8.8 69.2 29.7 71.3 606 LUMBAR 13.9 20.4 55.3 45.0 48.9 652 SACRUM 33.4 30.5 195 134 _ PATELLA 3.1 5.9 28.4 25.9 29.0 276 PELVIS 127 75.0 ... STERNUM 3.1 ^ _ _ _ SKULL 296 171 ^ _ _ JAW (1/2) 56.5 40.0 _ ^ _ _ SCAPULA 43.0 38.6 w 168 _ _ VERT. CENT. #1 5.0 4.5 40 25.0 33.0 300 #2 5.1 — 35 28.0 ULNA, PROX. — 6.4 ^^ 30.0 __ SESAMOID #1 — — ^ ^ 10.2 #2 - — — _ — 12.5 HUM. PROX. 41.0 47.0 313 170 275 2206 DIST. 41.0 47.0 313 170 275 2206 R/U. PROX. 32.0 31.0'^^ 192 127(R) 220 1434 DIST. 32.0 31.0"^^ 192 127<«) 220 1434 FEM. PROX. 47.0 79.0 269 203 433 2682 DIST. 47.0 79.0 269 203 433 2682 TIB. PROX. 41.0 79.0 143 220 298 1436 DIST. 41.0 79.0 143 220 298 1436 MT. PROX. 15.1 _ 90.0 88.0 _ DIST. 15.1 _ 90.0 88.0 ^ MC. PROX. 13.8 _ 85.0 122.0 DIST. 13.8 _ 85.0 122.0 SCAPULA (GLENOID) 21.5 19.3 153 84.0 132.0 - * M = Molar. East Rudolf Paleoecology • Behrensmeyer 573 APPENDIX 2 The value of r„ represents an idealized r^^i^..i^i- X Lj J r r- I quartz equivalent for the bone which dis- Calculation of Hydraulic Equivalence j .i rr .. r i regards the effects of shape. Processes of sediment transport are gen- It is difficult to predict the effects of erally explained in terms of quartz grains shape on bone-quartz equivalents. In some with a standard density of 2.65. Some work cases bone shape may decrease settling has been done on the hydraulic equivalence velocity by increasing the frictional drag ("equivalent settling velocity") of quartz on the bone, and this will reduce the size and particles with greater densities to show of the equivalent quartz grain. On the how small, dense grains sort out with larger, other hand, a bone shape (e.g., a stream- lighter ones ( Rittenhouse, 1943; Briggs, lined one) that reduces drag may increase 1962). However, there is a lack of infor- the size of the quartz equivalent. The mation on the hydraulic equivalence of orientation of a bone may have great effects quartz with particles of lower density such on settling velocit)^ and quartz equivalence, as bones. Thus, a metapodial dropped parallel to its Hydraulic equivalence can be considered lonig ^^^s may fall faster than a sphere of in terms of any two particles that have the equivalent volume, but the same bone same settling velocity. Given a particular dropped with its long axis horizontal could bone, it is possible to determine what size settle at a rate slower than that of a sphere, of quartz grain will settle at the same rate The same bone can alter from equivalence as the bone. For spheres, hydraulic equiva- to small or large quartz grains by slight lence to quartz can be easily calculated changes in orientation. In actual transport using the Impact law. If the settling veloc- situations, some bones tend to orient with ity for quartz (vq) is to equal the settling long axes parallel to current direction velocity of a bone (vb), then: (Voorhies, 1969:66-67), and these will rq = 1 '^07 / 1 ^ — 1 QA7 have maximum hydraulic equivalents for 160 / • ( pq - 1 ) • r„ - 1307 • ( pb - 1 ) • Tb tj^pjy volume. Bones also tend to orient per- "^Tfi^^"- l^^~n " pendicular to the current, and these will ^q ~ y^~ '' ^^ have smaller effective quartz equivalents. I Pb - J- ; • Th -pi^g bones that orient transverse to the 1-65 current should be more mobile in transport — one; situations. 7y = bone density ^^^^^^ ^^ ^ ^reat need for experimental r.i = radius of quartz grain '^°^^ '!^^^^^^ '""'^ ^^^°^ *,^^^ relationship he- rb = V. the nominal diameter ^^^^" ,^°"^ '^"^^j^S velocities and quartz of 1 ffiven bone equivalents and the actual current veloci- ties necessary for bone entrainment and If pb = l.5 and rb = 1.0 cm, then r,, = .30 cm. transport. 574 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 Plate 1. Surface textures of weathered and unweathered bones. A: a) Unweathered recent bovid radius, showing a smooth, "fresh" surface texture; b) Naturally weathered bovid femur from the recent East Rudolf thanatocoenose, showing slight roughening and cracking of the bone surface; c) Distal end of an equid femur from the recent East Rudolf thanatocoenose, showing extreme flaking and roughening of the bone surface. {Scale in 1 cm and 2 cm intervals) 6: a) Fossil astragalus from the 102-0201 channel sand, showing pre-burial abrasion; b) Fossil astrag- alus from the 103-0267 distributary channel and beach complex, showing considerable pre-burial abrasion; c) Recently weathered astragalus from the modern East Rudolf thanatocoenose, showing the typical cracked weathering pattern on its articular surface (Note: pattern lacking in a) and b)); d) Recent, unweathered astragalus; e) Unweathered and unabraded fossil astragalus from Lo- cality 8+6-0104 (floodplain). (Scale in 1 cm and 2 cm intervals) C: a) Distal end of a fossil humerus that was probably weathered prior to burial, showing the typical cracking pattern on its articular surface; b) Distal end of a recently weathered humerus, showing a similar cracking pattern; c) Recent, unweathered humerus; d) Fossil humerus from Area 8, East Rudolf, showing no sign of pre-burial weathering or abrasion. (Scale in 1 cm and 2 cm intervals) Plate 2. Fracture patterns in recent and fossil bones. A: a) "Sawtooth" fracture (right side of bovid pelvis); b) "Step" fracture (bovid metapodial); c) "Splin- tered" fracture (sheep rib); d) "Spiral" fracture (distal end of bovid humerus; e) Weathered bovid humerus (distal end) with a spiral fracture incurred prior to weathering. (Scale in 1 cm intervals) 6: Spiral fracture on the metatarsal of a recently killed giraffe, presumably caused by a hyaena. (Scale in 10 cm intervals) C: a) and b) Typical fracture patterns of bones after mineralization; c) Recent humerus (distal) show- ing spiral fracture; d) Fossil fragment of a diaphysis, showing a spiral fracture probably incurred prior to burial and mineralization. (Scale in 1 cm and 2 cm intervals) Plate 3. The trapping effect of surface vegetation on bones in the recent thanatocoenose on the delta of Laga Tulu Bor, lleret, East Rudolf. A: Bovid femur bound by shoreline grass and partially buried. (Scale in 10 cm intervals) B: Bovid skull and vertebrae, showing loose entrapment by grass. The horn cores are bound firmly to to the ground by warm tubes (just to right of camera lens cover). Plate 4. Recent sedimentary environments south of lleret. East Rudolf. A: A beach bar on the shore of the delta of Laga Tulu Bor, with the open lake to the right and a closed lagoon or back beach pond to the left. Pebbles and bone debris litter the beach. Beach bars such as this move shoreward seasonally with the annual rise in lake level (about 1 m fluctua- tion per year). Depositional environments such as this were probably active in the formation of Plio-Pleistocene deposits such as those of Localities 130-0201, 105-0208, 103-0267 and 103-0256. B: Laga Tulu Bor after a brief but heavy rainstorm, with a flow depth of about 1.5 m. The channel is normally dry for most of the year. A break in the gallery forest that fringes the channel is visible in the upper right of the photograph. This opens onto the grass-covered floodplain. Some charac- teristics of this depositional environment are probably comparable to Localities 102-0201 and 105- 1311. C: The upper part of the deltaic plain of Laga Tulu Bor, showing flooding of a low area (atrophied channel) after a heavy rain. This area lies in the transition zone between floodplain and deltaic plain. The sediment is primarily silt. This depositional environment is probably comparable to that of Locality 8+6-0104 (floodplain). East Rudolf Paleoecology • Behrensmeyer 575 B Plate 1 576 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 B Plate 2 East Rudolf Paleoecology • Behrensmeyer 577 Plate 3 578 Bulletin Museum of Comparative Zoology, Vol. 146, No. 10 Plate 4 827r 070 Harvard lllllll II II III I HI II MCZ LIbrarv llllllHI II III I II I II 2044 066 304 205 I i {Ml