HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology DOES NOT CiRPt a—. ogee wert RVARD UNI <, CAMBRI GE 30, w RETURN POSTAGE UARANTEED n eARP EBT Mn era ft Mera ei y ih i i : v a a NieME Pe PU i] ee 7 : ra a BREVIORA MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE, IN CAMBRIDGE NumMBERS 121-178 1960-1962 CAMBRIDGE, MASS., U.S.A. 1963 Edited By NExLpA E. WricHt No. No. No. No. 121. 123. . 126. 130. CONTENTS BREVIORA Museum OF COMPARATIVE ZOOLOGY NumBERS 121-178 1960 Three new species of Micrathena (Araneae, Argiopidae) from South America. By Arthur M. Chickering. 11 pp. March 10. Notes on certain species of Micrathena (Araneae, Argi- opidae) from South America. By Arthur M. Chick- ering. 7 pp. March 11. Alepisaurus brevirostris, a new species of lancetfish from the western North Atlantic. By Robert H. Gibbs, Jr. 14 pp. March 14. Anisian ammonoids from Malaya. By Bernhard Kum- mel. 8 pp., 1 pl. March 15. The luminous organs of Proctoporus (Sauria, Rep- tilia) — a re-evaluation. By Willard D. Roth. 12 pp. May 27. Mid-Seythian ammonites from Iwai formation, Japan. By Bernhard Kummel and Sumio Sakagami. 11 pp., 3 pls. June 3. Notes on the cranial anatomy of Necrolemur. By E. L. Simons and D. E. Russell. 14 pp. December 19. Size of endoceroid cephalopods. By Curt Teichert and Bernhard Kummel. 7 pp. December 20. Type and type locality of the Gulf Coast spiny softshell turtle, Trionyx spinifer asper (Agassiz). By Robert G. Webb. 8 pp., 2 pls. December 21. The mechanisms of carapacial and plastral hinges in chelonians. By R. V. Shah. 15 pp. December 22. No. No. No. No. plots . 132. 133. . 134. 136. 137. 138. ils . 140. . 141. . 142. A second record of the fossil rodent Palustrimus Wood. By Craig C. Black. 3 pp. December 30. The status of Sphaerodactylus pictus, with comments on the distribution of S. sputator and NS. sabanus. By Wayne King. 5 pp. December 30. 1961 On the generic limits in the family Pilidae (Proso- branchia: Mollusca). By Edward H. Michelson. 10 pp. February 27. Enzymatic constitution of Alsophis saliva and its bio- logical implheations. By George Hegeman. 8 pp. February 28. Notes on Hispaniolan herpetology. 2. A review of the Anolis semilineatus group with the description of Anolis cochranae, new species. By Ernest E. Wil- hams and A. Stanley Rand. 11 pp. April 7. Notes on Hispaniolan herpetology. 3. The evolution and relationships of the Anolis semilineatus group. By Ernest E. Williams. 8 pp. April 8. Notes on Hispaniolan herpetology. 4. Anolis koopmani, new species, from the southwestern peninsula of Haiti. By A. Stanley Rand. 4 pp. April 10. Pfeiffer’s unfigured species of Strophocheilus (Megal- obulimus). By T. E. Crowley and T. Pain. 8 pp., 2 pls. June 14. A new species of Sphaerodactylus from northern Haiti. By James D. Lazell. 5 pp. June 15. A preliminary review of the Nearctic species of Siero- lomorpha (Hymenoptera). By Howard EK. Evans. 12 pp. June 27. Three new toads from South America: Bufo mamcor- ensis, Bufo spinulosus altiperuvianus and Bufo quechua. By José M. Gallardo. 8 pp., 3 pls. June 28. Australian carabid beetles VI. The tropical and some subtropical species of Pamborus, Mystropomus, and Nurus. By P. J. Darlington, Jr. 13 pp. June 30. No. No. No. . 143. . 144. 145. . 146. 147. 5 148: . 149. /4152. . 153. Miocene lizards from Colombia, South America. By Richard Estes. 11 pp. August 20. A large ophiacodont pelycosaur from the Pennsyl- vanian of the Pittsburgh region. By Alfred Sher- wood Romer. 7 pp. August 21. A new species of the cetomimid genus Gyrinomimus from the Gulf of Mexico. By Henry B. Bigelow. 2 pp. September 5. New rodents from the early Miocene deposits of Sixty- Six Mountain, Wyoming. By Craig C. Black. 7 pp. December 14. Australian carabid beetles VIII. Leiradira, especially the tropical species. By P. J. Darlington, Jr. 12 pp. December 15. Australian carabid beetles IX. The tropical Noton- omus. By P. J. Darlington, Jr. 14 pp. December 18. A preliminary study of the Silurian ceratiocaridids (Crustacea: Phyllocarida) of Lesmahagow, Scot- land. By W. D. Ian Rolfe. 9 pp. December 19. 1962 The genus Bethylus in North America (Hymenoptera : Bethylidae). By Howard E. Evans. 12 pp. Jan- uary 5. A new phyllocarid crustacean from the Upper Devon- ian ot Ohio By ww. DD. lan Rolfe.) 7% pp. 1 pl: January 12. New Australian dacetine ants of the genera Meso- struma Brown and Codiomyrmex Wheeler (Hymen- optera-Formicidae). By Robert W. Taylor. 10 pp. January 15. Anolis scriptus Garman 1887, an earlier name for Anolis leucophaeus Garman 1888. By A. Stanley Rand. 5 pp. February 15. Notes on Hispaniolan herpetology. 5. The natural his- tory of three sympatric species of Anolis. By A. S. Rand. 15 pp. April 4. No. No. No. No. No. No. 2 158: 159. 160. 164. 167. Notes on Hispaniolan herpetology. 6. The giant anoles. By Ernest E. Williams. 15 pp. April 12. The fossiliferous Triassic deposits of Ischigualasto, Argentina, and preliminary description of Ischigual- astia, a new genus of dicynodont. By Alfred Sher- wood Romer and C. Barry Cox. 9 pp. April 13. A rhachitomous amphibian, Spathicephalus, from the Mississippian of Nova Scotia. By Donald Baird. 9 pp., 1 pl. May 28. A fossil gerrhosaur from the Miocene of Kenya (Rep- tila: Cordylidae). By Richard Estes. 10 pp., 1 pl. May 29. Age in a small sample of bluefish (Pomatomus salta- trac (Linnaeus)). By Richard H. Backus. 4 pp. May 31. ‘wo new arthropod carapaces from the Burgess shale (Middle Cambrian) of Canada. By W. D. Ian Rolfe. 9 pps, pl. June 12: A comparative study of the respiratory muscles in Chelonia. By R. V. Shah. 16 pp. July 16. Australian carabid beetles X. Bembidion. By P. J. Darlington, Jr. 12 pp. July 25. New worm-lizards (Ancylocranium and Amphisbaena) from southeastern Tanganyika Territory. By Arthur Loveridge. 6 pp. July 26. Notes on the herpetology of Hispaniola. 7. New ma- terial of two poorly known anoles: Anolis monticola Shreve and Anolis christopher Williams. By Ernest K. Williams. 11 pp. August 22. An extinet solenodontid insectivore from Hispaniola. By Bryan Patterson. 11 pp. August 22. Lectotypes of species of Ogcocephalidae selected from syntypes in the Museum of Comparative Zoology. By Margaret G. Bradbury. 4 pp. September 5. Bathyclupea schroedert, a new bathyclupeid fish from the western tropical Atlantic. By Myvanwy M. Dick. 4 pp. September 5, No. No. No. No. No. No. No. 168. eo Wf la 173. PLT. Two new species of fossil talpid insectivores. By Katherine M. Reed. 6 pp., 1 pl. September 7. New records of inshore fishes from the Atlantic coast of Panama. By Ira Rubinoff and Roberta W. Rubi- noff. 7 pp. October 15. The brain of the emu (Dromacus novaehollandiae, Lath). I. Gross anatomy of the brain and pineal body. By Stanley Cobb and Tilly Edinger. 14 pp., 4 pls. November 16. Notes on amphisbaenids (Amphisbaenia; Reptilia), 6. Redescription and range extension of Amphisbaena spurrell. Boulenger. By Carl Gans. 8 pp., 3 pls. December 14. A new species of the rodent Pipestoneomys from the Oligocene of Nebraska. By Raymond Alf. 7 pp. December 14. New species of land mollusks from the Republica Dominicana. By William J. Clench. 5 pp., 1 pl. December 24. A new arctocyonid from the Paleocene of Wyoming. By Bryan Patterson and Paul O. McGrew. 10 pp. December 24. A picrodontid insectivore (?) from the Paleocene of Wyoming. By Paul O. McGrew and Bryan Patter- son. 9 pp. December 24. On the races of Kinixys belliana Gray. By R. F. Laurent. 6 pp. December 27. Rhipidistian classification in relation to the origin of the tetrapods. By Keith 8. Thomson. 11 pp., 1 pl. December 27. On a new species of the earthworm genus Trigaster Benham 1886 (Octochaetidae). By G. E. Gates. 4 pp. December 27. INDEX OF AUTHORS BREVIORA MUSEUM OF COMPARATIVE ZOOLOGY NuMBERS 121-178 1960-62 No. PNUIGEO aia WARY SNM O)INUD) uc. Reaucenty Reisen usr thar ode cc cpeteotean eee Estee cence ee ee aaa G2 BACKUGs IGHARD tele: feet 2A) Ae esc kien Sus he oN ete ees 159 ANB D se DONAID sium otros eyes, toe ated te ties sie ont < eRe ee 157 BIG HWONWey bhEINGEO YUE tae ees cE PMc et. 1. tags eT a ee a 145 STAG Req CRAIG Os 7c ele oe es eens | eee BE, 131, 146 BRADBURY VIA RGAR BT Grsuece, fires. eee 2 na) au is Ree 166 CRICKERING, ATER ME 2.7.4 Jae 2 Ag 121, 122 (CETEEN CEU NG TEISTAUMI 2) ate Dis eS oc dep aon oh) Se ae oes cia ae ee 173 C@OBBs iS TANIEIDY (1214 FENG AGRE bs EIR RE Sag co 170 Corrs, BARRY: fs) och eek ta Mien ah fe Be eens oa ak eee 156 Crowe. "Ti ose ech INS a AR Es Ue Sy 138 DARTINGTON: Ps lie Siete set A Le” cae 142, 147, 148, 162 Drees Viy VAN WY Mia pend. Se See enn seed eee a ay ere 167 IpINGERY Tuan... he Sd, A Ee i 170 TESTS ESIC EDAD) 13555: 0.8.) 4, uss cat cnt See Ne Ag EE ye 143, 158 EVANS, FIOWARD! JM) 3% 2cdoce @ee or Fee ee ore ae 140, 150 GSDTAR DOs: OSH ANG sc. sir Sohin eae 2 eye eee ee eee 141 GANS Ss CARRIL RA Ses 5 Sorte Se eee ees Seneca sae ee eee 171 GEARS! AGES Bye deve atah ete chars tots tal Gas eo O e eee 178 CESES IOS MEM sl gence MRR e oa sis d < srsbid «Sa daw able Oe 123 EG ee ini COR GIN ec eS iar PAs eee ae kev boson Ad.» vo Salis Beek ee 134 ESSG NGG VN ASYONTEG aes Batt eS cae ce sate ret eet cay aiihia tera dsavaliie sehat tren 132 KGoMiMite SER NIEFARD) Sita, ssn etd coon Gatun Gi oheole sate 124, 126, 128 Oy AURUERETNE es rae a ea es rear arty acre Be rncue de aViActarennyetei cr sient 176 SSAC RTG Tr are ACTER) Wie Niece teats on Spe Fee em aiaunees Sas esha scalentah o/s es 139 [Era oraei ae eToR ys: 10 = Nop eagle eee gt vise Per anager cn ree eRe eds 163 Vie Gare wie tle AUilay OG Pa ctncn.< retreat wearer oes terre nt Seer 174, 175 MICHIE SON: MED WIARDy Ele 9 yyhs98 doco sation 6 sie sites steel = a 133 EAA ONe EIMS Peetep eee ep ee te Ree lactic acer ica PR eRe or REE Pa 138 LP EEDRSONGUESRVANG 67.) ous io Slate satiate ones & eon nee tae 165, 174, 175 EVAINDS Aces IPAS 4 20s heaalcns a wave, Ohare ore) 135, 137, 153, 154 FEE ya CAC RETO RTIN Nn oty a cuenta ns meee) occu sp 2c) ceiemeehc eens 168 ENOL SHV ewe) WANT gee 5 cuchn ete, cea reo ees wis ce ee 149, 151, 160 ROMER. AGERED SHERWOOD < Uacidass ae on she ce ye mene ne 144, 156 RO TEV MARDI! cross snide wickets eee ose eae 125 IERIUES ERS RUN RIAL Gactig. ice Mieiie sp) eek Mere kets Bh tats oye sea ee tte mee 169 EVUBINORE: ROBERTA Wie s..cx08 Snobs ols Sopa ac) aie hana ee ae arate 169 LRHGISST0 G0 TDs 1 Dis aa eae isis ote re Ree eatin meee oie noe bined oa 127 SAOACUAURUTE SS UINETO Sz suite ot «cote geloccoeas Sic ath dict sveyerata see shake tater 126 STeUNist Rt een echo OLA an Nea Rete erie oes co o cross 130, 161 STMT] ON Die a Beene gee Nolo gar IC) Seah ronan 127 PAVERS NVOBERN: Wiaseinic tres « cat inthe eros ae < he = ae 152 MOREE MGURT fe ose a d.acile aac Sas se emer ate noes ene 128 PEHOMGO NG INEETE. OS: J sepsis aes oats ise a sane ocala ef ps oc =n iT AN SETSESGPEVOBS SR Gress crates a. after asheiovoetea seh nota axe aucirater anche" < ops iat 129 Wit PAMS DRNEST Bhp isk set acs accmm aes eee 135, 136, 155, 164 ee seine Ho ddan on o-.. ae A: ¥ ¥] iy < ae iy ; 7 2 CPS on |) Mee ty mt: ites ve ¥ : _ Shy ae ° , ee YS nd a HL BOLT OND 44 a 2 Hib reeanth, ucaeet® BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. Marc# 10, 1960 NuMBER 121 THREE NEW SPECIES OF MICRATHENA (ARANEAE, ARGIOPIDAE) FROM SOUTH AMERICA By ArtHur M. CHICKERING Albion College, Albion, Michigan During the summer of 1959 I had the privilege of examining the South American Micrathenae in the Museum of Comparative Zoology at Harvard College. Among the specimens in this col- lection I found representatives of what I am compelled to regard as new species. I am describing these in this brief paper under the names: Micrathena hamata sp. nov.; M. lata sp. noy.; and M. shealsi sp. nov. I have also added a few notes together with figures illustrating features of the epigynum of MW. fissispina (C. L. Koch), 1836, with the hope that they will aid somewhat in a more precise identification of this species. The types of the new species are deposited in the Museum of Comparative Zo- ology. It is with pleasure and a deep sense of gratitude that I again acknowledge my indebtedness to members of the staff of the Mu- seum of Comparative Zoology at Harvard College for their con- tinued encouragement over a period of more than twenty-five years. Persons now active on the staff of the museum and chiefly responsible for this encouragement may be specifically named as follows: Dr. A. S. Romer, Director; Dr. P. J. Darlington, Jr., Curator of Insects; Dr. Herbert W. Levi, Associate Curator of Arachnology; Miss Nelda E. Wright, Editor of Publications. Several other members of the museum staff have also greatly aided me by providing privileges of the laboratories, collections, and library. i) BREVIORA No. 121 MIcrRATHENA FIssispIna (C. L. Koch), 1836 (Figures 1-3) Acrosoma fissispina C. L. Koeh, 1836 Plectana fissispina Walckenaer, 1841 M. fissispina Simon, 1895 M. fissispina Reimoser, 1917 M., fissispina Roewer, 1942 M. fissispina Bonnet, 1957 There are three mature females in the Nathan Banks Collection in the Museum of Comparative Zoology. These are simply la- belled as having come from Brazil with no dates of collection aS = —@e =~ =O) Sires SS SS SSS = External Anatomy of Micrathena Figures 1-3. M. fissispina (C. L. Koch). Figs. 1-3. Epigynum: from below; profile, right side; posterior surface, respectively. assigned. The abdominal spination is a fairly reliable feature for identification, but the epigynum is also important. Since this organ appears somewhat different than represented in Reimoser’s (1917) figures I am offering three more drawings with the hope that they will help others to a more exact identification of this species. The male remains unknown. 1960 NEW SPECIES OF MICRATHENA a MIcRATHENA HAMATA Sp. Nov. (Figures 4-7) The two mature males treated in this part of this paper were found filed in the Banks Collection in the Museum of Compara- tive Zoology with females considered to belong to MW. petersi (Taez.). The females are probably immature specimens of WM. scxspinosa (Hahn). The males appear to be new to science and, consequently, one has been selected as the holotype and is here- with deseribed in accord with my usual procedure. External Anatomy of Micrathena Figures 4-7. M. hamata sp. nov. Fig. Fig. Fig. Fig. Body of male, dorsal view. . Left palpal tarsus and tibia. Palpal tarsal hook; nearly retrolateral view. . Ibid., a different view. “SVD Ol 4 BREVIORA Nomizi Male holotype. Total length 4.68 mm. Carapace 1.92 mm. long, 1.105 mm. wide opposite second coxae where it is widest; .455 mi. tall; nearly level from behind PME to beginning of posterior declivity. Median thoracie fovea a shallow longitudinal depres- sion. Dorso-lateral foveae lacking. Eyes. Eight in two rows as usual in the genus; lateral ocular tubercles weakly developed; median ocular tubercles bearing ME quite pronounced. Viewed from above, both rows strongly recurved; viewed from in front, anterior row nearly straight, posterior row moderately procurved; central ocular quadrangle wider behind than in front in ratio of 22 : 17, about as long as wide behind. Ratio of eyes AME : ALE : PME : PLE =6 :6.5 : 8 : 5.5 (long diameter used when there are differences). AME separated from one another by four-thirds of their diameter, from ALE by nearly five-halves of their diameter. PME sepa- rated from one another by nearly three-halves of their diameter, from PLE by about three-halves of their diameter. Laterals separated from one another by two-thirds the diameter of AME. Clypeus strongly receding; height equal to about twice the di- ameter of AME. Chelicerae, Maxillae, and Lip. Apparently quite normal for males in the genus. Fragility of specimen precludes examination of fang groove for marginal teeth. Sternum. Very rugulose; with a marked transverse groove between third and fourth coxae; fourth coxae nearly touching. Legs. 4123. Width of first patella at ‘‘knee’’ .11913 mm., tibial index of first leg 11. Width of fourth patella at ‘‘knee”’ 12996 mm., tibial index of fourth leg 13. (All measurements in millimeters ) Femora Patellae Tibiae Metatarsi Tarsi Totals Ile .990 374 .748 594 418 3.124 2. 1.012 .396 .660 550 .396 3.014 3. .704 .264 .396 .330 O02 2.046 4. 1.078 330 .706 .660 1.450 3.224 Palp 400 .220 330 aan 2.682 1.632 1 Estimated because of loss of both fourth tarsi. 2 Including tarsal hook, 1960 NEW SPECIES OF MICRATHENA 5 There is no ventral hook on first coxa nor any corresponding proximal, prolateral femoral ridge and groove on the second femur. There also seems to be a complete lack of modified spines on the legs. Palp. Essential features shown in Figures 5-7. The tarsal hook and tibia appear to be quite distinctive. Abdomen. General shape as shown in Figure 4. It seems high- ly probable that the female of the species will be found to have a series of prominent paired spines, remains of which seem to be present in the male. Color in alcohol. Carapace a nearly uniform rich reddish brown. Sternum somewhat lighter. Legs and mouth parts with various shades of brown. Abdomen: Dorsum yellowish along each dorso-lateral margin; broadly and irregularly brownish in the middle with a central lighter irregular stripe and some indi- cation of reduced black spots; venter irregularly brownish, yel- lowish and nearly black. Type locality. Holotype male and one paratype male in the Nathan Banks Collection from Para, Brazil. No date of collection is given. The female is unknown. MICRATHENA LATA Sp. nov. (Figures 8-12) The specimen described below was filed in the Reimoser Col- lection in the Museum of Comparative Zoology at Harvard Col- lege as M. digitata (C. L. Koch). This is obviously an error and, since I ean find no record of it in the literature I am compelled to regard it as new and am describing it as such. Female holotype. Total length from anterior margin of base of chelicerae to posterior border of abdomen 7.15 mm. Width of abdomen at base of anterior spines 2.925 mm.; width between tips of larger posterior spines 13.325 mm. Carapace smooth; with median thoracic fovea a well defined pit; not markedly raised behind median fovea; with no dorso-lateral foveae. Eyes. Eight in two rows as usual in the genus. Viewed from above, both rows moderately recurved; viewed from in front, anterior row nearly straight, posterior row gently procurved. Central ocular quadrangle wider behind than in front in ratio of about 3 : 2; wider behind than long in ratio of 9 : 7. Ratio 6 BREVIORA No. 121 of eyes AME : ALE : PME : PLE = 8 :8 : 10: 8.5. AME separated from one another by five-fourths of their diameter, from ALE by six times their diameter. PME separated from one another by three-halves of their diameter, from PLE by nearly six times their diameter. Laterals separated from one another by three-eighths of the diameter of ALE. Height of clypeus equal to about seven-fourths of the diameter of AME. Chelicerae. Normal to genus; unable to observe teeth along fang groove because of fragility of holotype. External Anatomy of Micrathena Figures 8-10, M, lata sp. nov. Fig. 8. Outline of body of female holotype; dorsal view. Figs. 9, 10. Epigynum; profile, right side and posterior surface, re- spectively. Lip and Maxillae. Normal to genus; details considered unim- portant in the description. Sternum. Extended; with deep lateral notches and with six exaggerated marginal tubercles; with posterior end extended between fourth coxae as a prominent tubercle nearly one-third as long as entire sternum. Legs and Spines. Only two legs remain entire, hence details not recorded. In general quite normal to genus. True spines 1960 NEW SPECIES OF MICRATHENA =~) rare; the usual numerous setigerous tubercles moderately well developed. Abdomen. Extraordinarily broadened posteriorly ; with a pair of nearly straight anterior marginal spines extended nearly to PME; with a pair of very small lateral marginal spines; the posterior end is broadly bifurcated and each fork is subdivided into two spines thus making a total of eight. Epigynum. General features shown in Figures 9, 10, and 12. External Anatomy of Micrathena Figures 11 and 12, WM. lata sp. nov. Fig. 11. Posterior end of female holotype. Fig. 12. Epigynum, from below. Color in alcohol. Light reddish brown with dark streaks and irregular spots. Type locality. Holotype female from Theresapolis, Brazil, with no date of collection given. There are no paratypes and the male is unknown. 8 BREVIORA No. 121 MICRATHENA SHEALSI sp. nov. (Figures 13-17) Female holotype. Total length from AME to tip of posterior spines 9.23 mm. Carapace 2.79 mm. long; 2.08 mm. wide opposite second coxae where it is widest; about .98 mm. tall behind well External Anatomy of Wicrathena Figures 13-17, M. shealsi sp. nov. Fig. 13. Outline of female holotype, dorsal view. Fig. 14. Right lateral side of abdomen to show spines on bifurcation. Figs. 15-17. Epigynum: from below, right lateral side, and from posterior view, respectively. defined median fovea where it is strongly gibbous; with three pairs of dorso-lateral foveae as represented in Figure 13. Eyes. Kight in two rows as usual; viewed from above, both rows moderately recurved; viewed from in front, both rows 1960 NEW SPECIES OF MICRATHENA 9 gently proeurved, measured by centers. Central ocular quad- rangle wider behind than in front in ratio of about 8 : 7, only slightly wider behind than long. AME separated from one another by their diameter, from ALE by nearly 3.5 times their diameter. PME separated from one another by their diameter, from PLE by three times their diameter. Ratio of eyes AME ALE : PME : PLE=10 :10 : 12 :7 (long diameters used when differences exist). Laterals separated from one another by nearly the radius of ALE. Height of clypeus equal to nearly 1.2 times the diameter of AME. Chelicerac. In general, normal to the genus; promargin of fang groove with three teeth, the middle one bidental; retro- margin with three large teeth. Mazillae. Apparently completely typical of the genus in all observed features. Lip. Wider than long in ratio of about 4 : 3; transversely erooved in basal third. Sternal suture procurved. Sternum. Elongated scutiform ; moderately convex ; only with well developed antero-lateral tubercles: not continued between fourth coxae; sparsely supplied with moderately long stiff bristles. Legs. 4123. Width of first patella at ‘‘knee’’ .325 mm., tibial index of first leg 11. Width of fourth patella at ‘‘knee’’ .303 imm., tibial index of fourth leg 11. (All measurements in millimeters ) Femora Patellae Tibiae Metatarsi Tarsi Totals 1 2.860 .990 2.100 1.950 .910 8.810 2. 2.600 975 1.820 1.755 .845 7.995 3: 1.625 .650 .980 1.040 aA ts) 5.010 4. 3.250 .780 2.015 2.080 .910 9.035 Spines on lees apparently unnoteworthy ; many are lost and scars are difficult to locate with certainty. Abdomen. General features essentially as shown in Figures 13 and 14. Moderately flattened. With no anterior marginal spines; with a pair of lateral dorsal spines of moderate length ; bifurcated posteriorly with each fork subdivided into two spines with the larger below. 10 BREVIORA No. 121 Epigynum. Unlke that seen in any other species; anterior border of modified portion a granulated rim; features essentially as shown in Figures 15-17. Color in alcohol. Legs and mouth parts a dull reddish brown, lighter beneath. Sternum light brownish. Carapace with a brownish central stripe and a broad dark brown stripe on each side (both represented by stippling in Figure 13) ; remainder of central region yellowish. Abdomen: irregularly yellowish white dorsally with nearly black margins; venter light yellowish from genital furrow to base and lateral sides of cone surrounding spinnerets ; the cone with a nearly black circular ring ; remainder of venter irregularly dark brown or black with yellowish spots and streaks. Probably some loss of coloration from long preser- vation. Type locality. The holotype was simply labelled: Argentine, Sunchal, with no date of collection given. Apparently Cockerell was the collector. The species is named in honor of Dr. J. G. Sheals, Department of Zoology, British Museum (Natural His- tory). BIBLIOGRAPHY BONNET, PIERRE. 1957. Bibliographia Araneorum. Tome II, 3me partie. Les Artisans de 1’Imprimerie Douladoure, Toulouse, France. CAMBRIDGE, O. P. and F. P. CAMBRIDGE. 1889-1905. Arachnida-Araneida. Vols. I-Il. Im: Biologia Centrali- Americana. Dulau & Co., London. IXEYSERLING, GRAF EUGEN VON, 1892. Die Spinnen Amerikas. Epeiridae. Verlag von Bauer & Raspe, Niirnberg, Germany. Kocu, C. L. 1836. Die Arachniden. Niirnberg, vol. III, pp. 1-119. REIMOSER, EDUARD. 1917. Die Spinnengattung Micrathena. Verh. Zool. Bot. Ges. Wien, vol. 67, pp. 73-160, pls. 1-9, figs. 1-31. 1960 NEW SPECIES OF MICRATHENA 11 RoeweERr, C. FR. 1942. Katalog der Araneae. Vol. 1. Kommissions Verlag von ‘¢Natura.’’ Bremen. SIMON, EUGENE. 1892-1903. Histoire Naturelle des Araignées. Deuxiéme Edition. 2 Vols. Librarie Encyclopédique de Roret, Paris. WALCKENAER, CH. A. 1841. Histoire Naturelle des Inseetes. Apteres. Paris, vol. I1. ; sar bite! — a a Ne, fe. were De —- ayes Jaieaisi Borer ira | : in D4 di, ile ne ee oa od we opie 7 a] Se i at a an i.3t Saw + —- are pan - big . Berea ’ i ‘ . ’ ee 4 , > 7 ae we aS : , oT . a G : : it Pc 5 “> a 7 - 5 a 7 - rs ss win nh le “ oa ' 7 “~ ~m _ -_ t x e 9 a j / ba) > = = . @ - BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. Marcu 11, 1960 NuMBER 122 NOTES ON CERTAIN SPECIES OF MICRATHENA (ARANEAE, ARGIOPIDAE) FROM SOUTH AMERICA By ArtHuur M. CHICKERING Albion College, Albion, Michigan During a period of work in the Museum National d’Histoire Naturelle in Paris during the summer of 1958 Dr. Herbert W. Levi, Associate Curator of Arachnology in the Museum of Com- parative Zoology at Harvard College, examined types of nine species of Micrathena Sundevall 1833, all originally described from South America by the great arachnologist, Eugéne Simon. All of these species are poorly known and most of them have not appeared in collections since the originals were studied by their author. All were briefly described in 1896; four were mentioned in 1895 and accompanied by five simple figures. During the examination of the types mentioned Dr. Levi made free-hand drawings of the dorsal surface of the abdomens to show general form and spination. He also made careful drawings of the exter- nal genitalia when the latter were available; these were made with the use of a reticule with squares. All of the drawings made by Dr. Levi were turned over to the author to use as he saw fit in connection with his study of the genus. The figures appearing in this paper were made directly from Dr. Levi’s original pencil drawings. The outline figures of abdomens are freehand copies with enlargement; the drawings of genitalia were made with tracing paper directly from Dr. Levi’s originals. It has seemed worth while to present these data, thus obtained, with the hope that they will be of some help to others who may continue the study of this most interesting genus. 2 BREVIORA INowi22 MICRATHENA ACICULATA Simon, 1897 (Figure 1) M. aciculata Petrunkevitch, 1911 M. aciculata Roewer, 1942 M, aciculata Bonnet, 1957 The type is an immature female from Venezuela. Apparently the species has not been reported in collections since the original was taken. The general form of the abdomen with its spination, 2 ire EL External Anatomy of Micrathena Figure 1, M. aciculata, abdomen, dorsal view. Figures 2, 3. M. gaujoni, abdomen, dorsal view and epigynum, respectively. Figures 4,5. Mf. hamifera, abdomen, dorsal view and epigynum, respectively. seen in dorsal view, is shown in Figure 1. The stippled area is black and the postero-lateral corners are white. The male is unknown. MIcRATHENA GAUJONI Simon, 1897 (Figures 2, 3) M. gaujoni Petrunkevitch, 1911 M. gaujoni Reimoser, 1917 1960 CERTAIN SPECIES OF MICRATHENA 3 M. gaujoni Roewer, 1942 M. gaujoni Bonnet, 1957 Simon stated that the type was 8.7 mm. long and similar to M. fissispina (C. Koch), but this hardly seems correct. There are four pairs of spines, the second pair the longest (Fig. 2). The epigynum has a pair of depressions directed posteriorly be- neath an overhanging rim (Fig. 3). The type is from Ecuador and the male is unknown. MicraATHENA HAMIFERA Simon, 1897 (Figures 4, 5) The length of the holotype was given as 9 mm. The general form of the abdomen with its spination is shown in Figure 4. The dorsal surface is like white enamel in general appearance. Figure 5 shows the form of the epigynum from ‘‘slightly be- hind’’; just posterior to the curved boundary there is a large ‘*selerotized knob.’’ The female is known only from Peru and the male is still unknown. MIcRATHENA IMBELLIS Simon, 1895 (Figure 6) M. imbellis Simon, 1897 M. imbellis Petrunkevitch, 1911 M. imbellis Reimoser, 1917 M. imbellis Roewer, 1942 M. imbellis Bonnet, 1957 Simon (1895) included a figure showing the right side of the abdomen with no spines. Reimoser (1917) just mentioned the species and did not include it in his further treatment of the venus. The general appearance of the dorsal surface of the abdomen is shown in Figure 6; the stippled areas in the figure are black in the type. Dr. Levi has determined that the type is an immature female from Venezuela. The male is unknown. MIcRATHENA PERLATA Simon, 1895 (Figures 7, 8) M. perlata Simon, 1897 M. perlata Petrunkevitch, 1911 M. perlata Reimoser, 1917 4 BREVIORA No. 122 M. perlata Roewer, 1942 M. perlata Bonnet, 1957 The length of the female type is given as 6 mm. Dr. Levi has found that it also is immature. The general appearance of the dorsal surface of the abdomen is shown in Figure 7. The im- mature female is accompanied by a male which may or may not External Anatomy of Micrathena Figure 6. M. imbellis, abdomen, dorsal view. Figures 7, 8. M. perlata, abdomen, dorsal view and male palp, respectively. Figures 9, 10. M. pubescens, abdomen, dorsal view and male palp, respec- tively. be properly paired with it. Figure 8 shows certain features of the palpal tarsus. The specimens are simply labelled ‘‘ Amazon.’’ Simon (1895) stated that the type came from: ‘‘Brasilia: S. Paulo de Olwenea (de Mathan).’’ 1960 CERTAIN SPECIES OF MICRATHENA 5 MIcRATHENA PUBESCENS Simon, 1895 (Figures 9, 10) M. pubescens Simon, 1897 M. pubescens Petrunkevitch, 1911 M. pubescens Reimoser, 1917 M. pubescens Roewer, 1942 M. pubescens Bonnet, 1957 The female type is immature. The abdomen is hairy and sug- gests a close relationship with M. furcula (O. P. Cambridge) from Central America. There are no true spines but the posterior end of the abdomen is somewhat bifurcate. Simon (1895) fur- nished figures to show the right side of the abdomen and the bifureate posterior end. The immature female is accompanied by a mature male the palpal tarsus of which is shown in one position in Figure 10. Caution must always be exercised in matching the sexes in this genus and this male should be very carefully studied and compared with the growing number of different kinds of known males. The specimens are from Matto Grosso, Brazil. MIcRATHENA PUPA Simon, 1897 (Figures 11, 12) M. pupa Petrunkevitch, 1911 M. pupa Reimoser, 1917 M. pupa Roewer, 1942 M. pupa Bonnet, 1957 The length of the type is given as 8 mm. The form of the abdo- men and its spination are shown in Figure 11 and some of the features of the epigynum in Figure 12. The type female is from Ecuador. Simon apparently had a male associated with the female but Dr. Levi did not find it in the collection. Presumably it is lost and the original description does not give what we now consider to be the important diagnostic male features. MicrkATHENA TOVARENSIS Simon, 1897 (Figures 13, 14) M. tovarensis Petrunkevitch, 1911 M. tovarensis Reimoser, 1917 M. tovarensis Roewer, 1942 M. tovarensis Bonnet, 1957 6 BREVIORA No. i222 The length of the female type is given as 7.8 mm. The oeneral appearance of the abdomen and its spines, as seen in dorsal view, are shown in Figure 13. The appearance of the epigynum as shown in Dr. Levi’s drawing is given in Figure 14. The type is from Venezuela. The male is unknown aH A ie 15 External Anatomy of Micrathena Figures 11, 12. M. pupa, abdomen, dorsal view and epigynum, respectively. Figures 13, 14. M. tovarensis, abdomen, dorsal view and epigynum, respec- tively. Figure 15. M. ranthopyga, abdomen, dorsal view. MIcRATHENA XANTHOPYGA Simon, 1895 (Figure 15) M. xanthopyga Simon, 1897 M. ranthopyga Petrunkevitch, 1911 1960 CERTAIN SPECIES OF MICRATHENA ~ M. xanthopyga Reimoser, 1917 M. xanthopyga Roewer, 1942 M. xanthopyga Bonnet, 1957 The type is an immature female whose general appearance in dorsal view is shown in Figure 15. Simon (1895) published a figure of the type viewed from the right side. This figure shows four pairs of spines instead of three, as shown in Dr. Levi’s drawing. The type is from Venezuela. The male is unknown. BIBLIOGRAPHY BoNNET, PIERRE. 1957. Bibliographia Araneorum. Tome II (3me partie:G-M). Les Artisans de L’Imprimerie Douladoure, Toulouse. PrETRUNKEVITCH, ALEXANDER. 1911. A Synonymie Index-Catalogue of Spiders of North, Central, and South America, ete. Bull. Amer. Mus. Nat. Hist., vol. 29, pp. 1-809. REIMOSER, EDUARD. 1917. Die Spinnengattung Micrathena. Verh. Zool. Bot. Ges. Wien, vol. 67, pp. 73-160, pls. 1-9, figs. 1-31. RoEWER, C. FR. 1942. Katalog der Araneae. Vol. 1, Bremen. SIMON, EUGENE. 1895. Histoire Naturelle des Araignées. Vol. 1, Pts. 3-4. 1897. Descriptions d’Espéces de 1’Ordre des Araneae. Annales Société Entomologique de Franee, vol. 65, pp. 465-510. eet A : re Apts te ay es oe aft 4 or ¢ wvivihh we pity ire Guava ial ¥en a eeaeapan Hrd A NS sat | TIA coos ime 2M Sdalgien. wath, 1 Fgh oolulng? =iiol nel Yee aes "yc ' Ra a i ae ei ve. sea) uy! : . © +04 GOERS fr el ; a bilan te ‘ ; we an 8 oo i » = / 7 gsi a fan j— 7 i i ~ ehbal snrogemgy fe ..taey] : om iw snd digo yak. Ate is " i , = ’ + dyna, serene i vraag anigtt Jae, Tiay £ 3 & 7 US, nai OF 4 agli ighant ia i prea ares un cP ah yi nS Vid Mel! : bj (My ehaieneapenth HE ‘Dae! wa yah ue tet ? i x AD ¥ fe - - ae) vant, ee ae BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. Marcu 14, 1960 NUMBER 123 ALEPISAURUS BREVIROSTRIS, A NEW SPECIES OF LANCETFISH FROM THE WESTERN NORTH ATLANTIC By Ropert H. Gipss, JR. Department of Biology, Boston University Exploratory fishing for tunas in the western North Atlantic by the Fish and Wildlife Service vessel DELAWARE has re- vealed the presence of considerable numbers of lancetfishes, genus Alepisaurus. Through the courtesy of Mr. James L. Squire, I have been privileged to accompany most of these cruises. On the first two, the presence of Alepisaurus was recorded and stomach contents sampled, but only two specimens were saved. During the third cruise, it became apparent that two morphological types were represented, and from then on an attempt was made to measure and make counts on all possible specimens, and to pre- serve a large sample. At first the possibility was entertained that the differences might be due to sexual dimorphism. Gonads were therefore examined on all preserved specimens and on a large number of fresh ones. The appearance was almost exactly the same in all specimens, indicating that sexual dimorphism was probably not a factor. This has been borne out by histological studies of the gonads, which lead me to the rather surprising conclusion that both morphological types, which I am now certain represent valid species, are hermaphroditic. A study of nearly all type specimens and of all original de- scriptions leads to the conclusion that all previously described Atlantic species are conspecifie with A. ferox Lowe. The second form is described here. bo BREVIORA No. 123 ALEPISAURUS BREVIROSTRIS, Sp. NOV. (Figures 1-2) Holotype. U.S. National Museum 186197, 682 mm. in standard length when fresh, 684 mm. preserved; taken on longline with 20-fathom buoy lnes by the M/V DELAWARE at 38° 49’ N, 64° 02’ W, on September 13, 1957. Hl 4 HH Hid Us kif J la HH Ute, Jia. i me Figure 1. Left lateral view of Alepisaurus brevirostris (top), and A. ferox (bottom ). Paratypes have been distributed to the Museum of Compara- tive Zoology, Academy of Natural Sciences of Philadelphia, Cornell University, University of Miami Marine Laboratory, Tulane University, Scripps Institution of Oceanography, Stan- ford Natural History Museum, Museum of Zoology University 1960 ALEPISAURUS BREVIROSTRIS 3 of Michigan, California Academy of Sciences, British Museum (Natural History), Museum National d’ Histoire Naturelle Paris, and Museu Municipal do Funchal. Diagnosis. A dark-hued species of Alepisaurus with a grad- ually arcuate dorsal fin profile, the dorsal origin well forward of the rearmost margin of the operculum, a short head (6.5 or more in standard length), and a short snout (2.5 or more in head). Description. Dorsal fin high, originating over the middle of the opercle, its longest ray (about number 25-30) about three times the greatest depth of the body; its rays flexible, easily bent and broken, not branched, joints difficult to observe, though present. Leading dorsal ray thickened, its anterior edge finely serrated. Anal fin highest at the second and third rays, the rays branched distally. Pectoral fin pointed, its middle rays longest ; first ray thickened, serrated anteriorly, unbranched, the rest branched and obviously jointed. Pelvie fin also pointed, the middle rays longest, the first ray thickened, serrated anteriorly, and unbranched, the remaining rays jomted and_ branched. Caudal fin strongly forked, with eight procurrent rays in each lobe, ten upper principal rays and nine lowers; uppermost prin- cipal rays considerably elongated in some specimens. Snout inclined downward more sharply than the rear of the head, its length more than 2.5 times in head length. Nostrils a little less than halfway from tip of snout to anterior edge of orbit, the anterior opening round, the posterior crescent-shaped. Upper and lower jaws subequal. Upper jaw with a row of many small thin teeth on the premaxilla, one or two large fangs on anterior palatine, about three smaller fangs on the rear of the same bone, followed by about 7-10 low, triangular teeth. Lower jaw with an anterior large fang, followed by about 10 small caniniform teeth, one to three large fangs, and about 10-15 low, triangular teeth. No teeth on poorly developed tongue. Two patches of pharyngobranchial teeth. Gill arches with 3-6 upper, 0-1 middle, 17-23 lower groups of low, spinous rakers, totaling 23-28 groups. Branchiostegals mostly 7. Eye about 5 in head length, with vertical adipose eyelids anteriorly and posteriorly. Preopercle smooth. Opercle large, sculptured with lines radiat- ing from its antero-dorsal corner; subopercle also with striae 4 BREVIORA No. 123 radiating from its anterior point ; interopercle apparently absent. Interorbital space bounded by prominent parietal ridges, the area forming a flat to slightly concave surface, gradually widen- ing posteriorly. Body elongate, its greatest depth, at level of pectoral fins, about 12 in standard leneth. A low lateral keel occupying most of the rear half of each side. Lateral-line pores opening along the keel and continuing forward beyond it; lateral line on head forming prominent supraorbital, suborbital, and preoperculo- mandibular systems; supratemporal apparently lacking. Anus posterior to pelvie insertion by less than half the length of the pelvie fins. Fin-ray counts and morphometric data are given in Tables 1 and 2. Coloration. Body iridescent brownish-black dorsally, becom- ing eradually lighter laterally. Region above lateral line liberal- ly sprinkled with both large and small melanophores, many of the former ocellated. Below the lateral ne, and particularly on the belly, many small melanophores present. General coloration decidedly dark in comparison with A. ferox. Lateral keel black. Dorsal fin membrane iridescent black, often with a horizontal row of white spots a short distance above base. Other fins, including adipose dorsal, black. Head dark above, becoming lighter ventrally. Abdominal cavity marked externally by alter- nating light and dark bands, the light ones representing strips of musele, between which the dark peritoneal lining shows through. Visceral Anatomy. Peritoneal lining black. Liver relatively small, covering only the most anterior portions of the stomach and intestine. Stomach black, highly distensible, forming a long, blind sac. Intestine arising at the anterior end of the stomach, continuing straight, without bends, to the anus, divided at about one-third of its leneth into anterior thick-walled and posterior thin-walled portions. Kidneys occupying the entire leneth of the body eavity, lying retroperitoneally along the ventral side of the vertebral column. Ureters enter a thin-walled urmary bladder which extends about from the level of the pelvic insertion to the anus. Gonads consisting of a prominent pair of elongate, con- tinuous ovaries, which occupy the posterior third of the body 1960 ALEPISAURUS BREVIROSTRIS 5 cavity above the intestine, and a pair of thin testes, almost in- visible, lying in the dorsal groove formed by the two ovaries. The ducts of the ovaries, and presumably also those of the testes, join the urinary bladder and open by a urogenital pore immediately behind the anus. Swimbladder absent. Related Species. Other than A. brevirostris, the only recog- nized Atlantic species of the genus Alepisaurus is A. ferox Lowe. The two species are distinguishable by many characters. The most trenchant ones are shown in Table 3, and may be visualized in Figures 1 and 2. In addition to these, many less-perfect ones are demonstrable. Characters associated with head length show significant differences (snout to dorsal origin, snout to pectoral insertion, see Table 2). The pectorals average about one-fifth of the standard length in A. brevirostris and are slightly shorter in A. ferox. The eye is relatively larger in A. brevirostris, doubt- less correlated with the shorter snout. Meristic characters show consistent modal differences: dorsal rays most commonly 42-45 in A. brevirostris, 39-42 in A. ferox; anal rays usually 14-15 vs. 15-17; pelvic rays 13-14 vs. 14-15. Alepisaurus is common in the Pacific, but at present it is not possible to ascertain the species. I have examined seven Pacific specimens in the U.S. National Museum and find them extremely similar to, if not conspecific with A. ferox (see Table 1). The only disconcerting element was the low number of dorsal rays in four specimens, which indicates at least some degree of dif- ferentiation. I have seen no examples of a form resembling A. brevirostris from the Pacific. Young Specimens. Five specimens, 85.0 to 190.5 mm. standard length, present a confusing array of characters which defy posi- tive identification. Tables 1 and 2 show for these specimens many characters within or beyond the ranges displayed by adults of both A. brevirostris and A. ferox. There is obviously a great change between 200 and 500 mm. in the relative proportions, the caudal end in particular increasing proportionally greatly in this time. Presumably the shape of the head also changes, as these five are all extreme, even to A. brevirostris, in having short heads and snouts. [I am inclined to eall them all A. ferox, but with considerable doubt. BREVIORA IN@, 12 ght). a 1. ferox () Ps eae sani (eft), and “LS sti “eEviro: ‘ight. to the 1 a il side « Do rete Sear oo) ee Nw Vs WAS ? Figure 2. Photographs showing melanophore patterns of A. bi ~I 1960 ALEPISAURUS BREVIROSTRIS Mensural Discrepancies. Counts and a few selected measure- ments were made at sea on most of the specimens which came aboard the M/V DELAWARE. A later check has shown that the measurements cannot be used in conjunction with others taken on preserved specimens. Among the 17 preserved speci- mens of each species used in the descriptions here, were 11 A. ferox and 16 A. brevirostris which were also measured at sea when fresh. In these, the measurements of standard length and head length were consistently less in preserved specimens, In A. ferox, the standard length of one specimen was 0.7 per cent greater, the others 1.0-16.2 per cent less, averaging 6.6 per cent less. The head lengths were 3.4-6.6 per cent less, averaging 5.1 per cent. The corresponding figures for A. brevirostris were: standard length 0.3 and 1.1 per cent greater in two, others 2.0- 13.1 per cent less, averaging 4.5 per cent less; head length 0.5-2.8 per cent greater in three, 0.5-9.2 per cent less in 13, averaging 2.7 per cent less. Synonymy. Previous descriptions of Alepisawrus leave much to be desired. Eight nominal species have been described. I be- lieve all these descriptions refer to a single, perhaps polytypic, species. These names are discussed below. Alepisaurus ferox Lowe, 1833. The original description was made from two specimens from Madeira (Lowe, 1835a). In all important characteristics the written description is nearest A. ferox as recognized in the present study, but the accompanying drawing shows an arcuate dorsal fin profile, an absolute character of A. brevirostris. The matter was further complicated by a description and drawing of a third specimen from Madeira (Lowe, 1835b) which shows a dorsal fin profile characteristic of A. ferox. N. B. Marshall has kindly examined two types in the British Museum. His observations leave no doubt that both are A. ferox as I understand it. In the specimen labeled ‘‘TYPES!’’ (no registered number) the standard length is 1125 mm., head length (to tip of lower jaw, as upper is damaged) 191 mm. (17%), dorsal rays 39 or 40, anal 17, pectorals 14. In the specimen labeled ‘““SYNTYPE”’ (number 1852.9.13.98) the standard length is 1225 mm., head length 206 mm. (17%) ; snout 90 mm. (44% of head); dorsal 41, anal 17, pectorals 15. In both, the / / 8 BREVIORA No. 128 dorsal origin is over or shehtly behind the posterior edge of the operculum and no large, ocellated melanophores are present ; both are larger than any A. brevirostris I have seen. The name Alepisaurus feror is thus reasonably established for the long- snouted species. Alepisaurus azureus Valenciennes, 1849. No type is extant, but the following parts of the original description strongly sug- gest A. ferox: ‘‘. . . la dorsale est d’égale hauteur jusque vers le trentiéme rayon . . .’’? (Cuvier and Valenciennes, 1849: 531) ; dorsal rays 38, anal 16. The pectoral count of 10 is presumably in error. The length of 5 feet 3 inches is larger than any known A. brevirostris. The description was based on a specimen from the Canary Islands. Alepisaurus richardson Bleeker, 1855. Based on the deserip- tion by Sir John Richardson (1844) of a head from Van Diemens Land. The drawing of the head shows a snout most like A. ferow. Alepisaurus altivelis Poey, 1861. It is difficult to be certain of this description. The anterior rays are all said to be the same height, the posterior ones decreasing rapidly ; dorsal rays 40, anal 17, pectorals 16; all most like A. ferox. The pelvic count of 13 1s presumably in error. Based on a Cuban specimen. Alepidosaurus borealis Gill, 1863. Based on a head, dorsal, caudal, and pelvie fins from a Pacifie specimen. I have examined this specimen and find it close to A. feroxr except for a dorsal count of 35. The pelvic count of 13 is apparently an error. The stated snout/head ratio of 2/5 definitely precludes A. brevi- rostris. Alepidosaurus serra Gill, 1863. Deseribed from a head, caudal, and pelvie fins of a Pacific specimen. I can detect no significant differences between Gill’s descriptions of this species and of A. borealis. He places much emphasis on opercular sculpturing, which seems to be quite variable and not a good specific charac- ter. Again the pelvie count of 13 is presumed to be an error. The distance from eye to snout, stated as 2/5 of head length, excludes A. brevirostris. Alepidosaurus poeyi Gill, 1863. Described on the basis of drawiness (which I have not seen) of the second specimen from Cuba mentioned by Poey (1861) in his description of A. altivelis. 1960 ALEPISAURUS BREVIROSTRIS 9 The specimen was described as having the first dorsal rays be- coming longer, the fourth very long, rays 6-24 high and equal. This suggests A. feror, as does the dorsal ray count of 41. Poey’s presumably erroneous pelvic count of 13 is restated. Alepidosaurus aesculapius Bean, 1883. I have examined the type and find it similar to A. ferox. The dorsal rays could not be counted accurately, but Bean (1883) gave 39; anal rays 16, both A. ferox characters. The snout length (41% of head length) rules out A. brevirostris with absolute certainty. Described from a Pacific specimen. Distribution of Alepisaurus in the Atlantic. Among the speci- mens examined and definitely identified, the most southerly was one A. feror from off southern Puerto Rico, now in the Museum of the University of Miami Marine Laboratory. Several of the same species from the Gulf of Mexico are in the U. S. National Museum, and specimens from the Gulf of Maine (La Have Bank the most northerly) were seen at the Museum of Comparative Zoology. The remaining positively identified specimens of I & 6 42° @ ml jb il 1, te) 3} A. ferox 2 B10 WO ag 7 2b BF al ae) 7 Pacific (USNM ) 222 1 1 SS Small 1 HW op Anal Rays Pectoral Rays Gill Rakers 183 V8 aly a ali ash 11) aes awe als TMG) } BEL sy A) Arh ts} 249) A. brevirostris i 8) IB} al i ales 10) ms 46 a al 8} A. ferox Wz) Pal Wey ala oy SO al Dy t4e Ae 23 ill Pacific (USNM ) 6 1 4 3 Small 3 2 Zell! 1960 ALEPISAURUS BREVIROSTRIS 13 Table 2. Proportional dimensions of Alepisaurus expressed as per cent of standard length. Range, with mean in parentheses. Based on seventeen adult specimens of each species and five small specimens. small 7.5) 85.0-190.5 (144.2) brevirostris 551-894 (670.6) ferox Standard length, mm. Per cent of standard length Snout to anal origin Snout to pelvic insertion Snout to pectoral insert Snout to dorsal origin Head length Greatest depth Caudal peduncle depth Pectoral length Pelvic length Anal base Anal height Per cent of head length Snout to fleshy orbit Snout to bony orbit Fleshy orbit length Bony orbit length Least interorbital width Snout to anterior nostril Number of gill raker groups (total) 77-83 (79.5) 42-50 (46.3) 13-16 (14.7) 9.4-13 (11.4) 12-16 (14.5) 7.2-10.4(8.5) 1.9-3.2(2.4) 14-20(16.3) 7.2-12.4(9.5) 8.5-11.4(9.8) 5.9-7.9(6.9) 31-37 (34.5 ) 26-31(28.4) 19-23 (20.9) =O (2 (cs) 15-20(17.5) 14-16(14.9) 23-28 (25.4) 31-1088 (77 76-88 (80.1) 43-53 (47.5) 17-23 (19.4) 16-22 (17.9) 16-23 (18.6) 8.0-12.5 (9.5) 2.3-4.0(2.8) 17-24(20.5) 7.5-10.4(9.0 ) 9.2-13.8 (10.7) 7.4-10.3 (8.6) 41-46 (43.3 ) 35-42 (38.1) 13-20(17.8) 17-28 (23.4) 14-18 (16.2) 21-24(22.4) 23-29 (25.8) 80-81 (80.0) 53-58 (55.3 ) 29-95 (24.0) 21-25 (23.2) 23-30 (25.6) 13-16 (14.4) 2.7-4.0:(3.4) 15-19 (17.3 ) 7.4-8.5(7.9) OF 5 asi Glee) 6.5-9.1(7.8) 36-41 (38.4) 27-37 (31.4) -2.8:( 26.2) -36 (33.4) Sala) £18172 2) -27(26) 1/4) BREVIORA No. 1238 Table 3. Principal differential characters of A. brevirostris and A. ferox A. brevirostris Coloration dark, more and larger melanophores, many of them ocellated (Mig. 2) Dorsal fin gradually arcuate, without free anterior rays (Mig. 1) Irregular horizontal row of white spots often present on dorsal membrane Dorsal origin well in advance of rear margin of the operecu- lum Head 6.5 or more in standard length Snout 2.5 or more in head leneth A. ferox Coloration heht, fewer, most- ly small melanophores, few or none ocellated (Fig. 2) Dorsal fin with several ante- rior rays elongated, free from membrane; rest of fin about equal in height until sudden posterior drop, or slightly higher before drop (Fig. 1) No white spots on dorsal mem- brane Dorsal origin about level with the rear margin of the opereu- lim Head less than 6.5 in standard leneth Snout less than 2.5 in head length BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. Marcu 15, 1960 NUMBER 124 ANISIAN AMMONOIDS FROM MALAYA By BERNHARD KUMMEL Fossiliferous marine strata are sparsely represented or known from Malaya. Even the Triassic system which is one of the better known systems to yield fossils is represented by extremely small faunas consisting mainly of peleeypods, and only indeterminate ammonoids have been reported by Newton (1923, 1925). The first discovery of determinable ammonoids was reported by Savage (1950) from mudstones near Kuala Lipis, Pahang. The initial collections were submitted to L. EF. Spath of the British Museum (Natural History) who made the following report (in Savage, 1950): ‘‘Quite a number of common Middle Triassic (Anisian) genera can be recognized in the collection, including Paraceratites (dominant), Sturia, Ptlychites, and Acrochordi- ceras, so that the age of the assemblage is the trinodosus zone. Specific identifications would be more difficult but are unneces- sary; Paraceratites trinodosus (Mojsisovies) and such close allies as the Himalayan Ceratites thuilleri (Oppel) and P. winterbot- toni (Salter) are probably all represented.’”’ The rarity of Triassic ammonoids in Southeast Asia, lying as it does at the eastern end of Tethys between the richly fossil- iferous Triassic horizons of the Himalayas and the island of Timor, warrants a more substantial record of these faunas than Spath was able to give. Through the kindness of Dr. M. K. Howarth of the British Museum (Natural History) all of the best preserved material from Kuala Lipis, Pahang, was loaned to the writer. Close examination of this small collection (29 specimens ) showed that Spath’s conclusions as to genera present and age assignment are correct in spite of the rather poor preservation. 2 BREVIORA No. 124 The fauna contains the following species : Paraceratites trinodosus (Mojsisovies ) Sturia sansovinit Mojsisovies Acrochordiceras sp. indet. Ptychites sp. indet. Data on the geographic and geologic occurrence of this fauna can best be quoted from Savage (1950, p. 76) : ‘One of the newly recorded areas is some 10.5 miles south-south-west of Kuala Lipis on one of the branches of the Sungei (= River) Tua, where it flows through Budu Estate (approximately lat. N. 4°02/30”, long. EH. 102°00’15”; Malayan Survey Department Topographi- cal Sheet No. 2 0/13). The rocks are mudstones, rarely laminated sufficiently to be called shales. They are fairly homogeneous, slate grey, almost black and carbonaceous; but some slightly sandier strata (muddy siltstones) weather to a pale buff or brown. The beds show minor flexures but in the main strike 40°-220° and dip to the north-west at angles of about 40°. They are strongly jointed along several directions, the two main joint systems being vertical and striking 60°-240° and 155°-335°. The beds are fossiliferous over a distance of at least 50 yds. in a horizontal direction normal to the strike, equivalent to a strati- oraphical thickness of about 100 ft.’’ No data are available with the specimens as to their precise position in the fossiliferous horizon. The species recognized in this small fauna are common forms widely distributed in Tethys and the general cireum-Pacifie region. Paraceratites trinodosus and Sturia sansovinii were originally established on Alpine specimens. The specimens as- signed to Ptychites and Acrochordiccras are too poorly preserved to enable specific identification but there is no doubt as to the generic assignment. The two previous records of Triassic ammonoids from Malaya are not as satisfactory. A small fragment of an ammonoid was recorded by Newton (1923, p. 302, pl. 9, fig. 29); it is quite indeterminable. The specimen came from argillaceous sandstone at Mount Faber, Singapore, and was reported to have a ‘‘de- pressed whorl with indications of straight ribs and furrows con- necting with some well-separated knob-like tubercles situated within a short distance of the inner margin.’’ Newton likewise 1960 ANISIAN AMMONOIDS FROM MALAYA 3 did not beheve the specimen was determinable but thought that it resembled the genus Balatonites. There does not appear to be any justification for this suggestion. The second record of am- monoid remains from Malaya is also in a paper by Newton (1925) on a small Upper Triassic fauna from the Province of Kedah. The first of the two specimens available to Newton was cited as Ammonites sp. indet. *‘A’’ and he suggested that it may be referred to a form of the Arcestidae. It seems more likely that it isa Juvavites or possibly an Anatomites but the specimen lacks the characteristic constrictions of these genera (Spath, 1951, p. 106, footnote). The second specimen was listed by Newton as Ammonites sp. indet. “‘B’’ and suggested a resemblance to Balatomtes. Spath (1951, p. 15) thought that this specimen might be Hannoceras nasturtium (Mojsisovies). In the adjoming regions of southeast Asia only Indochina has yielded a large and varied fauna of Triassic ammonoids which, however, are generally not well preserved. The literature on the stratigraphy and faunas of this area is very large and need not be reviewed here. A brief summary can be found in Saurin (1956). The only other really new discovery of Triassic am- monoids in southeast Asia has been made in Thailand where Anisian and Karnian faunas are now known. The geology of this Triassic region has been described by Pitakpaivan (1955) who includes a preliminary list of the species present, identified by Kummel. Full description of this fauna will be published shortly. In the following description of the species from Kuala Lipis the extensive svnonymies for Paraccratites trinodosus and Sturia sansovini have been omitted; essentially complete synonymies can be found in Diener (1914a) and Kutassy (1933). SYSTEMATIC DESCRIPTIONS Family CERATITIDAE Mojsisovies, 1879 Genus PARACERATITES Hyatt, 1900 PARACERATITES TRINODOSUS (Mojsisovies) Plate 1, figures 3-6 The collection contains no less than twenty erushed and in- complete specimens that can be assigned to the well known 4 BREVIORA No. 124 Paraceratites trinodosus (Mojsisovies). Allowing for the general faulty preservation, these specimens agree well in most details with the type of this species and with other specimens assigned to it. In his preliminary examination of the fauna, Spath (in Savage, 1950, p. 76) considered that in addition to Paraceratites trinodosus the fauna also contained P. thuilleri (Oppel) and P. winterbottomi (Salter). These are very closely allied forms occurring with P. trinodosus in the Himalayas. However, con- sidering the poor preservation of the specimens, it seems that a more conservative approach is desirable and I am recognizing only the better known and more widely distributed P. trinodosus. This species is particularly widespread in the Alps, Balkans and the Middle Kast. It is hkewise recorded from the Himalayas and Nevada. Identical or closely related species are also known from Japan. This species gives its name to the upper Anisian trinodosus zone. veposittory. BMNH —C 55672, C 55673, C 55674, C 55675 (figured specimens); C 55653, C 55654, C 55655, C 55657, C 55658, C 55661, C 55662, C 55663, C 55666, C 55667, C 55668, C 55670, C 55671, C 55676, C 55678. Family PTYCHITIDAE Mojsisovies, 1882 Genus PTYCHITITES Mojsisovies, 1875 PTYCHITES sp. indet. Plate 1, figure 7 The collection contains three crushed and fragmentary speci- mens that without question belong in Ptychites but identification at the specific level is not possible nor advisable. The most complete specimen is actually only the impression of one side of a conch. The illustration on Plate 1, figure 7 is of a latex cast of this impression. It shows the funnel-shaped umbilicus, broad- ly arched lateral areas, and the radial ribs — all features which are very characteristic of the genus Ptychites. Ptychites is known from Middle Triassie¢ strata throughout the world. It likewise includes a very large number of species based largely on differences in shape of the conch, character of ribs, degree of involution, and details of the suture. In the Himalayan 1960 ANISIAN AMMONOIDS FROM MALAYA 5 Muschelkalk, Piychites, of the group of P. rugiferus (Oppel) to which these Malayan specimens most likely belong, is one of the most abundant forms present, being represented by seven species (Diener, 1895, 1907). In southeast Asia the record of Ptychites is very fragmentary and represented mostly by indeterminate species. Even the rich Middle Triassic faunas of Timor appear to have only one species, P. amarassicus Welter (1915; Arthaber, 1928). The genus is, however, also present in Thailand and Indo- china. It is also known to be present in Japan and New Zealand. Repository: BMNH — C 55659 (figured specimen) ; C 55664, C 55656. Genus STURIA Mojsisovies, 1882 STURIA SANSOVINH Mojsisovies Plate 1, figure 2 The most easily recognizable species in the collection is this strigate form which is widely distributed throughout Tethys. The specimen consists only of the impression of slightly more than one third volution of one side of a whorl; no suture or whorl section is preserved. In spite of this fragmentary preser- vation the ornamentation is so characteristic that there is no reason to doubt its identity with this species. The ornamentation consists of broad, flattened strigations separated by broader, rounded grooves which bear a fine spiral line down the center. The pattern of ornamentation on the Malayan specimen is identi- cal to the fine specimen from the Shalshal Cliffs in the Himalayas, figured and described by Diener (1895, pp. 61-62, pl. 15, figs. la, b). In his description of the Himalayan specimen Diener was quite emphatic as to the identity of his form with the type from the Alpine Middle Triassic. In this conclusion he had confirma- tion from Mojsisovics who also examined the Himalayan speci- men. Sturia sansovini is known from Anisian and Ladinian strata at many localities in the Mediterranean region, Bibliographic citations to these can be found in Diener (1915a) and Kutassy (1933). The distribution in the region of eastern Tethys and in the cireum-Pacifie region is not so well known and is of special 6 BREVIORA No. 124 interest here. This species is the only form of Sturia from the Himalayas proper in the so-called Himalayan facies, where Diener (1895, 1907) has recorded specimens from the upper Muschelkalk at the Shalshal Cliff and at Spiti. However, in Tibet in some of the exotic blocks of Malla Johar near Chitichun Peak No. 1 (17,740 ft.), Diener obtained a specimen of Sturia which he identified as Sturia monogolica (Diener, 1895, p. 113, pl. 29, fig. 4). This form is quite distinct from other species of Sturia in its open umbilicus and the suture, characterized by long, slender, pyramidal saddles. At a later date Diener (1916) erected the genus Psilosturia with S. mongolica as the type species. Indeterminate species have been recorded from Middle Triassic horizons in upper Thailand (Kummel, in Pitakpaivan, 1955). These particular forms are small, poorly preserved, and crushed specimens whose relationship to S. sansovini is impossible to determine. Amone the numerous Middle Triassic faunas de- scribed from Indochina, Sturia has as vet not been recorded. Welter (1915) records S. ef. sansovinii from a Ladinian horizon on Timor based on a fragmentary specimen. Sturva japomca Diener (1915b, pp. 18-20, pl. 6, figs. 1-2) is based on a highly distorted specimen from Middle Triassic forma- tions at Inai, Japan. It is quite similar to S. sansovinii differing in minor features of the suture and character of the strigations. Sturia sansovinu is thus found widely distributed throughout the Tethyvan geosyncline where it occurs in strata of Anisian and Ladinian age. Repository: BMNH — C 55669 (figured specimen ). Family ACROCHORDICERATIDAE Arthaber, 1911 = Genus ACRCCHORDICERAS Hyatt, 1877 ACROCHORDICERAS sp. indet. Plate 1, figure 1 A single, large, crushed and elongated specimen can be as- signed to the genus Acrochordiceras but its poor and incomplete preservation prevents determination of its specific relationship. The whorl sides bear strong radial to shehtly curved sharp ribs. 1960 ANISIAN AMMONOIDS FROM MALAYA Ff Some of the ribs begin at the umbilical shoulders where they increase in height forming somewhat of a tubercle beyond which they bifureate. Other ribs lack the umbilical protuberances and are slightly less prominent. The poor preservation prevents de- termining the pattern of alternation of these two types of ribs. The conch was no doubt slightly evolute but the shape of the whorl section is not possible to determine nor is any part of the suture preserved. Close comparison of this Malayan specimen with the known species of Acrochordiceras is not very satisfactory but one fea- ture is notable — that is, the rather sharp ribs on the Malayan form. The extent to which these sharp ribs may be due to the deformation of the specimen is hard to determine, however. In most species of Acrochordiceras the ribs tend to be rounded and in some cases broadly rounded. Even though specific comparisons are not possible there is no question of the generic assignment of this form. The genus Acrochordiceras is widely distributed in the Tethy- an belt from the Alps to Timor and is likewise known from a number of localities in the cireum-Pacifie region. Repository. BMNH — © 55660 (figured specimen). REFERENCES ARTHABER, G. V. 1928. Ammonoidea Leiostraca aus der oberen Trias von Timor. 2. Nederl. Timor Expedite 1916 onder leiding von Dr, H. G. Jonker. Uitgegeven door Dr. H. A. Brouwer. IV. Jaarb. Mijnw. Neder. Ind., vol. LV, no. 2, pp. 1-174, pls. 1-20 (1926). DIENER, CARL 1895. Himalayan Fossils. The Cephalopoda of the Muschelkalk. India Geol. Survey, Palaeont. Indica, ser. 15, pt. 2, pp. 1-118, pls. 1-31. 1907. Fauna of the Himalayan Muschelkalk. India Geol. Survey, Palaeont. Indica, ser. 15, pt. 5, pp. 1-410, pls. 1-17. 1915a. Fossilium Catalogus. I. Pt. 8. Cephalopoda triadica. 369 pp. Berlin. 1915b. Japanische Triasfaunen. Denkschr. Akad. Wiss. Wien., vol. 92, pp. 1-30, pls. 1-7. 1916. Einige Bemerkungen zur Nomenklatur der Triascephalopoden. Centrabl. Min. Geol. Paliont., pp. 97-105. 8 BREVIORA No. 124 Kutassy, A. 1933. Fossilium Catalogus. I. Animalia. Pt. 56, Cephalopoda triadica I1., pp. 371-832, Berlin. Newton, R. B. 1923. On marine Triassic shells from Singapore. Ann. Mag. Nat. Hist., ser. 9, vol. 12, pp. 300-321, pl. 9. 1925. On marine Triassic fossils from the Malayan Provinces of Kedah and Perak. Geol. Mag., vol. 62, pp. 76-85, pl. 3. PITAKPAIVAN, KASET 1955. Occurrences of Triassic Formation at Mae Moh. Royal Dept. Mines, Bangkok, Rept. Investigations No. 1, pp. 1-11, pls. 2-4, map. SAURIN, E. 1956. Lexique Stratigraphique International, vol. 8, Asie, Fasc, 6a. Indochine, pp. 1-141. SAVAGE, H.: E. F. 1950. Triassic fossils from near Kuala Lipis, Pahang (Malaya). Colonial Geol. and Min. Resources, vol. 1, no. 1, pp. 76-77. SpatrH, L. F. 1951. Catalogue of the fossil Cephalopoda in the British Museum (Natural History). Part 5, the Ammonoidea of the Trias (D). London, pp. 1-228. WELTER, O. A. 1915. Die Ammoniten und Nautiliden der ladinischen und anisischen Trias von Timor. Paliont. von Timor, vol. V, pp. 71-136, pls. 83-95 Explanation of PLATE The specimens illustrated on this plate are from mudstones of Anisian age from near Kuala Lipis, Pahang, Malaya. They are deposited in the 3ritish Museum (Natural History), London. Figure 1. Acrochordiceras sp. indet. BMNH — C 55660. X 0.5. Figure 2. Sturia sansovinii Mojsisovics. BMNH — C 55669. X 0.5. Figure 3-6. Paraceratites trinodosus (Mojsisovies) BMNH — € 55672 — CO! NO, Oo: ale Figure 7. Ptychites sp. indet. BMNH — C€ 55659. X 1. PLATE BREVIORA Museum of Comparative Zoology eS ee ee eee ee eee CAMBRIDGE, Mass. May 27, 1960 NuMBER 125 THE LUMINOUS ORGANS OF PROCTOPORUS (SAURIA, REPTILIA) — A RE-EVALUATION By Wriuuarp D. Rotu Department of Anatomy, Harvard Medical School, Boston, Mass. and CARL GANS Department of Biology, The University of Buffalo, Buffalo, N. Y and Carnegie Museum, Pittsburgh, Pa. ” INTRODUCTION The herpetological literature contains two reports describing the first luminous organs in a terrestrial vertebrate. The two papers discuss identical specimens of the Trinidad lizard Procto- porus shrevei Parker. Sanderson (1939, and observations cited by Parker, same date) claimed that light was emitted by black bordered ocelli on the sides of a male, and Parker (1939) sup- ported this on the basis of his histological examination of the preserved animal. No new observations have been published since that time, but a number of workers have commented upon the original observations. Thus Pope (1955, p. 306) remarked that ‘‘other te1ids have spots somewhat like those of P. shrevei, so it is highly probable that they, too, can light up.’’ In contrast to this, Harvey (1952, p. 494) in his monograph on bioluminescence stated that he believed ‘‘all reports of luminescence in higher vertebrates to be false or spurious due to reflection of light or infection by luminous bacteria.’’ The divergence of opinion on this interesting point prompted a re-examination of this question. The present paper reports a few additional field observations, and includes as well a detailed examination of the histological structure of the ocelli. Since Proctoporus shrevei is very rare, this re-examination had to be carried out on two related and superficially similar forms. Z BREVIORA No. 125 OBSERVATIONS ON LIVING SPECIMENS Parker (1939, p. 659) mentioned that Sanderson’s field notes contained only a brief reference to color pattern involving “‘five, black spots each containing a small, vivid white, sometimes luminous bead.’’? Parker further stated that Sanderson in con- versation informed him ‘‘that the animal was kept alive in captivity and could be stimulated to emit light from the lateral spots; the hght was of a pale greenish hue, similar to that pro- duced by the hands and figures of a luminous watch. Exeitement produced by flashing an intermittent beam of light on the lizard was found to be a very effective stimulus to ight production.”’ A more specific first-hand report was given by Sanderson (1939, pp. 41-48) in his popular, considerably amplified and shehtly different, account. When initially observed the lizard ‘‘turned its head away from me and both its sides lit up for a few seconds like the portholes of a ship.’’ ‘‘ After one brilliant display on the night of its arrival in camp it refused to shine with full brightness though the beadlike spots remained plainly discernible in a darkened box when the rest of the animal was invisible.’’ A ‘‘loud whistle, sudden winds, and flashes of light greatly agitated our lizard, causing it to switch on its ‘‘port- holes’’. .. . The light was much brighter the first time it was switched on after the animal had been quiescent for a period, and more especially after it (the lizard) had previously been sub- jeeted to intense illumination.”’ We have been able to obtain four sets of further observations on live specimens of the genus Proctoporus. Julian §8. Kenny (V. C. Quesnel, personal communication ) some time ago repeated Sanderson’s experiments on the original species (P. shrever) with entirely negative results. Kenny’s field notes also indicate that the lizards are diurnal and inhabit rela- tively open spaces on El Tueuche, Trinidad. Dr. Janis Roze (in litt.) states that a specimen of Proctoporus achlyens Uzzell (M.C.Z. 53128. later used for histological exami- nation reported herein) did not glow when placed in a darkened room after capture. He adds that exposure to ultraviolet light did cause the spots to shine faintly. The test was carried out in an incompletely darkened room, and the results seem to be open to some question. Harold Heatwole and Owen J. Sexton (personal communica- tion) performed a number of experiments at a field station in Venezuela. They tested one adult male of P. Juctwosus (Peters) 1960 THE LUMINOUS ORGANS OF PROCTOPORUS Ww and two adult males of P. achlyens for a period of one month. The spots of the first species were yellow and those of the second were red in life, both series of spots bleaching to white after formalin preservation. The specimens were repeatedly moved from light to dark environments and were observed at night. The animals were disturbed. No luminescent effect was ever noted. Ultraviolet illumination was not attempted. The most extensive series of observations on live animals was made in Ecuador by James A. Peters (in litt.). Specimens of Prionodactylus vertebralis (O’Shaughnessy ) and Neusticurus ec- pleopus Cope were observed while free in the field, durine the collecting process, and for several weeks in the laboratory. He reported that neither luminescence nor any other kind of heht could be noted in broad sunlight, dim or artificial light, or in the complete absence of light. No reflection could be noted under various types of lighting (sun, fluorescent and incandescent), in quiescent, active or deliberately disturbed animals. The evidence is most valuable because Peters was aware of the lizards’ reputa- tion and was deliberately testing the hypothesis of luminescence. SUPERFICIAL APPEARANCE AND PHYLOGENETIC DISTRIBUTION OF THE OCELLI The supposedly luminous ocelli (Fig. 1) are rather similar in the species of Proctoporus and Neusticurus here discussed. They are arranged in a single row along the side of the animal; each ocellus always shows a_ light-colored, sharply-defined, black- bordered, circular center. They may be restricted to adult males, with juvenile specimens and females showing only traces. There is usually a size decrease of the ocellar center posteriorly along the series. There may be a marked irregularity in the width of the black border. There is no correlation between the ocellar and the scale patterns. While sharplv-defined ocelli are commonly well developed only in Boulenger’s (1885, p. 332) telid group II, a check of the more than 130 species of telid lizards as well as forms of other families represented in the collection of the Museum of Comparative Zoology at Harvard College indicated that patterns with sharply contrasting Heght and dark colors are extremely common. Vv PIRATE 2 BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 19, 1960 NUMBER 127 NOTES ON THE CRANIAL ANATOMY OF NECROLEMUR By K. L. Stuons! anp D. BE. Russet ? INTRODUCTION The large number of well-preserved skulls of Necrolemur antiquus of the late Eocene Querey phosphorites of south central France allow for much more detailed study of cranial anatomy in this primate than is possible for most early members of the order. In spite of the fact that cranial osteology can be studied in great detail, views as to the taxonomie position of this primate, and of the allhed genera Microchoerus, Nannopithex, and Pseudoloris show considerable variance. Although not all of the same provenance, little dental varia- bility is evidenced in specimens of Necrolemur antiquus examined by us. In the course of this study, however, a number of differ- ences in position and size of basicranial foramina have been observed, which are in line with mutability of cranial foramina (in individuals of the same species) reported by other authors (see Edinger and Kitts, 1954). Also, in Necrolemur the proba- bility remains that known specimens differ considerably in age (from early Bartonian to late Ludian provincial ages, at least), but locality data are inadequate for precise age determinations. Comparison of upper dentitions in the M.C.Z. and Paris skulls has failed to show any dental basis for species distinctions among them. To date, the most detailed studies of the cranium of Necrolemur have been by Stehlin (1916) and by Hiirzeler (1948). Stehlin’s thorough and excellent description can scarcely be improved on, +Zoological Laboratories, University of Pennsylvania. *Muséum National d’Histoire Naturelle, Paris. y BREVIORA No. 127 but copies of this work are not as generally available as could be wished. Also, the implications of some of his early observa- tions seem to have been neelected in later literature. In some rather significant points, recent examination of more and dif- ferent skulls permits comments supplementary to his work. ACKNOWLEDGEMENTS The authors would like to take this opportunity to thank Drs. J.-P. Lehman, Curator of Fossil Vertebrates at the Muséum National d’Histoire Naturelle in Paris, and A. 8S. Romer of the Museum of Comparative Zoology at Harvard for generously eiving permission to publish on the specimens in their respec- tive charges. Preparation of the figures, by Miss Ellen Cole, was supported by a grant from the Wenner-Gren Foundation for Anthropological Research. ABBREVIATIONS In the absence of specimen numbers, the Paris Museum skulls of Necrolemur have been numbered 1 through 5 for convenience of reference. Abbreviations used in this paper are as follows: M.C.Z., Museum of Comparative Zoology at Harvard College. Montauban, Natural History Museum (Geological Collection), Montauban, France. Paris, National Museum of Natural History, Paris. CRANIAL CHARACTERS I. AUDITORY REGION Most of the information, published to date, regarding the com- ponents of the auditory bulla in Necrolemur comes from Mon- tauban 9, which has been discussed by both Stehlin and Hiurzeler. Even though this skull was prepared with considerable skill, the crystalline calcite filling was apparently confused with the very similar appearing bone in the region of the epitympanic recess. Consequently, the route of the stapedial artery across the tympanic cavity of the middle ear was lost just anterior to the fenestra ovalis. The extrapetrous portion of the Fallopian aque- duct was lost as well. Preparation of Paris 2 has revealed more details of the epitympanie region (Fig. 1). Exposure of the inside of the bulla in Paris 2 and 5 indicated primarily the lack of a free annular tympanic ring. Moreover, studies by Simons (in press) on a specimen of Necrolemur at the NECROLEMUR OF ANATOMY CRANIAL 1960 “NOITTITULIS TRULLI XoO is ): wey “yy twnrtioj,uoword “gq fuasue1osy promos “yg + Adoyre Tetipodeys “prog {Adoyre ALOZUOWMOId © Va ‘popRiossuxo Ydop otdoosoatojyg “G X “VPN YYstr JO AOLIoJUL ‘Z SLE :gyonponbe uvido[Rgq JO woryt0od snowjodeayxo JO ¢..f,, “yp tStpvao vajysou G cl T Huo ag ‘ i “| RURo Lv] Ow + Bpunjor Biyso 7*q ‘Suorpeiaoiqqy SSUNDUUD MNWAIOLII N 4 BREVIORA Now 127 British Museum (Natural History) demonstrate that the ecto- tympame element is tubular, and medially fused to the ventral bulla wall. Huirzeler’s evidence (1946:353: 1948:28) of the presence of a free ring, therefore, can no longer be accepted. The bone he identified as such had to be removed during prepa- ration to expose the carotid canal and thus cannot be re-exam- ined. In M.C.Z. 8879 the meatal tubes are reasonably well pre- served but one of the Paris skulls shows an even more complete osseous meatus. Together, these indicate that the ectotympanic (external to the bulla) is about as lone as the transverse diam- eter of the foramen magnum, curves slightly backward, and may be broadest at the external aperture. None of the fossil or recent Lemuroidea have this sort of meatus. Hurzeler (1948:27) cites M.C.Z. 8879 as not showing any evidence of a fused tympanie ring. Nevertheless, four and pos- sibly more transverse struts are exposed on the ventrolateral face of the right bulla of the Harvard skull. These bars are supports for the internal rim of the tubular ectotympanic. Also of interest is the fact that the anterior route of the promontory artery (true entocarotid) is apparently variable. In Montauban 9, it curves sharply anteromedially shortly after leaving the promontory of the petrosal. In Paris 2, this curving is much less accentuated. Some erushing is to be allowed for in the tympanic region of the latter specimen but the amount of curvature illustrated by Hiirzeler (1948, figs. 30 and 31) for Montauban 9 is not indicated in the Paris skull. It should be further noted that this bony tube does not lie in a horizontal plane in Montauban 9 and Paris 2, but slopes anterodorsally at an angle of about 45°. The stapedial artery, like the promontory artery, remained enclosed in a bony tube throughout its route within the bulla. Branching from the promontory artery just inside the carotid foramen, the stapedial artery curved dorsally, lying on and following the form of the petrosal promontory. It then passed anterior to the fenestra rotunda to the bottom of the fenestra ovalis. There it diverged laterally, crossing the fenestra ovalis, and continued anteriorly nearly parallel to the promontory artery (Fig. 1). The groove mentioned by Hiirzeler (1948 :31) is surely a remnant of the stapedial tube, as he suggested. The exit of the stapedial artery appears to be at the dorsoanterior base of the external auditory meatus. 1960 CRANIAL ANATOMY OF NECROLEMUR 5 The extrapetrous portion of the Fallopian aqueduct is also enclosed up to, or nearly up to, its exit at the stylomastoid fora- men. Its route lies lateral to the fenestra ovalis, just above the stapedial artery and continues posteriorly under the external semicircular canal. At this point, the tube forms a ‘‘T”’ giving rise to the small anterolateral opening and a larger posterome- dian branch. The former is a natural foramen. Damage to the latter region makes it impossible to say whether or not the more posterior branch continued as an enclosed tube to the stylo- mastoid foramen. In Paris 2, this foramen is single, in Paris 1, double. Without exposing the interior of the bulla in Paris 1 a possible connection between the branching Fallopian aqueduct and the double stylomastoid opening cannot be confirmed. That the anterolateral foramen of the ‘‘T’’ in Paris 2 could have given passage to the chorda tympani seems likely. Regarding the foramen designated FX, ‘‘Foramen von un- bekannter Bedeutung,’’ by Hiirzeler (1948 :fig. 28), it seems probable that this represents the opening of the inferior petrous sinus, as originally stated by Stehlin (1916 :1355). Contrary to the views of a few students, we find little distine- tion between Tarsius and Necrolemur in the major carotid rela- tionships, both inside and outside the bulla. The two genera appear to agree in those features of the carotid circulation which distinguish tarsiers from Malagasy lemurs, adapids and lorises (see Le Gros Clark, 1959:151). Location of the internal carotid foramen in Tarsius on the ventral surface of the bulla (instead of on the median wall) constitutes a slight difference from Necrolemur, but it should be stressed that placement of this foramen in the fossil species does approximate the situa- tion in Tarsius, being more ventral than in most, if not all, other prosimians. Furthermore, Necrolemur and Tarsius are alike in having both stapedial and promontory branches within the bulla, encased in bony canals or tubes, with the promontory division the larger. In typical Lemuriformes (Malagasy lemurs, adapines and notharctines), the carotid foramen has a quite dif- ferent location at the posteroexternal angle of the bulla and, inside it, the promontory division is very small (Gregory, 1920 :174-180). Lorises and the cheirogaleine lemurs differ also, in that the carotid divides outside the entotympanic and the main branch enters the skull through the foramen lacerum medium instead of going through the bulla. A middle lacerate foramen is not present in Tarsius and Necrolemur. The few differences be- tween these two genera to be observed in the auditory region 6 BREVIORA INO RAZ 7 seem best understood with reference to the effects of the anterior shifting of the foramen magnum and greater inflation of the anterointernal part of the bulla in Tarsius. Some primitive fea- tures are also to be seen in the Eocene form. For instance, the canal for the promontory artery in Necrolemur is only slightly thicker than that for the stapedial. In Tarsius this difference is more pronounced. What is of general significance is that when Necrolemur differs from Tarsius it is usually intermediate be- tween the latter and yet more primitive prosimians. One could hardly expect an Eocene tarsioid to be otherwise. II. BASICRANIUM Stehlin (1916:1851) mentioned that the alisphenoids partici- pated in the composition of the anterior wall of the bulla. How- ever, Htrzeler (1948:26) has pointed out that, although the bulla is overlapped by the alisphenoids, this does not permit the definite statement that the alisphenoids constitute a part of the true bulla wall. In Montauban 9, it is possible to follow the suture between the bulla and its neighboring elements from the carotid foramen around the anterior end to the squamosal. The diverticulum D 2 (of Hurzeler) appears to lie outside this suture. If then, as Hirzeler suggested, the alisphenoid forms no part of the anterior bulla wall, this diverticulum (D 2) is extra-bullar. The broad overlapping of the external pterygoid plate of the alisphenoid onto the anterolateral bulla wall in Necrolemur (Fig. 2) is a feature of some interest in relating the Querey form to the modern Tarsius. Cope (1885 :467) long ago stressed the distinetiveness of this region of the tarsier basicran- ium when comparing it with the then newly discovered skull of an American Wasatchian prosimian, Tetonius homunculus. In both Necrolemur and Tetonius these external pterygoid plates overlap the bullae, as in Tarsius. Gregory (1920:227) gives the following as a general character of lemuroid Primates: ‘‘The elongate pterygoid plates of the alisphenoids extended back to the auditory bullae, whereas in the Anthropoidea they are well separated from them.’’ A further distinction here is possible in that the posterior extremities of the external pterygoid plates in lemurs and lorises, including such fossil forms as Smilodectes, Notharctus, Adapis, and Pronycticebus, typically (although not in all cases) reach back to the anterior tip of the bulla, but the area of contact is very small and cannot be described as over- lapping. Necrolemur, Tarsius, and Tetonius differ in this respect 1960 CRANIAL ANATOMY OF NECROLEMUR il ~ from lemuriform, lorisiform, platyrrhine, and ecatarrhine Pri- mates. Stehlin could not find the stylomastoid foramen in his speci- mens; Hiurzeler (1948) shows it in figures 27 and 28 at the posteroexternal angle of the bulla, but does not label it (see Mig. 1). Just anterior to this foramen is a fossette, probably for reception of the stylohyal, if, as in Tarsius, the tympanohyal was not distinct (van der Klaauw, 1931:239). This foramen and fossette lie in the same depression and have a somewhat variable degree of separation. In M.C.Z. 8879 the external appearence is as a single oblong foramen, while in Paris 1 the two are more distinctly set off (Fig. 2). A specimen at the British Museum is intermediate in this regard. Contrary to Stehlin’s suggestion (1916 :1348) that a true post- vlenoid process does not exist in Necrolemur, Paris 1 and 5 ex- hibit a process that can justly be termed postglenoid. Also a postglenoid foramen is present (M.C.Z. 8879, Paris 1, 2 and 5) median to this process and between the posterior limit of the glenoid area and the external auditory tube. Two large foramina, one on either side of the alisphenoid pterygoidal wing, were deseribed by Stehlin (1916 :1353-1354) as the imner and outer openings of the canalis civinninii (or foramen pterygospinosum, Stehlin 1912:1205). He named a smaller opening situated anterodorsally in the same region the foramen ovale. The position of this latter small foramen is vari- able, but is always anterior to the elenoid fossa instead of being approximately on a line with it, as is the foramen ovale in Primates generally. In those specimens in which this foramen is relatively large, a groove extends laterally and slightly pos- teriorly from it, which would probably not be the case if it were the foramen rotundum. Removal of matrix from Paris 1 and 2 in the region of the external pterygoid plate or wall has re- vealed a foramen opening directly into the cranial cavity, lying within the wall at the juncture of canals from the three foramina. Given this information, probably not known to Stehlin, we sug- vest that the posteroexternal foramen (outer opening of Stehlin’s canalis civinninil) is the foramen ovale. In a footnote, Stehlin (1916:1354) cited Gregory (1915 :430) as confirming his identifications in the region of the foramen ovale. Gregory does this only partially, committing himself no further than to say that the foramen ovale is on the external (as opposed to the internal) side of the external pterygoid wall. me CO bo BREVIORA Nos 227 FIGURE 2 Necrolemur antiquus x 3.5 Foramen for branch of internal 6. Hypoglossal foramen maxillary artery 7. Alisphenoid canal Foramen ovale 8. Opening of eustachian tube Postglenoid foramen 9. Internal carotid foramen Stylomastoid foramen and fos- 10. Inferior petrous sinus sette for ? stylohyal 11. Posterior lacerate foramen Foramen for auricular branch of pneumogastric 1960 CRANIAL ANATOMY OF NECROLEMUR 9 He did not definitely say that Stehlin’s small foramen is the foramen ovale. However, he identified as the foramen rotundum the smaller foramen (Stehlin’s f. ovale) regarded by us as being for the internal pterygoid branch of the internal maxillary artery. it ORBIM Along its median wall the orbit is composed principally of the frontal and the maxillary. Careful search of Paris 1 revealed no os planum present in the orbital wall (Fig. 3). The lacrymal forms a narrow band within the orbit along the anterior rim. Frontal, parietal and alisphenoid comprise the posterior wall. No anterior sutures between the small orbitosphenoid, frontal, and palatine, respectively, could be made out in Paris 1, in which this region is entirely undistorted. Only a small palatine com- ponent is present in the orbit, and this does not separate the frontal from the maxillary. Stehlin (1916 :1345) noted that his material was not adequate to allow determination of the maxillo-malar suture. This led him to suggest that the malar might reach the lacrymal. Paris 1 and 2 show that this is not the case; the maxillary makes up part of the orbital rim. When discussing the Eocene lemuroid Noth- arctus, Gregory (1920:227) remarked ‘‘. . . the malar if not in actual contact with the lacrymal certainly came very close to it, whereas in tarsioids and anthropoids it becomes widely separated from the lacrymal and limited to the outer side of the orbit.’’ Consequently, Necrolemur resembles the higher Primates in this regard, and not the majority of prosimians other than Tarsius. The absence of an ethmoid component in the rostral orbital wall of Necrolemur has been taken by some students as an indi- cation of a lack of affinity between it and Varsius, since in the latter the os planum is large. To the writers this distinction does not seem to have much significance. A primitive prosimian condition, where the ethmoid has no orbital plate, is retained in such forms as Necrolemur, Pronycticebus, Smilodectes ete., in which, perhaps, there has not been enough orbital expansion to effect an expression of this bone in the orbit by impinging on the anterior part of the interorbital septum. In TVarsius and some Lorisiformes the interorbital septum is very narrow — ev1- dently an accommodation for relatively large eyes, and in both groups an os planum occurs. Moreover, only in Cheirogaleinae, 10 BREVIORA No. 127 among living and fossil Lemuriformes, is the os planum present. In these small lemurs also, the occurrence of an os planum is coupled with large orbits (compared to body size) and a thin interorbital septum. For the Anthropoidea, a similar origin for FIGURE 3 Necrolemur antiquus x 3.5 approx. 1. Sphenopalatine foramen and &. Ethmoid foramen posterior palatine canal Abbreviations: 2. Foramen rotundum and ante- as, alisphenoid rior lacerate foramen (?coal- fr, frontal esced ) ma, maxilla os, orbitosphenoid pl, palatine pr, parietal this orbital element may be considered. In most Ceboidea the orbits restrict the interorbital septum to a thin plate on which the os planum is exposed laterally. Although in Old World Anthropoidea the rostrum between the orbits is occasionally 1960 CRANIAL ANATOMY OF NECROLEMUR ital rather broad, it is possible to posit that such breadth is secondary and that they descend in common from a form in which rela- tively large eyes impinged on the interorbital area enough to induce the appearance of an orbital ethmoid component. This hypothesis is strengthened by observed interorbital narrowness in the only known part of an Oligocene catarrhine skull (Simons, 1959:8). If the foregoing suggestions apply, then it is not neces- sary to expect the presence of an os planum in the stock from which Tarsius may have arisen. In Necrolemur (Paris 1), a small venous foramen can be seen situated near and beneath the median dorsal rim of the orbit. As in Tarsius, but apparently not in other Primates, below this foramen a deep groove curves posteroventrally and (in both) hes at a juncture between the plane of the lateral wall of the rostrum and that of the back of the orbit. This is another un- usual feature (occurring in both Necrolemur and Tarsius) which has to be attributed to independent acquisition, by those who doubt that any known fossil prosimians have a close phyletic relationship to Tarsius. A eranio-orbital foramen exists in Paris 1, but could not be found in Paris 2, 3 or 5. Running anteroposteriorly and shghtly above the optic foramen is another groove. A small opening near its anterior end appears to be the ethmoid foramen. The sphen- opalatine foramen and the posterior palatine canal in Paris 1] are combined to open posteriorly through a common large fora- men in the suture between maxillary and palatine. Apparently coalescence of the foramen rotundum and _ the anterior lacerate foramen oecurs in Paris 1 but they are separate in Paris 5. In both cases, however, the foramen rotundum hes within the orbit, as Stehlin noted (1916 :1353), and not lateral to the postorbital part of the alisphenoids, as Gregory (1915 :450) suggested. The orbital region of Paris 1 is entirely undistorted and shows that the postorbital opening was small. It is of some interest that neither lemurs nor lorises, nor any other known fossil prosimians of similar size possess a smaller aperture here. As with so many eranial characters the primate showing the most interesting re- semblance to Necrolemur in respect of the structure of the post- orbital region is T'arsius. However, because of the huge flanges that encircle the orbit in mature specimens of Tarsius, and the ereater degree of postorbital closure seen in such adults, much more revealing comparisons can be made between skulls of 12 BREVIORA INOS 2a Necrolemur and those of juvenile tarsiers. In the latter, the circumorbital flanges are not yet very pronounced and resemble the slight flanges seen in Necrolemur (Fig. 3). It seems necessary to assume, first, that these flanges in Tarsius are concerned pri- marily with the support of the enormous eyes, and second, be- cause of their very uniqueness, that they were not so developed in Kocene forerunners of the living genus. Necrolemur clearly has such fianges in an incipient stage. Closure behind the orbit in Tarsius (on the outside) can be seen to proceed in successively older juveniles from three main centers: 1, ventrolaterally, by an upgrowth of the posterior mid- region of the orbital plate of the maxilla; 2, laterally, by an anteroposterior spreading of the middle of the postorbital bar; and 3, dorsolaterally, by growth of a flange from the frontal, which arises beneath the frontal insertion of the postorbital bar. Ossification proceeds downward as this flange, or plate, grows alongside the postorbital bar, and it eventually fuses with the bar, leaving no sutural indication. In most specimens of adult Tarsius, the frontal and jugal components of the postorbital wall can be distinguished by the fact that the region where they fuse is much thinner and consequently more translucent. In WNec- rolemur, at least two of these components effecting closure ap- pear to be partially developed. Paris 1 has an uncrushed post- orbital bar which shows an anteroposterior spreading at the middle, much as in Tarsius. The evidence is less clear because of breakage, but the posterior part of the orbital plate of the maxilla also bears a flange in Necrolemur. The third center of closure seen in Tarsius, the frontal element, is indicated in Necrolemur only by a distinct angulation along the lateral wall of the cranium betwen the orbital and temporal fossae. Although such characters as the greatly reduced paraconids and the loss of certain anterior lower teeth eliminate Necrolemur from the direct ancestry of Tarsius1, the incipient circumorbital flanges and characteristics of postorbital closure in Necrolemur are sug- gestive of a stage to be expected in the Tarsius ancestry. Perhaps Pseudoloris or Nannopithex are nearer the actual line leading to the modern form, but incompleteness of known specimens pre- vents the sort of comparisons here made with Necrolemur and leaves this possibility imsoluble at present. 1Byven if this were not so it would be almost irrelevant, in the absence of intermediates, to urge an ancestor-descendent relationship for forms so separate in geographic distribution and in time. 1960 CRANIAL ANATOMY OF NECROLEMUR ils; It may be noted in passing, that the manner of postorbital closure in Tarsius (insofar as the malar and frontal are con- cerned) is distinct from that seen in catarrhines and platyrrhines. Closure in this area in Tarsius is chiefly effected by an outward and downward growth of a flange of the frontal (with relatively little malar expansion) while in higher Primates the greater part of the dorsolateral area of enclosure is contributed by the de- velopment of an orbital plate of the malar. These differences strongly imply that the partial postorbital closure of Tarsius only parallels that of the Anthropoidea and is not a character of their common inheritance. If it be agreed that some postorbital clo- sure arose at least twice among Primates, the possibility that this feature also was independently acquired in the ancestral platyr- rhine and eatarrhine stocks can be more seriously entertained. EXPLANATION OF FIGURES The lateral and ventral views of the skull of Necrolemur are based on Paris 1, to which details of missing regions have been added from other specimens, principally M.C.Z. 8879. Mislead- ing stains and fractures in Paris 1 are largely omitted from these illustrations. Certain details of the anterior dentition and mandible are drawn from specimens figured by Stehlin (1916). REFERENCES Cops, E. D. 1885. The Lemuroidea and the Insectivora of the Eocene period of North America. Amer. Naturalist, 19: 457-471. EDINGER, T. and D. B. Kirts 1954. The foremen ovale. Evolution, 8: 389-404. GREGORY, W. K. 1915. I. On the relationship of the Eocene lemur Notharctus to the Adapidae and to other Primates. II. On the classification and phylogeny of the Lemuroidea. Bull. Geo. Soc. Amer., 26: 419-446. 1920. On the structure and relations of Notharctus, an American Eocene primate. Mem. Amer. Mus. Nat. Hist., (n.s.) 3: 49-248. HURZELER, J. 1946. Zur Charakteristik, systematischen Stellung, Phylogenese und Verbreitung der Necrolemuriden aus dem europiischen Eocaen. Eclogae geol. Helvetiae, 39: 352-354. 1948. Zur Stammesgeschichte der Necrolemuriden. Schweiz. palaont. Abh., 66: 3-46. 14 BREVIORA No. 127 KLAAUW, C. J. VAN DER 1931. The auditory bulla in some fossil mammals. Bull. Amer. Mus. Nat. History., 62: 1-352 Lr Gros CLARK, W. E. 1959. The antecedents of man. 374 pp. Edinburgh University Press. Simons, E. L. 1959. An anthropoid frontal bone from the Fayum Oligocene of Egypt: the oldest skull fragment of a higher primate. Amer. Mus. Novit., no. 1976: 1-16. 1960. Notes on Eocene tarsioids at the British Museum. Bull. Brit. Mus. Nat. Hist., Geol. Ser., in press. STEHLIN, H. G. 1912. Die Siugetiere des schweizerischen Eoeaens. Abh. Schweiz. Paldont. Gesell., 38: 1165-1298. 1916. Die Siugetiere des schweizerischen Eocaens. Jbid., 38: 1299- 1552. BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. DECEMBER 20, 1960 NuMBER 125 SIZE OF ENDOCEROID CEPHALOPODS! > By Curt TEICHERT? AND BERNHARD KUMMEL®? The maximum size of fossil animal groups, whether mammals, reptiles, or invertebrates has always been a fascinating subject of inquiry, because phyletic size increase is one of the important trends that dominate the evolution of living things. In the case of large animals, the evidence is often hard to assemble because their remains are difficult to obtain, to transport, and to store. Squids are the largest living invertebrates and a tradition has been handed down in paleontological literature that the largest fossil invertebrates likewise are to be found amone the cephalo- pods, but few accurate data are to be found in published sources which are now readily available. Amone the nautiloid cephalopods, it has long been suspected that the Endoceratida furnished the real giants, but no accurate measurements in support of this statement are available. Clarke (1897) stated that entire shells of Cameroceras pro- teiforme, 10 to 15 feet long (3 to 5 meters), had been found in the Middle Ordovician of Minnesota. In the same publication, Clarke figured an internal cast of part of a siphuncle, from the base of the body chamber to the adapical end of the spiess, which was 3 feet and 3 inehes lone. Miller and Kummel (1944) deseribed and illustrated additional species of these Middle Ordovician endoceroids from Minnesota, which are deposited im the Carnegie Museum. One of their paratypes of Endoceras clarkei measured 750 mm lone, is septate throughout and is not complete, adapically or adorally. The holotype of Hndoceras gracillimum Miller and Kummel (1944) measured 670 mm in LPublication authorized by the Director, U. S. Geological Survey. 2U. S. Geological Survey, Denver, Colo. 3Museum of Comparative Zoology. 2 BREVIORA No. 128 length, again an incomplete specimen consisting only of phrag- moeone. These same authors described a new species, HE ndoceras decorahense, on two portions of the internal mold of the phrag- mocone from the Decorah formation, Winneskiek County, Iowa. The larger portion is about 625 mm lone and the length of the smaller measures about 320 mm. They estimated the interval between the two pieces as about 115 mm, so the total length of this phragmocone was about 1,060 mm. These authors also men- tioned that there is on display in the Chicago Natural History Museum a larger endoceroid that measures 6 feet in length. Teichert (1927) noted the occurrence, in Middle Ordovician limestones of Estonia, of endoceroids as much as 5 meters long, but gave no further details. Flower (1955) stated that specimens 12 feet in length had been colleeted and added that he was ‘‘not wholly inclined to discredit a report of an endoceroid found in a quarry near Watertown, New York, which was measured before it was broken up and found to attain a leneth of 30 feet.”’ As far as we have been able to ascertain, these somewhat vague statements are all that is presently available in the published rec- ord on the subject of the maximum size of endoceroid cep- halopods. It does not seem to be generally known that the Museum of Comparative Zoology at Harvard University possesses what ap- pears to be the largest fragment of an endoceroid cephalopod on display anywhere in the world. As Flower (1955) has stated, “the removal of even reasonably complete specimens involves something very close to quarrying operations, storing them is another problem.’’? The specimen in the collections of the Mu- seum of Comparative Zoology is, therefore, probably unique in museums of the world. The specimen measures 3,000 mm in length but is not com- plete, adorally or adapically. In general the preservation is fair, but as a result of weathering and crushing the full diameter of the conch is preserved only in one plane, and in the other plane the outer shell is removed exposing traces of septa and in places the siphuncle. The first recognizable septa are 500 mm from the adoral end but the whole specimen could well be phragmocone as this adoral 500 mm is shghtly erushed and weathered and one cannot tell whether septa are present or absent. The adoral diameter of the specimen is 280 mm. The coneh tapers at a uniform rate and the adapical diameter meas- ures 120 mm. 1960 SIZE OF ENDOCEROID CEPITALOPODS i ‘ Figure 1— Large endoceroid on exhibit in the Museum of Zoology. Comparative 4 BREVIORA No. 128 The septa slope adapically at an angle of about 45° and in the mid-part of the specimen are spaced 17 to 20 mm apart. The siphunele is visible only on the adapical half of the specimen. About 1,000 mmm from the adoral end of the shell the siphunele has a diameter of about 95 mm; at 1,750 mm from the adoral end of the shell the siphunele has a diameter of 75 mm. The first endocones appear 2,000 mim behind the adoral end of the shell. The spiess measures 510 mm in leneth. The surface of the shell bears faint annulations that are spaced approximately 10 to 12 mm apart. SUMMARY OF MEASUREMENTS Length pe ....3,000 mm Adoral diameter : eee oman Diameter 1,000 mm from adoral end 220 mm Diameter 1,750 mm from adoral end ale. O moa Adapical diameter .. 120 mm Diameter of siphuncle 1,000 mm from adoral end 95 mm Diameter of siphunele 1,750 mm from adoral end 75 mm Spiess length ae e ..510 mm A eraphical reconstruction of the shell on the basis of these measurements shows that the entire fossil from its presently pre- served adoral end to the apex may have measured about 6,500 mim. The total leneth of the body chamber is a matter of guesswork. There are few published and illustrated records of any straight fossil cephalopod shells, complete from apex to aperture, which are more than a foot or so lone. In short shells the ratio of body chamber to phragmocone may be high, even larger than 1:1. With increasing total leneth of coneh, however, ratio of body chamber to phragmocone is likely to decrease, although no defi- nite figures can be stated. ina specimen of Actinoceras beloitense (Foerste and Teichert, 1930, pl. 28), which was 450 mm long, the ratio of body chamber to phragmocone was about 1:2. Leith (1942) described a specimen of Lambcoceras lamba (Whiteaves ) which was 45.5 in. (1,155 mm) lone. He estimated the total leneth of the shell at 1405 mm. The body chamber was almost wholly preserved and not more than 250 mm long. Ratio of body chamber to phragmocone was thus 1:4.6 in this specimen. It should be noted, however, that both Actinoceras beloitensc and Lambcoceras lambii have body chambers with constricted 1960 SIZE OF ENDOCEROID CEPHALOPODS 5 apertures, whereas no endoceroids with constricted apertures are known. It seems physiologically plausible that in large straight cephalopod shells the animal should have a_ better ‘“‘orip’’? on a body chamber with constricted aperture than on one with an uneonstricted aperture; therefore, in shells which expanded uniformly from the apex to the aperture, like the endoceroids, the animal itself, and thus its body chamber, should have been relatively larger. In a juvenile specimen of a straight ammonoid, Baculites ovatus, Trueman (1941) determined the ratio of length of body chamber to phragmocone as 1:0.7, but in adult shells this ratio becomes much smaller. If we assume the ratio of length of body chamber to phragmocone in endoceroids to be more lke that of Actinoceras beloitense we arrive at a length of the body chamber for the Harvard Endoceras of 2,650 mm and for the en- tire shell of 8,150 mm, or 28 feet. This is a conservative esti- mate, yet close to the possible maximum figure of 50 feet men- tioned by Flower. Add to this the length of the tenacles which must have ex- tended a considerable distance in front of the aperture, cer- tainly no less than half the length of the body chamber, and we have an invertebrate animal considerably longer than 30 feet —a truly imposing size. Today’s giant squid, Architeuthis, rivals and slightly exceeds in length the largest extinct endocer- oids. Spirek (1928) records specimens of Architeuthis dux from the North Atlantic, washed ashore on the Norwegian Coast, that have body lengths of up to 2 meters and tentacles as much as 10 meters long. The largest specimen to our knowledge is that of Architeuthis harveyi? recorded by Verrill (1879, p. 196) which measures 624 inches (17 meters). There is a model in the Museum of Comparative Zoology of a specimen of Archi- teuthis princeps, which was washed ashore in Newfoundland, which measures about 15 meters in length. More recently, Lane (1960, pp. 198-227) has critically reviewed a larger number of reports of finds of and encounters with giant squids. He is in- clined to believe that individuals of Architeuthis or some other venus, as yet undescribed, may reach overall leneths of some 70 feet. While the Harvard specimen represents by far the largest nautiloid cephalopod on which accurate data are now available, it is interesting to compare it with the largest ammonoid on record. This is Pachydiscus seppenradense Landois from the 6 BREVIORA No. 128 Upper Cretaceous of western Germany (Landois, 1895, 1898). In 1895, Landois first described this fossil ammonoid whose shell was 1,800 mm im diameter and in which the last camera was 590 mm high. Landois’ reconstruction provided the animal with a body chamber equivalent to only one-fourth of a complete whorl. From this he concluded that the total diameter of the complete specimen of his ammonoid had been about 2,550 mm. From later studies (Trueman, 1941) it is, however, likely that Landois’ estimate of the leneth and bulk of living chamber was too low. If the body chamber of Pachydiscus seppenradense was equivalent, as is more likely, to three-fourths or one full volu- tion of the shell, the diameter of the adult shell of this ammonite would have been of the order of 3,500 mm, or more than 10 feet. A very approximate graphic plot of a shell of this kind shows that the total length of the shell of Pachydiscus seppenradense, when unrolled, would have been of the order of 60 feet, or roughly twice as long as that of the largest endoceroid. In another paper Landois (1895) attempted to estimate the weight of these giant cephalopods. On the basis of his estimated measurements he arrived at a total weight of the ammonite as 1,455 ke, or 750 ke for the weight of the animal itself, and 705 ke for the weight of the shell. We shall abstain from any attempt to indicate exact weights of the large endoceroids. The order of magnitude was almost certainly the same as that inferred by Landois for the giant Pachydiscus, something of the order of 1 ton (about 1000 ke). It must be assumed that the weight of shell and siphunele, which for a leneth of over 5,000 mm was entirely filled with calcareous deposits, balaneed the buoyancy provided by the empty camerae and confined the animal to a strictly benthonic existence. Few, if any, fossil invertebrates ever surpassed them in bulk weight and size. One other point deserves attention: Phyletic size inerease is a trend that as a rule continues until the end, or very close to the end of the evolutionary life of a particular group of organisms, as, for example, in the ammonoids. The endoceratids, however, reached their maximum size lone before the time of extinetion, in fact relatively early in their evolution. In North America, as well as in northern Europe, endoceroid cephalopods survive to the end of the Ordovician period, but reach their maximal size during Middle Ordovician time (Teichert, 1930, pp. 235-236). 1960 SIZE OF ENDOCEROID CEPHALOPODS Ti REFERENCES CLARKE, J. M. 1897. The Lower Silurian Cephalopoda of Minnesota. Geology of Minnesota, vol. 3, pt. 2, Paleontology, pp. 761-812, pls. 47-60. _ ov ple Ee 1955. Status of endoceroid classification. Jour. Paleontology, vol. FLOWER, 29, no. 3, pp. 329-371. Forrstr, A. F., and TEICHERT, CURT 1930. The actinoceroids of east-central North America. Denison Univ. Bull., Sei. Lab. Jour., vol. 25, pp. 201-296, pls.: 27-59. LAnNpbotrs, H. 1895. Die Riesenammoniten von Seppenrade, Pachydiscus, Zittel, Sep- penradense H. Landois. Westfal. Provinzial. Ver. Wiss. und Kunst. f. 1894/95, vol. 23, pp. 99-108, 2 pls. 1898. Gewichtsverhiltnisse der Riesen-Ammoniten. Ibid., vol. 26, pp. 27-28. GAN, BY OW: 1957. Kingdom of the Octopus. Jarrolds, London. 286 pp. = 4 Lrita, HE. I. 1942. Notes on the cephalopod Lambeoceras lambii from Manitoba. Jour, Paleontology, vol. 16, no. 1, pp. 130-132, 1 text-fig., pl. 22. Miuuer, A. K., and KUMMEL, BERNHARD 1944. Some large straight Ordovician cephalopods from Minnesota. Annals Carnegie Museum, vol. 30, pp. 19-38, 4 pls. SPARCK, RAGMAR 1928. Nordens Dyreverden. Henrik Koppel, Copenhagen, 658 pp. TBICHERT, CuRT 1927. Der estlindische Glint. Natur u. Museum, vol. 57, pp. 264- 272, 7 figs. 1930. Biostratigraphie der Poramboniten. Neues Jahrb. f. Mineral. etc., Beil. Bd., Abt. B, pp. 177-246, 8 figs., 4 pls. TRUEMAN, A. EH. 1941. The ammonite body-chamber, with special reference to the buoyancy and mode of life of the living ammonite. Quart. Jour. Geol. Soc. London, vol. 96, pp. 339-378. ViRRILL, A. HB. 1879. The cephalopods of the northeastern coast of America. Trans. Conn. Acad. Sci., vol. 5, pp. 177-476, pls. 25-56. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 21, 1960 NUMBER 129 DYE BANDS LY PH LOCALITY OF TEE GULE (COAST SPINY SOFTSHELL TURTLE, TRIONYX SPINIFER ASPER (AGASSIZ) By Ropert G. WEBB Museum of Natural History, The University of Kansas The currently accepted type locality of Trionyx spinifer asper, Lake Concordia, Louisiana, is in an area of intergradation be- tween three subspecies of Trionyx spinifer. Of the nine available syutypes of asper, none has been designated as a lectotype, and only one of the syntypes of 7’. s. asper is recognizable as belong- ing to that taxon. Abbreviations of names of museums from which specimens are mentioned are: KU, Museum of Natural History, University of Kansas; MCZ, Museum of Comparative Zoology, Harvard College; SM, Strecker Museum, Baylor University; TU, Tulane University ; and USNM, United States National Museum. Agassiz (1857:406) described Aspidonectes (= Trionyzx) asper as having: ‘¢ | . very coarse and large tubercles of the front and hind part of the carapace, which extend, behind, even over the bony shield, and are there supported by prominent warts of the bony plates. These bony warts exist in no other species with which I am acquainted: their form is very irregular, sometimes oblong and sometimes orbicular; they also project more or less. Another marked peculiarity of this species consists in the greater bluntness of the extremi- ties of the jaws, which are more rounded than in Asp. spini- fer. The jugal arch is also broader. The difference between the males and the females is more striking in this species than in any other, the males being regularly oval, whilst 2 BREVIORA No. 129 the females are almost circular in their outline . . . in younger specimens of Asp. asper there are ... two or three black lines separating the pale rim of the posterior mar- Cali Tyg ea Several of the syntypes show ‘‘prominent warts of the bony plates’? (supra), which appear posteriorly and principally on the seventh pair of pleurals toward the midline on the bony carapace (Plate 1). I have seen bony elevations, which are circular or elongate resembling short ridges, on the carapaces of specimens of Trionyx ferox (USNM 43878, 55316, 62217); a photograph of a ferox (Stejneger, 1944 :pl. 7) clearly shows bony prominences on the posterior part of the bony carapace. These bony elevations also occur in populations of 7. spinifer (SM 2502, 2558, Texas; USNM 54731, Iowa; USNM 100396, 100404, Louisiana). Bony warts and ridges on the posterior part of the carapace are not diagnostic for 7. s. asper. To my knowledge, the subspecies of Trionyx spinifer are not distinguishable by characteristics of the skull, bony carapace or plastron. T. s. asper closely resembles the subspecies hartwegi and sprnifer but differs in usually having two or more blackish lines paralleling the rear margin of the carapace, and usually in having the postocular and postlabial stripes united on the side of the head. Other characters mentioned above by Agassiz seem not to be of taxonomie worth. Agassiz (op. cit.:405-06) did not designate a type, mention precise localites, or state the number of specimens that formed the basis of his description of Aspidonectes asper. Baur (1893: 220) restricted the type locality of asper to Lake Concordia, Louisiana, but did not mention any specimens; presumably Baur’s action was based upon an examination of the bony cara- pace of USNM 012349, which shows the prominent bony ridges described and considered diagnostic by Agassiz and has ‘‘Lake Concordia, Louisiana’’ written in ink on the underside of the carapace. Barbour and Loveridge (1929 :225) listed MCZ 1597 and 1622 as cotypes. Stejneger (op. cit.:56-58) discussed some of the syntypes of asper, and also regarded the type locality as “‘Lake Concordia, la.,’’ designating USNM 12349 (= 012349) and MCZ> 37173 as ‘‘cotypes.’’ The type locality of T. s. asper is currently accepted as Lake Concordia, Louisiana (Schmidt, 1953:109). T. s. asper intergrades with 7. s. hartwegi and T. s. spinifer in the lower Mississippi Valley (Conant and Goin, 1948:11). 1960 TRIONYX SPINIFER ASPER 3 The softshell turtles inhabiting the Mississippi River and its tributaries in Louisiana (including Lake Concordia) and Mis- Sissippl represent an intergrading population of spinifer and hartwegi, and, to a lesser extent, asper. Most turtles from the Pearl River drainage and rivers that drain into Lake Ponchar- train adjacent to the east are typical of asper. Lake Concordia is a large oxbow on the west side of the Mississippi River in Concordia Parish, Louisiana. I was a member of a field party, from Tulane University, which collected three specimens (TU 16524, 16524.1, 16524.2) of Trionyx spinifer from Lake Con- cordia on August 1-3, 1954. Because none of these turtles has the postocular and postlabial stripes united on the side of the head or any indication of more than one marginal line parallel- ing the rear margin of the carapace, none is considered referable to asper. Some of the nine syntypes discussed below were mentioned by Stejneger (op. cit.:57-58). (1) USNM 012349 (Plate 1) is represented by a bony eara- pace and may be considered the present type (lectotype), although never designated as such. ‘‘Trionyx Ferox?, Lake Concordia, Louisiana, BLC Wailes, 1851’’ is written in ink in the same handwriting on the underside of the bony carapace. Subsequently, ‘‘Ferox?’’ has been crossed out and ‘‘asper’’ added in pencil, and ‘‘012349’’ inked on the second pleural. > gular scales ca. = ventrals ventrals ventrals 14-17 enlarged dorsal 11-13 enlarged dorsal 10-12 enlarged dorsal scales in distance snout scales in standard scales in standard to middle of eye distance distance (standard distance ) median rows of enlarged median rows of enlarged median rows of enlarged dorsal scales about as dorsal scales mostly dorsal seales about as broad as long longer than broad lhroad as long 17-21 ventrals in standard 11-14 ventrals in 11-14 ventrals in distance standard distance standard distance Other differences have been listed by Mertens or Cochran, but they are at best modal differences or they alter markedly with age. These species are indeed close, and females and juveniles are sometimes difficult to distinguish. Discussion: Anolis cochranae combines in new ways charac- ters of A. semilineatus and A. olssoni. It is in no sense an intermediate; its characters are either those of one or the other or are somewhat exaggerated versions of a trend present in one. It is necessary to admit that we know very little about this species beyond its existence. Its distribution would appear, on present evidence, to be extraordinarily limited. It may well be confined to the high interior, but its real range is surely more extensive than known at present. The area from which it comes is remarkable for certain peculiar forms: Celestus darlingtom and Audantia shrevei in the higher elevations, Anolis aliniger (described as a subspecies of A. chlorocyanus by Mertens in 1939 but in reality a full species) in the vicinity of Constanza itself, Anolis darlingtoni both from Constanza and from higher elevations. The region merits extensive and systematic collect- ing. The biological relation of A. cochranae to the other two mem- bers of the semilineatus group is equally unknown. We do not 10 BREVIORA No. 135 know its contacts with either form. Its relationship to A. semilineatus in particular is puzzling. In squamation it differs strongly enough that we have called it, as a matter of judgment, a distinct species. The scale differences from both semilineatus and olssoni are as great or greater than the differences between other closely related sympatric fully valid species. But in other Anolis such (or lesser) differences are correlated with color and dewlap differences that are evident visual cues to species recog- nition. In color and dewlap A. cochranae exactly resembles one of the neighboring species — semilineatus. If A. cochranae is indeed a full species that is at some point in contact with semv- lineatus, it is necessary to suppose that there is some unknown behavioral difference that maintains the distinctness of the population in the absence of color cues. REFERENCES CITED CocHRAN, D. M. 1941. The herpetology of Hispaniola. Bull. N. S. Nat. Mus., 177: 1-398. MERTENS, R. 1939. Herpetologische Ergebnisse einer Reise nach der Insel His- paniola, Westindien. Abh. Senckenberg. Naturf. Ges., 449: 1-84. ScuHMIpT, K. P. 1919. Descriptions of new amphibians and reptiles from Santo Domingo and Navassa. Bull. Amer. Mus. Nat. Hist., 41: 519-525. 1921. The herpetology of Navassa Island. Bull. Amer. Mus. Nat. Hist., 44: 555-559. ala ANOLIS COCHRANAE 1961 “RpoTuBdsiyT ut dnois snjyoauywuwas syoup ey4 JO sotoeds oe1y} 94} JO UOKYNGII4sIq : dey 069 U Od Sé_09 Sr of SI_O SddILIWOTNY 40 3719S IOINBdSITH old old \uOSS[|O @ snpeautiMmas O 2eueiyso0o of td a . . om « oe 7 s - aa - - a ; f 4 ~'s e shod . z r ‘ ? a ha . ‘ . ; F pe: bolas . - + bs , sae i : 2 r * ' ay ae . ? = - 1 - me.5 . se + " ' ae aia . a arg oo os ¥ ; ‘i i bene BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. ApRIL 8, 1961 NUMBER 136 NOTES ON HISPANIOLAN HERPETOLOGY 3. THE EVOLUTION AND RELATIONSHIPS OF THE ANOLIS SEMILINEATUS GROUP By Ernest E. WILLIAMS The discovery of a third species of the Anolis senvlineatus group, confined apparently to the high interior of the Dominican Republic, poses problems in the distribution, biology and evolu- tion of the group. The distributional data for the semilineatus group has been given in Williams and Rand (1961) and need not be repeated in detail here. A. semilineatus and A. olssoni are both widely distributed north of the Cul de Sae Plain but occupying eco- logically somewhat different situations and thus with but limited actual contact or overlap; only A. semilineatus at present is known south of the Cul de Sac Plain in the southwest and Barahona peninsulas. A. cochranae is found in the center of Hispaniola in the Cordillera Central — geographically in the midst of the other two species though its contacts with these others are not known. The biological peculiarity in the relation of A. cochranae to A. semilineatus has also been pointed out in Williams and Rand (1961). Thus, though differing strongly from the closely related A. semilineatus in certain scale characters, A. cochranae is iden- tical in body and dewlap color. This phenomenon is highly unusual in the genus Anolis in which body and dewlap color dif- ferences are important cues in species recognition. (There are, for example, strong body and dewlap color differences between A. semilineatus and A. olssoni.) A. cochranae, if it is in contact with A. semilineatus, as A. semilineatus and A. olssoni are in contact with one another, would seem to be a most anomalous case in which it would be necessary to provide some ad hoc explanation — such as some unknown behavior difference — for the maintenance of the species distinction. bo BREVIORA No. 136 The problem is thus to provide an explanation of the central veographic position of Anolis cochranae in Hispaniola and of the curious absence in cochranae of the usual anoline species recog- nition characters contra a related species that occurs literally on every side of it. I propose below a suggested history of the semilineatus group that appears to solve this problem. It must be admitted that this proposed history depends upon taking at face value the distributions of the three species as they are known at present. This is patently unsafe, but it provides a useful starting point. On our present knowledge of distribution it is simplest to suppose that the postulated biological problem has not arisen, that cochranae and semilineatus are nowhere in contact. This is at the moment only a brave hypothesis. Anolis cochranae is known from only two collections; our more extensive knowledge of the distributions of semilineatus and olssoni is by no means 2ood enough to prove contact or absence of contact with coch- ranae. Critical to the proposed history is the supposition — uncon- tradicted by the available evidence — that olssont is really absent from the southwest and Barahona peninsulas. It does appear to be absent from the moist coastal zone at Aux Cayes (observations by A. S. Rand and J. Lazell in 1960) and Rand did not collect it in the dry area of Oviedo on the Barahona peninsula in 1959. It is not present in Hassler’s collections from these two areas. Let us then take the present distributional evidence at face value. Let us assume then that semilineatus is the only grass anole of the southwest and Barahona peninsulas and that olssona just touches this area at the southern edge of the Cul de Sac Plain. The southwest and Barahona peninsulas taken together are just that portion of the island which was cut off from the mass of Hispaniola by the Pleistocene seaway through what is now the Cul de Sae Plain. Residual salt lakes and coral rocks still testify to this former seaway. The division of Hispaniola into two parts which resulted from this seaway provides two suitable theatres —a main island and a southern counterpart — for the classic pattern of speciation during separation, and intensification of species difference (‘‘character displacement’’) during renewed contact. On this hypothesis semineatus is the autochthonous. grass anole of the southern cut-off portion of: Hispaniola and: oalssont 1961 ANOLIS SEMILINEATUS GROUP 3 and cochranae autochthons of the northern main mass of the island. Semilineatus has infiltrated the northern island all but completely, while olssoni is not known to have invaded the south- ern island. The spread of semilineatus through much of the northern island is not too surprising in view of its eurytopic ecology (Mertens, 1939, Williams and Rand, 1961). Though character- istic of a specialized open habitat, it seems to be sufficiently tolerant of forests that these would be less efficient barriers to its spread than they would to stenotopic olssont. It is somewhat more surprising —if it is true—that olssoni has not spread alone the dry north coast of the southwest peninsula or the east coast of the Barahona peninsula, but it would be stopped easily by discontinuities in suitable habitat and would for this reason be unlikely to reach localities otherwise quite suitable to it on the southern island. The different coloration in olssoni, including the dewlap color, and the large size of the dewlap scales may well have developed after olssont came into secondary contact with semilineatus dur- ing the latter’s invasion of the northern island fragment. In suggesting this we assume that the features in common of cochranae and senuilineatus are primitive and that modification in these features took place exclusively or almost so in olssoni. (Surely the lack of enlargement in the eular scales is primitive in senilineatus and cochranae; this leaves only color in ques- tion. ) What, however, about the origin and relationship of cochranae and olssoni? It must first be noticed that there is some plausi- bility in considering these two more closely related to each other than to semilineatus. In body squamation (i.e. scale size), coch- ranae and olssoni are very similar. This is a feature which, unlike the characters of the dewlap or of body pattern, is un- likely to be a matter of intra- or inter-species recognition. We do not know that it is per se adaptive: the difference in scale size between semilineatus, on the one side, and cochranae-olssoni, on the other, is more likely to be the external expression of more fundamental genetic divergencies. No physiographic barrier, however, will account for the divi- sion of the grass anole population of the northern or main Hispaniolan island into two species. It is necessary to suppose that the barrier was an area of unsuitable ecology, i.e. moist dense forest. Olssoni may then be supposed to have arisen in 4 BREVIORA No. 136 the arid coastal lowlands while cochranae arose in the open areas of the high pine woods‘ of the interior valleys of the Cordillera Central. (We note that Wetmore and Swales, 1931, p. 24, describe the natural vegetation of the Valle Constanza as ‘‘ forests of open pine mingled with areas of dense rain forest.’’) The known habitat of cochranae — Valle Constanza —is a high interior valley of the Cordillera Central. Though the floor of this valley is not very high (ca. 8000 feet) it is surrounded by some of the highest peaks in Hispaniola and ingress to it at moderate elevations is somewhat narrow and limited. In such an area a grass anole population might indeed enjoy a measure of isolation from other grass-bush populations — the more so if we suppose that the separation of olssont and cochranae dates from a period in which the density of the hardwood forest of intermediate elevations was at a maximum. Relationships of the semilineatus group. There are no other anoles in Hispaniola which either very much resemble or seem very closely related to the semilineatus vroup. A search for close relatives and ancestors takes us at once outside Hispaniola. Two Greater Antillean groups of Anolis are structurally simi- lar —the alutaceus-clivicolus-cyanopleurus-spectrum group im Cuba and the krugi-pulchellus-poncensis series in Puerto Rico. (None of the anoles of Jamaica or the Bahamas are similar either ecologically or structurally.) Both the Cuban and the Puerto Rican series share with the semilineatus group the middorsal zone of enlarged scales (least developed in krugi of Puerto Rico). All except alutaceus-clivi- colus have keeled ventrals. The Cuban anoles are all forest species, A. alutaceus occurring in rather deep shade, A. spectrwm in less deep shade. But, though in this regard they differ from the Hispaniolan species which are fonder of open areas, they are closer to the semilinea- tus group in structure than are the Puerto Rican species. Like the semilineatus group they are small, usually under 40 mm snout-vent length, slender, with large dewlaps and well developed postanal seales in the males. In color, however, they differ in never possessing the flank stripe so characteristic of the semi- lineatus group, tending instead to emphasize the light middorsal stripe. 1 Pine in Hispaniola, in contrast to e.g. Cuba, is confined to higher elevations. 1961 ANOLIS SEMILINEATUS GROUP 5 Of the alutaceus series, clivicolus, which may be a subspecies of alutaceus, has the least slender habitus and the least specialized squamation. It is easy to envision this as representing the primi- tive stock of this series. The Puerto Rican series is, on the other hand, more similar to the Hispaniolan species in habits. Two of the three species — pulchellus and poncensis — are ‘‘grass anoles’’ or at least anoles of open reaches. The third species — krugi — is an anole of denser brush. All are larger than any species of the semi- lincatus group — nearer 50 mm than 40 mm snout-vent length. They are perhaps not as slender as their parallels in Hispaniola (though this is a character difficult to estimate objectively) ; the dewlaps are relatively small and the postanal scales poorly developed. All three have a flank stripe passing forward through the eye more or less well expressed. In both Cuba and Puerto Rico the series exhibit a wider range of structure than do the Hispaniolan forms. In each series there is a species with the middorsal zone of enlarged keeled scales less developed than in any Hispaniolan species (in Cuba — clivicolus-alutaceus, in Puerto Rieco—krugi) and one with this zone much more strongly developed than in any Hispaniolan species (in Cuba — spectrum, in Puerto Rico — poncensis). One difference appears in this regard: in all the Cuban forms the width of the zone of enlarged dorsal scales is about the same (ca. 8 scale rows as compared with ca. 10 in Hispaniolan forms), while in the Puerto Rican forms concurrently with increase in the size of the middorsal scale zone, there is an increase in the number of rows enlarged (ca. 4 in krugi, ca. 12 in pulchellus, 15+ in poncensis). The evaluation of these resemblances, which are in each case beset with significant differences, is difficult. Parallelism is very probable, and it is especially likely that the Puerto Rican series is an independent radiation within Puerto Rico from the same stock that gave rise to A. cristatellus, A. stratulus, A. gundlachi and A. evermanni. The primitive member of the Puerto Rican series, A. krugi, is not very different from cristatellus and gundlachi and would certainly be classed with them except for its obvious position at the base of a small grass anole radiation on Puerto Rico. The Cuban anoles which display a strong structural affinity im spite of some habiiat difference are more probably close relatives of the Hispaniolan series. There is in fact no substan- tial reason for doubting the relationship. 6 BREVIORA No. 136 It must be pointed out that the squamation pattern with a strongly developed middorsal zone of enlarged keeled scales, smaller laterals, and strongly keeled ventrals as large or larger than the middorsals is common in mainland Anolis, particularly so in Central America. This pattern occurs also in the Greater Antilles in three species which, though certainly anoline, are currently referred to other genera: the Cuban species (ophio- lepis) to Norops, and a species from Navassa (barbourt) along with one from Hispaniola (wetmorei) to Chamaelinorops. The mainland forms exhibit a whole spectrum of conditions in regard to the distinctness, number of scale rows, size of scales involved in the dorsal zone, ete. No described form seems close enough to the Hispaniolan or Cuban grass anoles to be worth serious consideration as representing the ancestral stock. Norops ophiolepis, which occupies the grass anole habitat in Cuba, does not seem related either. It has some features peculiar to itself — the reduction of the canthal ridge scales to two, the small number of scales in the loreal area (ca. 10-12), the very elongate scales between the nostrils, the large mental scales — that are unlike not only the semilineatus-alutaceus groups but its supposed congeners on the mainland. The relationships of ophiolepis are probably with Anolis sagret and more remotely with the homolechis complex; there are certainly no grounds for postulating close affinity to the semilineatus-alutaceus set. Chamaclinorops barbouri and C. wetmorei are even more dis- tinet. The basic pattern of squamation is quite heterogeneous and yet upon this has been imposed a second pattern of enor- mously enlarged keeled dorsals and hugely enlarged keeled ventrals exaggerated beyond that seen in any other forms. This picture, like the apparent radiation of forms on Puerto Rico and the extraordinarily varied array of forms on the mainland, suggests strongly that the pattern — enlarged mid- dorsal zone, enlarged keeled ventrals — is one of several stereo- types that the anoles have again and again produced, that this is one of a limited set of squamation patterns possible to the anolines and therefore produced in parallel fashion in many times and places. It is this parallelism that contributes to the notorious ‘‘dif- fieulty’’ of the genus Anolis. Narrow groups are rather easy to recognize (though the specific and infraspecifie structure within bas | 196] ANOLIS SEMILINEATUS GROUP the zroup may be puzzling in the extreme) but wider relation- ships (at least when externals only are considered) are problem- atical, becoming obseurer with each step more distant from the species group. Origin of the semilineatus group The species of the semilineatus group are more uniform than the related Cuban series. They most resemble cyanopleurus, the middle term in the morphological series of Cuban forms. This species has its range in extreme eastern Oriente and is thus geographically closest to the Hispaniolan group. It therefore seems probable that the semilineatus series on Hispaniola has been rather recently derived from a cyanoplewrus-like Cuban ancestor but has been on Hispaniola long enough to achieve island-wide dispersal and moderate differentiation at the specific level. Acknowledgments I have had the advantage of discussions with A. Stanley Rand and with Dr. Richard Etheridge. The latter’s osteological evi- dence for species groupings within Anolis (unpublished thesis, University of Michigan) has in part confirmed, in part guided my own thinking on the wider relationships of Anolis species. The map-diagram was prepared by Patricia Grubbs. REFERENCES CITED MERTENS, R. 1939. Herpetologischer Ergebnisse einer Reise nach der Insel His- paniola, Westindien. Abhandl. Senckenberg. naturf. Ges., 449: 1-84. WETMORE, A. AND B. H. SwWALES 1931. The birds of Haiti and the Dominican Republic. Bull. U.S. Nat. Mus., 155: 1-483. Wiiiams, E. E. AnD A. S. RAND 1961. Notes on Hispaniolan Herpetology. 2. A review of the Anolis semilineatus group with the description of Anolis cochranae, new species. Breviora, No, 135: 1-11. No. 136 BREVIORA ‘soroeds 9014} [[@ IOF Sp10d01 AJI[BIOT [BNJOV IOF (TOGL) PuUvY PUv SUTBIT[IA Ul dew veg ‘poyvorpUT ST UvYy snonut}u09 AyIvou aloU 10 dATSUO}Xo9 o10M oq ABUT AY} ‘UOTJLVMIOFUT JUOsoId YIIM puUOdsoat10d 0} ‘payzepOsT 10 jyounfstp sv pejzuoseidait “ysviqUOd UT ‘aIe aDUDLYO0O ‘Pf puke WwOss}O *P JO SuOTINIAYSIp OY, ‘QOUITUBATOD JO 9YVS OJ A[UO SNONUTJUOD SB PoJMasoIdar ST snynaurjwas *P JO WOTYNQLAASIP OY, “snppaurjiuuas syoup Aq Bpotuedsty]T JO sSvul ULeVUT 9Y4 JO WOTSVAUT [VoTJOYZOUAY oy} JO tvISVIG, aeuesyo0o set snjzeaul||was Se re? Ss puneiedee wel abe ab nste eases 3 Sr] © SL 09 Sy OF StILIWOTIA JO FIWIS IVW SHL BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. AprRIL 10, 1961 NuMBER 137 NOTES ON HISPANIOLAN HERPETOLOGY 4. ANOLIS KOOPMANI, NEW SPECIES, FROM THE SOUTHWESTERN PENINSULA OF HAITI By A. STANLEY RAND Biological Laboratories, Harvard University Among the reptiles and amphibians collected for the Museum of Comparative Zoology in Haiti, with the support of an Ameri- ean Philosophical Society grant, during the summer of 1960, are seven specimens that appear to represent an undescribed species of Anolis. These lizards are small, moderately proportioned, dull colored in life, as well in preservative, and rather nondescript animals. In life the adult males possess a dark gray gular fan and an orange-pink chin and throat that distinguish them immediately from any other Hispaniolan Anolis. When this chin and throat color disappears in alcohol, the combination of seale characters distinguishes the species, but there are no unique characteristics. Dr. Karl Koopman, Assistant Curator of Mammals, Chicago Natural History Museum, provided financial assistance and encouragement that helped to make the collection of these specimens possible. In recognition of his aid the new species is called : ANOLIS KOOPMANI new species Type. MCZ 62541, adult male. Type locality. Carrefour Canon, 350 m. altitude, near Ducis, N. of Aux Cayes, Haiti. Collector. A. S. Rand and J. Lazell, 4 August 1960. Paratypes. Adult males, MCZ 62542-3; adult females, MCZ 62544-5 ; young males, MCZ 62546-7. All from Les Platons, 750 bo BREVIORA No. 137 m. altitude, above Carrefour Canon, Haiti, A. 8S. Rand and J. Lazell, 5 August 1960. Diagnosis. The presence of a zone of much enlarged middorsal scales, keeled head scales, and keeled, imbricate, and pointed ventrals distinguish this species from all the Hispaniolan Anolis except those of the semilineatus group (semilineatus, olssoni, and cochranae). It is distinguished from the latter in having 6-8 (not 10) enlarged middorsal rows, 5-8 (not 3-5) loreal rows, and 3-5 (not 1-3) scales separating the interparietal from the supraorbi- tal semicircles. It differs also in coloration, the males having pinkish-orange chin and throat. Description. (In the following description variations occurring in the paratypes follow, in parentheses, the condition in the type. ) Head. Wead seales strongly keeled. Frontal depression mod- erate. Scales across snout between second and third canthals 8 (7-9). Nares anterior to canthal ridge; separated from rostral by 1 scale. Canthal ridge distinct, not exaggerated, composed of 4 (4-5) large scales preceded by 2 (1-3) small ones. Second canthal largest, third next in size, first and fourth subequal. Posterior frontal subequal to (shghtly smaller than) anterior supraorbital ; separated from canthals by 2 (1-2) scales. Anterior supraorbital separated from canthals by 3 (2-3) scales. Supra- orbital semicircles separated by 3 (1-3) scales; separated from supraocular disc by one row of small scales (occasional narrow contact). Supraocular dise of 5-6 (5-7) enlarged keeled scales; separated from superciliary by 5-6 rows of granules. A single elongate superciliary. Interparietal scale slightly smaller than ear (14 to slightly smaller) ; separated from supraorbital semi- circles by 4-5 (3-5) seales. Seales in center of supratemporal area, granular, smaller than flank scales, smallest in center. Scales over temporal bar not (very slightly) enlarged. Temporal scales like supratem- poral seales. Suboculars broadly in contact with supralabials, separated from canthals (very narrow contact), not continued behind eye as a series of large scales. Supralabials to center of eye 5 (5-6). Loreal rows 5 (5-8). Loreal scales subequal in size. Mentals broader than long, in contact with 6 (5-6) throat scales posteriorly. No series of enlarged sublabials. Central throat scales small, elongate, keeled. Gular fan: Gular fan small. Seales slightly smaller than (sub- equal to) ventrals, keeled. Trunk: Middorsal seales much larger than flank seales, grad- 1961 ANOLIS KOOPMANI 3 ing into them. Rows of enlarged middorsals 6-8. Ventral scales larger than middorsals, imbricate, keeled. Limbs and digits: Scales on arms and legs larger than ven- trals, multicarinate. Hand and foot scales multicarinate dorsally. Lamellae under phalanges 2 and 3 of fourth toe 18 (17-19). Interdigital pads narrow. Tal: Tail round in cross-section. Verticils indistinct. En- larged postanal scales present in males. MEASUREMENTS Snout-vent Total Head Tibia Hind leg Sex MCZ # length length length length length male 62541 34mm 114mm 9mm l1imm 28mm (type) as 62542 39 135 10 12 33 as 62543 38 --- 10 12 31 62546 23 68 6 7 18 oe 62547 21 —- 6 7 17 female 62544 33 7 9 26 a 62545 33 109 8 10 26 In life a low nuchal and dorsal crest seems permanently raised (absent in females and young males). Color in life: Male, MCZ 62542, uniform gray-brown above; a whitish line, black edged above, from over shoulder to hind leg, indistinet for the posterior half of its length, below this line the flanks with seattered dark spotting. Belly light brown, chin and throat pale pinkish-orange, with a few scattered black spots, gvular fan scales colored like the chin, but the skin dark gray ; iris blue. Female, MCZ 62544, plain brown above, a middorsal stripe with a scalloped margin outlined with darker brown, a yellow stripe from below eye to over shoulder, continued faintly to hind leg; venter yellowish with faint dark spotting on throat; iris blue. - ae Habitat: The type was taken in a bush along a trail,at the edge of a coffee, grove. It was found at dusk among small twigs about three feet above the ground where it probably had climbed to spend the night. Of the six paratypes, three adults were found in heavily shaded coffee groves. All were on the ground among damp leaf litter. One juvenile was found six inches up in low herbaceous vegetation at the edge of the coffee grove. The others were purchased from a small boy for two cents each. Relationships: The relationships of this species are obscure ; 4 BREVIORA No. 137 it does not seem to be particularly close to any species now known either from Hispaniola or elsewhere. The only Anolis outside of Hispaniola that are at all similar to this species are the alutaceus, clivicolus, spectrum, cyanopleu- rus groups of Cuba, and this similarity lies primarily in the presence of a zone of enlarged middorsals and does not extend to other details. I interpret this as parallelism. Within Hispaniola, A. koopmani is superficially most like semilineatus and olsson. A zone of enlarged middorsals, keeled imbricate, pointed ventrals, keeled head scales, a lateral stripe, narrow digital expansions and small size occur also in semilineatus and olssoni and suggest a relationship to them. However, most of these characters are not unique to semilineatus and olssoni even in Hispaniola. The nature of the zone of enlarged middorsals (fewer rows that decrease in size laterally), the strong sexual dimorphism in color, the less attenuate body shape, and generally smaller scales all argue against this rela- tionship. Anolis ricord:, distichus, cybotes, armourt, shrevei, chloro- cyanus, coelestinus and Chamaelinorops wetmorei all have special- izations that seem to exclude them from close relationships with koopmant. The remaining species, Anolis monticola, darlington, christo- phet, hendersoni, baharucoensis and Xiphocercus darlingtoni are poorly known. It is possible that some if not all of them are closely related to one another and that koopmani’s relationships lie with these. However, until more information is available for this assemblage of species, particularly in regard to color in life, behavior and ecology, this hypothesis must remain very tentative. Acknowledgments: I am grateful to Dr. Ernest E. Williams for his advice and to Mr. James Lazell and M. Luc Whiteman for their assistance in the field. I wish also to thank M. Leonce Bonnfil fils and the other members of the Department of Agri- culture of Haiti who helped us in so many ways. BREVIORA Museum of Comparative Zoology CAMBRIDGE, MASs. JUNE 14, 1961 NUMBER 138 PFEIFFER’S UNFIGURED SPECIES OF STROPHOCHEILUS (MEGALOBULIMUS) By T. E. CRowLEY AND T. PAIN This paper is a supplement to J. C. Bequaert’s ‘‘Monograph of the Strophocheilidae, a Neotropical Family of Terrestrial Mollusks’’ (Bull. Mus. Comp. Zool., Harvard, 100: 1-210, 1948). When Dr. J. C. Bequaert published his monograph on the Strophocheilidae he was unable to deal in detail with two species described by Pfeiffer from specimens in the Cuming collection, now in the British Museum (Natural History), London. Neither of these had ever been figured, and the types are so far the only specimens known. Opportunity has, therefore, been taken to complete Dr. Bequaert’s monumental work with figures of Pfeiffer’s almost unknown species. In addition we are describ- ing and figuring S. (J/.) capillaceus (Pfeiffer), so far unfigured, and S. indigens Fulton. The authors wish to express their grateful thanks to the Brit- ish Museum authorities for permission to examine and photo- eraph the types for reproduction herein, to Dr. R. Zischka for specimens of S. (M.) indigens Fulton, to Drs. W. Blume and W. Weyrauch for the loan of material, and to Mr. S. P. Dance and Mr. J. A. Willson for their generous assistance in photo- eraphine specimens. STROPHOCHEILUS (MEGALOBULIMUS) HECTOR (Pfeiffer) Plate 1, Figure 1 Bulimus hector Pfeiffer 1857, Malak. Blatt., 4, p. 157 (Brazil); 1859, Monogr. Helic. Viv., 4, p. 367; 1868, Op. cit., 6, p. 11; 1876. On. cit., 8, p. 15; 1877, Op. cit., 8, p. 604. Paetel, 1889, Cat. Conch. Samml., 4th ed., 2, p. 212. Bulimus (Borus) hector von Martens 1860, in Albers, Die Heliceen, 2nd ~ 2) BREVIORA No. 138 ed., p. 192; 1876, Novit. Conchol., Abt. 1, 5, pts. 50-51, p. 21. Pfeiffer, 1879, Nomencl. Helic. Vic., p. 224. Strophocheilus (Thaumastus) hector Pilsbry 1895, Man. of Conch., (2) 10, p. 00. Thaumastus hector Pilsbry 1902, Man. of Conch., (2) 14, Classification, p. XX1. Strophocheilus (? Megalobulimus) hector Pfeiffer, Bequaert 1948, Bull. Mus. Comp. Zool., Harvard, 100: 118. Original description: ‘‘T. subimperforata, elongato-ovata, solidula, sub epidermide decidua, fulvida alba; spira coniea, apice rotundata; anfr. 6 convexiusculi, summi conferte capillaceo striati, ultimus spiram yix superans, plicato-striatus et obsolete decussatus ; columella leviter arcuata, non plicata; apertura sub- verticalis, acuminato-ovalis, intus albida, nitida; perist. album, marginibus ecallo albo junctis, dextro subincrassato, brevissime expanso, columellari superne dilatato, adnato. — Long. 71, diam. 35 mill., Ap. 36 mill. longa, 1914 lata.”’ New Measurements of Adult Holotype Greatest Aperture Aperture Length Width Length Width 71 mm. 39 mm. 35 mm, 26 mm. 6 whorls Specimen examined: Brazil (Miers Coll.), holotype (Brit. Mus. Nat. Taste.) Remarks. Bequaert (1948, p. 118), who had seen no speci- mens, was inclined to exelude hector from the Strophocheilidae, and Pilsbry (1895, p. 50) placed it in Thaumastus. Von Mar- tens (1876), however, suggested that it might be related to Strophocheilus (M.) oblongus (Muller). Careful examination of the type has convinced us that it is indeed correctly referred to the Strophocheilidae, being by reason of its nepionic sculpture a member of the subgenus Mega- lobulimus. It does not appear, however, to be in any way related to S. (M.) oblongus, the shell being longer and narrower in proportion, much paler and with a white lip. It is further- more much thinner, the apical sculpture finer, and is covered with a brown periostracum. S. (M.) hector would appear to us to be quite distinct from any other species of Megalobulimus so far known. 1961 STROPHOCHEILUS 3 STROPHOCHEILUS (MEGALOBULIMUS) COCAPATENSIS (Pfeiffer) Plate 1, figure 2 y= Bulimus cocapatensis Pfeiffer 1855 (August), Proe. Zool. Soc. London, (for 1855), p. 115 (Coeapata, Bolivia); 1859, Monogr. Helic. Viv., 4, p. 367; 1868, Op. cit., 6, p. 11; 1876, Op. cit., 8, p. 15. Paetel, 1889, Cat. Conch. Samml., 4th ed., 2, p. 209. Bulimus (Borus) cocapatensis Pfeiffer 1856 (January), Malak. Bla&tt., 2, (for 1855), p. 147. Von Martens, 1860, in Albers, Die Heliceen, 2nd ed., p. 192; 1876, Novit. Conchol., Abt. 1, 5, pts. 50-51, p. 9. Pfeiffer, 1879, Nomenel. Helic. Viv., p. 224. Strophocheilus (Borus) cocapatensis Pfeiffer, Pilsbry 1895, Man. of Conch., (2) 10, p. 20. Strophocheilus (Borus) cocopatensis Pfeiffer, Pilsbry 1895, Man. of Conch., (2) 10, p. 12; 1902, Op. cit., (2) 14, Classification, p. v. Misspelling of cocapatensis. Bulimus corapatensis Pfeiffer, Paetel 1889, Cat. Conch. Samml., 4th ed., 2, p. 10. Misspelling of cocapatensis, Strophocheilus cocopatensis Pfeiffer, Pilsbry 1930, Proce. Ac. Nat. Sci. Phila- delphia, 82, p. 855. Misspelling of cocapatensis. Strophocheilus (Megalobulimus) cocapatensis Pfeiffer, Bequaert 1948, Bull. Mus. Comp. Zool., Harvard, 100, no. 1, p. 126. Original description: ‘‘B. testa imperforata, ovato-oblonga, solida, minutissime decussata, sub epidermide virenti-fulvida violaceo-carnea; Spira convexo-conica, apice obtusa; sutura al- bida, irregulari; anfr. 514 superis radiatim costatis et minutis- sime granulatis, sequentibus peroblique descendentibus, parum convexis, ultimo spiram sub-aequante, basi rotundato; columella recedente, leviter arcuata; apertura subverticali acuminato- ovali, intus margaritacea; perist. Incrassato, breviter expanso, marginibus callo nitido junctis, columellari dilatato, adnato. Long. 67, diam. 30 mill.’’ MEASUREMENTS OF ADULT SHELLS Greatest Aperture Aperture Length Width Length Width 67 mm. 33 mm. 31 mm. 21 mm. 5% whorls. Holotype 67 33 31 20 51% whorls. Paratype 66 30 32 20 514 whorls. Paratype Specimens examined: Cocapata, Bolivia (Bridges Coll.), 3 types from the Cuming Collection (Brit. Mus. [Nat. Hist.]). Remarks. As pointed out by Bequaert (1948, p. 127), the radially ribbed and minutely granulated nepionie whorls, men- tioned in Pfeiffer’s original description, are characteristic 4 BREVIORA No. 138 of Megalobulimus, to which subgenus S. cocapatensis un- doubtedly belongs. Pilsbry (1930, Proc. Ac. Nat. Sci. Philadel- phia, 82, p. 355), in describing his S. carrikeri, infers that it may be related to S. cocapatensis, but we are unable to see any justifi- cation for this assumption. Bequaert suggests that it is not im- possible that cocapatensis may be the same as S. intertextus Pilsbry, but comparison of a specimen of the latter with Pfeif- fer’s type has convinced us that they are in no way related. The shell of cocapatensis is imperforate, long, thin and deli- eate, brown in color, with a very streaky, pink-flushed appear- ance. The spire is attenuated, the apex pointed, the mouth long and narrow, the aperture brown within, and the outer lip thin, white, shehtly reflected. Columella and callus white. Pfeiffer compared cocapatensis with S. rosaceus, but, as pointed out by Bequaert, the nepionic sculpture is typical of Megalobulr- mus and is not found in Chiliborus, to which subgenus SN. rosaceus belongs. Pfeiffer later 1856) placed it between ‘‘S. matthewsi’’ (= leucostoma) and S. capillaceus but, as he does not show it as being then in his collection, this opinion would seem of little value. STROPHOCHEILUS (MEGALOBULIMUS) CAPILLACEUS (Pfeiffer) Plate 1, figure 3 Bulimus capillaceus Pfeiffer 1855 (July), Proce. Zool. Soe. London, (for 1855), p. 93. Strophochetlus (Borus) capillaceus Pfeiffer, Pilsbry 1895, Man. of Conch., (2) 10, p. 31, Pl. 14, fig. 69. Strophocheilus (Megalobulimus) capillaceus Pfeiffer, Bequaert 1948, Bull. Mus. Comp. Zool., Harvard, 100, p. 120 (full synonymy ), Pl. 14, fig. 5. The type of S. (J.) capillaceus is in the British Museum (Nat. Hist.), from the Cumine Collection. It consists of three syntypes, of which that figured herein is now chosen as leetotype, no holotype having been selected by Pfeiffer. MEASUREMENTS OF ADULT SHELLS Greatest Aperture Aperture Length Width Length Width 64 mm. 38 mm. 37 mm. 24 mm. 5 whorls. Lectotype 60 39 35 30 ) whorls. Syntype 53 30 23 19 5 whorls. Syntype 67 40.5 40.5 22 5% whorls. Huanaco -—1/ yy 69 42 4] 21.5 5% whorls. Santa Ana on 1961 STROPHOCHEILUS Plate 1 Fig. 1. Strophocheilus (Megalobulimus) hector (Pfeiffer). Holotype. nat. size. Fig. 2. Sirophocheilus (Megalobulimus) cocapatensis (Pfeiffer). Holotype, nat. size. Fig. 8. Strophocheilus (Megalobulimus) capillaceus (Pfeiffer). Lectotype, nat. size. Fig. 4. Strophocheilus (Megalobulimus) maximus indigens Fulton. Apical aspect of immature shell, much enlarged. 6 BREVIORA No. 138 Specimens examined: ‘‘ Banks of the River Solimoes,’’ Peru (Cuming Collection). Near Santa Ana, Rio Urubamba, Peru, 3900 ft. (W. Weyrauch Coll., in Pain Collection). Huanaco, Peru (Pain Collection). Remarks. Bequaert (1948, p. 120) has dealt at length with the probable relationships and position in the subgenus of S. capil- laceus, and gave an excellent figure of it, together with a com- plete synonymy. To this very full account there is nothing which we ean profitably add. STROPHOCHEILUS (MEGALOBULIMUS) MAXIMUS INDIGENS Fulton Plate 1, figure 4; Plate 2, figures 5, 6 Bulimus kremnoicus d’Orbigny 1837, Voyage Amér. Méridion., 5, pt. 3, Moll., p. 300 (in part only: some specimens from Yuracare, Bolivia, the locality given in Explanation of Plates, p. 695, for fig. 3), Pl. 35, fig. 3 only. Not Helix kremnoica d’Orbigny 1835. Strophocheilus (Borus) maximus ? var. kremnoicus d’Orbigny, Pilsbry 1895, Man. of Conch., (2) 10, p. 16, Pl. 5, fig. 28 (copy of d’Orbigny’s fig. 3); 1902, Op. cit., (2) 14, Classification, p. iv. Strophocheilus (Borus) indigens Fulton 1914, Proe. Mal. Soe. London, 11, pt. 3, p. 165, fig. (Peru). Strophocheilus (Megalobulimus) indigens Fulton, Bequaert 1948, Bull. Mus. Comp. Zool., 100, No. 1, p. 98, Pl. 24, fig. 1 (copy of Fulton’s 1914 fig.). Strophocheilus indigens Fulton, Blume 1952, Arch. f. Mollusk., Frankfurt, 81, pts. 4-6, p. 105. Strophocheilus (Borus) kremnoicus subsp. vestitus Pilsbry 1926, Proe. Ac. Nat. Sci. Philadelphia, 78, p. 6 (Bolivia, probably in Dept. Cochabamba), Jeu Py infec Te Strophocheilus (Megalobulimus) maximus vestitus Pilsbry, Bequaert 1948, Bull. Mus. Comp. Zool., Harvard, 100: 94, Pl. 19, fig. 4. Original description: ‘‘Shell ovate-oblong, yellowish brown, moderately solid; spire about 13 mm. longer than the aperture ; whorls 614, apex smooth, the second and third whorls with prom- inent oblique plications, last two volutions polished and appar- ently smooth, but under the lens are seen to be finely granulated, the granulations being strong on the middle whorls and gradu- ally becoming weaker towards the aperture; the lower whorls have also some irregular and almost obsolete plications ; aperture sub-oval, whitish within; peristome thickened and very slightly expanded, white, margins joined by a moderately thickened white callus. Alt. 110, Diam. Maj. 47 mm. The nearest species to this is NS. (Borus) huascart Tschudi, which is broader, has a wider aperture, a rougher and duller surface, and its apical plieations are much finer and closer together than in indigens.’’ ~ 1961 STROPHOCHEILUS MEASUREMENTS OF ADULT SHELLS Length Greatest Aperture Aperture Width Length Width Whorls 135 mm. 57.5mm. 60mm. 41mm. 7 Bolivia: Sacha, Yungas, 800-1500 m. (Bavarian State Mus. ) 132 63 59.5 37 7 SG BG 127 58 56 39 7 oe Be 128 58 55 Olle 61% Bolivia (W. Weyrauch Collection ) 118.5 61 57.5 40.5 6 Bolivia: Chapare, 400 m. (Bavarian State Mus.) 118 58 56 24 — Type of vestitus Pilsbry 116 48 47 31 6% Holotype of sndigens, Peru 101 46 47 26 5% Bolivia (T. Pain Collection ) 100 49 48 27.0 5% Bolivia (W. Weyrauch Collection ) Specimens examined: Peru, type of indigens Fulton (British Museum [Nat. Hist.], No. 1915-1-5-199). Yungas de Palmar, 1200 m., Bolivia (R. Zischka Coll., in W. Blume, T. Pain, and W. Weyrauch Collections). Remarks. Comparison of the type of indigens, together with the shells from Bolivia, with Bequaert’s (1948) figure of vestitus Pilsbry, leave us in no doubt that they are identical. All show the strong, prominent oblique plications on the second and third whorls, noticeably absent on both the typical maximus and the subspecies huascart. As pointed out by Bequaert (1948, p. 94), indigens (=vestitus) is of considerable interest in that it bridges the gap between typical maximus and subspecies huascari in re- spect of its relatively wider spire, narrower body-whorl and smaller mouth. Fulton, who described indigens from a unique holotype, did not apparently connect it with maximus, although he drew at- tention to its close resemblance to huascart. Bequaert (1948, p. 94), dealing with vestitus, makes no men- tion of the prominent sculpture, although this is easily recog- nized in his excellent photograph of the shell he selected as holotype from Pilsbry’s type set (Ac. Nat. Sci. Philadelphia No. 138105). Dr. Weyrauch informs us (7m litt., 1958) that he considers the elongated shell from Oxapampa, Peru, referred by Bequaert to vestitus, to be a typical maximus on account of the aperture being much longer than in indigens (= vestitus). A similar elongated shell from Peru, without more definite locality, (os) BREVIORA No. 138 kindly sent by Dr. Weyrauch, shows traces of a dark periostra- eum and, although much worn about the spire, has the long aper- ture characteristic of typical maximus. S. (M.) maximus indigens Fulton has not so far been ob- tained in Peru by Dr. Weyrauch, but, from the similarity of the fauna of southeastern Peru and northeastern Bolivia, there can be little doubt that indigens occurs also in Peru. ane nh ek ae ne a tt Plate 2 Fig. 5. Strophocheilus (Megalobulimus) maximus indigens Fulton. Hol- otype, nat. size. Fig. 6. Strophocheilus (Megalobulimus) maximus indigens Fulton. Yungas de Palmar, Bolivia, nat. size. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. June 15, 1961 NumBeEr 139 A NEW SPECIES OF SPHAERODACTYLUS FROM NORTHERN HAITI By James D. LazEeuu Although the genus Sphaerodactylus on Hispaniola is sut- ficiently diversified and confused to warrant at least a partial revision, the species here described is so remarkably different from any other form that I am confident in naming it at this time. The new species is named for Mr. Benjamin Shreve of the Museum of Comparative Zoology for his current work on the sphaerodactyls of Hispaniola. SPHAERODACTYLUS SHREVEI Sp. nov. Type: MCZ No. 62548, Mole Saint Nicolas, Haiti. Coll.: J. Lazell and A. 8. Rand, 16 July, 1960. Diagnosis: The combination of the following characters serves to distinguish this species from any other found in the Antilles: the presence of a highly convex snout as seen from the side (loreal region also somewhat convex) ; very large keeled dorsal scales beginning at the level of the axilla; and the paravertebral arrangement of these dorsals, producing a middorsal zone not of granules but of small and large, irregularly placed scales. Description of type. Snout short. Eye nearer tip of snout than ear. Snout, as seen from the side, highly convex. Loreal region also somewhat convex. Rostral large with a partial median cleft. Nostril between rostral, first supralabial, a large supranasal (= internasal) and two small postnasals. A single scale between the supranasals (= internasals), which border the rostral posteriorly. Granular scales on top of snout somewhat larger than interocular or nape scales. Four supralabials, of 2 BREVIORA No. 139 about equal length, to the center of the eye, followed by two smaller ones. Four infralabials gradually decreasing in size followed by two abruptly smaller ones. Mental short, wider than long, bordered posteriorly by two postmentals. Supraciliary spine small. Squamation of head and neck granular to the level of the shoulder; at that level a moderately rapid change to large, flattened, heavily keeled dorsals. Twenty-five dorsals from the level of the axilla to the posterior level of the hind limb. Five dorsals in the standard distance. A very ill-defined middorsal zone of smaller keeled scales not forming a continuous row but with the large dorsals meeting irregularly along the middorsal line. Middorsals subimbricate or not imbricating, the laterals more distinctly imbricating. Throat scales smooth, granular, juxtaposed. Chest and belly scales larger, smooth, eycloid, broadly imbricate, about nine ventral scales in standard distance. Seales of anterior surfaces of limbs imbricate, cycloid, smooth, somewhat smaller than ventrals. Seales of posterior surfaces of limbs granular, smooth. Digital pads approximately twice as broad as the subdigital lamellae. Ten infradigital lamellae under fourth toe. The type, the only specimen yet col- lected, lacks the tail. It is a female and the eseutchecn there- fore cannot be described. Coloration in life. S. shrevei is a dull-colored animal with a pattern composed of three basic hues—each tendimg to be unique on an individual scale. There are very irregular dark grey-brown blotches across the dorsum; beginning at the back of the head there are three such markines to the shoulders. There are three more crudely ‘‘Y’’ shaped markings on the body, the most anterior of which bifurcates to the right, the remaining ones bifurcating to the left. There are two small blotches on the right side of the rump and one on the left. The second transverse blotch, on the nape, is broken by a lght middorsal line that continues down through the fourth marking and then fades out. The ground color of the dorsum is ash grey. There are scattered over the dorsal surface short transverse series of white or partly white scales — from two to four in a row — that appear to have no correlation whatever with the rest of the animal’s pattern. The top of the head is ash grey except for a very irregular, dark, grey-brown blotch on the parietal area. Coming back from the eye are two stripes, one of which runs downward across the cheek; the other nearly connects with the first transverse marking on the back of the head. Not 1961 SPHAERODACTYLUS SHREVEI 3 including the stripe across the cheek, there are five vertical dark markings across the pale labials, the anterior two of which are connected at the edge of the mandible. There are dark streaks on the lateral edges of the chin and throat, the underside of the hind limbs, and across the venter just anterior to the anus. The ventral surface is white; a close examination reveals that on each scale there are tiny black dots. This peppering becomes more noticeable laterally and posteriorly. Along each side of the animal is a line of partially connected, small, dark, grey- brown blotches; just ventral to this row is another composed of widely spaced, single, dark, grey-brown scales. All three of the animal’s hues: white, grey and grey-brown, are simply variations in the intensity of speckling on each scale with tiny black or brown dots. The pattern of the animal bears no resemblance to that of the young or females in the species to which it has been com- pared, or to any other Sphaerodactylus I have seen. Habitat. The type specimen was taken from a large circular rock pile about two-and-one-half feet deep; this sort of rock pile is the result of removing the debris from a heap of charcoal after burning, and is composed of rocks that vary in size from that of a golf ball to nearly the size of a football. This particu- lar heap was of some age, for even in the arid terrain of Mole Saint Nicolas several fair-sized thornbushes had sprouted up in it. Collecting was very difficult, for any animal uncovered could generally manage to dart back into the pile before the collector could safely ascertain that it was not a scorpion or some other unpleasant handful. In order to get best results we excavated areas through the pile, dividing it up into more manageable smaller piles; this system netted Celestus, Typhlops. and Tropidophis, as well as the type of Sphaerodactylus shrevei. Another specimen of apparently the same species escaped, leav- ing only its tail behind. Due to the relative inaccessibility of peninsular north-western Haiti it may be some time before additional specimens can he obtained. Comparisons. From S. copet, the only other comparably large- scaled Hispaniolan form, S. shrevei differs in the following characters: (1) Snout seen from the side highly convex; (2) No middorsal zone of granules; (8) Dorsal scales flatter, not swollen, apt not to imbricate, especially in the middorsal area ; (4) Pattern a series of irregular dark dorsal blotches with 4 BREVIORA No. 139 a line of often connected smaller blotches along each side; dorsal blotches broken by a light middorsal line anteriorly. Krom S. scaber of Cuba, S. shrevei differs in all the men- tioned characters and in snout length, which averages slightly shorter in the Cuban form. S. samanensis Cochran of the Dominican Republic resembles S. shrevet somewhat in squamation but the dorsals are smaller and the ventrals larger. There are no smaller middorsal scales and the pattern is very different. S. shrever ditfers from S. beck: of Navassa again in the absence of a middorsal zone of granular scales and in the flatness of the dorsal scales, which are rounded, swollen, and rather tubercular in S. beck. Two Jamaican forms, S. richardson and S. parkeri, ocea- sionally possess dorsal squamation similar to that of S. shrevei in that while there is no middorsal zone of granules there may be small scales irregularly scattered along the middorsal line, but in general the arrangement of the scales is much more regular and not of a paravertebral nature. From both of these species S. shrever ditfers in the following characters: (1) Snout seen from the side highly convex; (2) Head granules extending posteriorly to level of axilla instead of just onto nape; (3) No enlarged, clearly defined canthal scale; (4) Pattern and colora- tion. From 8S. parkert it differs also in having only a single small scale between the internasals. There is a vaguer resemblance to S. shrever in some Lesser Antillean forms. S. vincenti and S. microlepis, for example, show a tendency towards reduction in size of the middorsal scales, but in these forms the pattern of squamation tends to be very regular and the scales are much smaller. The new species has been compared with these forms largely because there are apparently no closer ones, although it bears little resemblance to any of them. Its relationships at the moment are not at all clear. Acknowledgments. My thanks are due to Mr. A. S. Rand for help in collecting the specimen described, to Mr. B. Shreve tor checking the description and comparisons made and to Dr. E. EK. Williams for the loan of comparative material and for readiny the manuscript. A grant from the American Philo- sonhical Society supported the expedition which resulted in the discovery of S. shrevet. 1961 SPHAERODACTYLUS SHREVEI 5 REFERENCES BaRsBour, T. 1921. Sphaerodactylus. Mem. Mus. Comp. Zool., 47: 217-277. CocHRAN, D. M. 1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177: 1-398. RUIBAL, R. 1959. A new Sphaerodactylus from Oriente, Cuba. Herpetologica, 15: 89-93. TABLE 1 Comparisen of the snout length ratios of six species of Sphaerodactylus. The ratio is obtained by dividing the standard distance (i.e., the distance from the center of the eye to the tip of the snout) into the distance from the center of the eye to the ear opening. The higher the figure obtained the shorter the snout. Specimens Mean Value Range S. richardsoni 5 WI .67- .81 S. parkert 3 .76 .73- .78 S. copet 10 76 .70- .80 S. shrevet 1 80 S. scaber 6 82 ale ey S. becki 4 90 -78-1.00 ree > ahd k Gry ad eS ae ARAN Gall Shah, Leni 2 aa eaten Linea, i 4 ie oe ee eb eal al Bad speeangallad vay hntlh sant p8dee 2 b | ies Oe Ga icaeitns ita : > Ay aT tr 6 1 ie, “ee Pan , gm uited! ye wits) ut) ten) oes ’ ph Pith) pA Fa Windy Whore flaaacania eee pana nnn! alt ilies! “Diam eet) @f tociatty ah etn ie DI GAP VB Ch mew ae OEE ti~ = ibosttin, soegieds qnayé o@F geldyr tiene: doin Don... rea Y pean ale J GAL Mma ¢ hote* ar ae a | by ie Wh eye @ oY Eby. Gun, ‘8 St ri 4 ‘ ibe | Ms i : _ _— rae | pei ia) : 2 7 Pet i. ee, ete ‘ ae , | :- ailiiend tens ; ae se wet " j ‘ ; t is a ja oY ,; , . : ee St. ~ dl es Uw a) eae J: ee _ "i we _ S 7 . Aa: 4)! 2 of Sa aes rene? tis 1s OS sae Pyle Ar (} i ar j oe ij a ae Bi Ie ety) ae dive Un | Hb(hd Fel) } Ue = f | a 7 ; » BaP e 1p Qa Or - iy 4 Cem % 5 BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. JUNE 27, 1961 NuMBER 140) A PRELIMINARY REVIEW OF THE NEARCTIC SPECIES OF SIEROLOMORPHA (HYMENOPTERA) By Howarp E. Evans In the course of collecting Bethylidae and examining material in various museums, I have encountered a good many specimens of the curious bethylid-like genus Sierolomorpha Ashmead. Some of these specimens are so different from the type, ambigua, for a long time the only known species, that it seems desirable to provide names for them. Krombein (1951, U. S. Dept. Agri. Monogr. 2, p. 748) has already pointed out that there appear to be several species in North America. The present paper does not pretend to be an exhaustive treatment of this genus. Names are provided for several of the more distinctive species, but before a definitive revision is possible more collecting must be done and more detailed studies made of structural details and their vari- ation. Actually, Sierolomorpha is not a bethylid or even especially close to the Bethylidae. Schuster (1949, Ent. Amer., 29: 124) is correct in pointing out its close similarity to certain Mutillidae and Tiphiidae, and the arrangement in the Synoptic Catalog (Krombein, 1951, op. cit.), in which the genus is placed in a monogeneric family between the Tiphiidae and Mutillidae, is probably the best that can be achieved at present. Sierolomorpha is related to the most primitive Scoloidea, and may not be far from the stock which gave rise to the Bethylidae. However, in virtually all structural details it stands very much closer to the Tiphiidae than to the Bethylidae. Only one species of the genus, hospes Perkins, has been de- seribed from outside the Nearctie region (from Hawaii). Krom- bein has suggested that this species may have been introduced from North America. In the collection of the U. S. National Museum there is a single male of an undescribed species from 2 BREVIORA No. 140 Panama. I have seen no other specimens from outside the United States and Canada. The most useful structures for the separation of species in this genus appear to be the antennae, especially the tyloides of the male (Figs. 1-6), the propodeum, and the first two abdominal tergites. There seems to be little variation within the genus with respect to the mandibles, clypeus, and most details of the thorax including the legs and wing venation. I have therefore made little mention of these structures in my descriptions. I have studied the male terminalia of selected specimens and found some minor variation, particularly in the volsellar cuspis. However, the differences are so slight and subtle that further study to determine the extent of individual variation did not seem war- ranted. I have therefore made no mention of characters of the terminalia in the keys and descriptions. The following abbreviations are used for the museums and private individuals that supplied material for this study: AMNH, American Museum of Natural History, New York; CIS, Cali- fornia Inseet Survey, Berkeley; CNC, Canadian National Col- lections, Ottawa; HKT, Henry K. Townes, Ann Arbor, Michi- n; MCZ, Museum of Comparative Zoology, Cambridge, Mass. ; UCD, University of California at Davis; USNM, U. 8S. National Museum. Key to Species Females } 1. Abdomen with the constriction between the first two tergites strong, the first tergite with a weak to strong apical trans- verse depression which is longitudinally striate ; propodeum with a moderately wide median groove which is irregularly margined by carinae. . . a 42 Abdomen with the constriction peer een ae one “ane senses weak, first tergite smooth, not depressed or striate apically ; propodeum with median eroove absent or linear and not margined as above... . lest Pte 2. Legs beyond the coxae bright allo ne OR: pasal segments of antennae suffused with light yellowish-brown; notauli strongly diverging anteriorly (Florida and Arizona to Alberta, Ontario, and Massachusetts) . : canadensis G@azovenchen) 1 The females of two species, apache and brevicornis, are unknown. 1961 NEARCTIC SPECIES OF SIEROLOMORPHA 3 Legs brown except front tibiae and tarsi yellowish-brown; antennae brown, sometimes weakly suffused with paler brown on sides of basal sezments; mesoscutum rather flat, notauli weakly diverging anteriorly (California, Arizona, and Colorado to Saskatchewan and Yukon) . nigrescens N. sp. Front, pronotum, and mesopleura with strong punctures; pro- and mesonota bright rufocastaneous (Arizona ) bicolor n. sp. Front, pronotum, and mesopleura with only minute, widely spaced punctures; thorax entirely black (Georgia to Con- mechicut and Kansas)’. < 2 a4. = .ssemilisi nesp) Males Abdomen with the constriction between the first two tergites strong, the first tergite with a weak to strong apical trans- verse depression which is longitudinally striate; tyloides present on antennal segments seven through ten, short and rather prominently projecting (Figs. 1, 2); propodeum as in female (see couplet 1 of key to females) . ... 2 Abdomen with the constriction between the first two tergites weak or absent, the first tergite smooth, not depressed or striate apically; tyloides not present on antennal segment seven or, if so, very ee and low on segments eight and: nine 460% : st ialass Tibiae and tarsi nasi or sities light cre ini te ae own in most specimens; temples rather short, as seen from above much shorter than eyes (Florida and Arizona to Alberta, Ontario, and Massachusetts) . . . canadensis (Provancher) Legs brown except front tibiae (sometimes also front tarsi and middle tibiae) light yellowish-brown; temples strongly developed, as seen from above nearly as wide as the eyes (California, Arizona, and Colorado to Saskatchewan and VAL) We eee - = « Magrescens n.'sp. Antennae eee short and nian segment eleven about 1.2 X as long as thick, segment thirteen about 1.5 X as long as thick; tyloides present on antennal segment eleven (Fig. 3); head extremely broad, about 1.25 X as wide as high (SouthsCarolina)) (2) 2) +2 3 ‘brevmecorms nosp. Antennae elongate, segment eleven about twice as long as thick, segment thirteen much more than twice as long as thick; tyloides not present beyond segment ten; head alee ASW GevaNm bdo a (eps oe) Cement! cots, Ma fl 4 BREVIORA No. 140 4. Ocelli distinctly enlarged (diameter of anterior ocellus about 22 X minimum width of front); tyloides present on antennal segments seven through ten (Fig. 6); front with a strong median groove in front of anterior ocellus CAriz0na)) | ae a2 . /4 “a apache nssp: Coals not enlarged (Gate a sobenton ocellus Jess than 22 X minimum width of front); tyloides present on seg- ments eight through ten (Figs. 4, 5); front with a weak median impression ifianyie, — A io = Soe eo 5. Punctures of front moderately strong; tyloides rather short (Fig. 5); middle and hind tibiae and tarsi dull brown GAwizona) . -. ie 2 fe bieoloram: ‘sp: Punctures of Seach very weak: Paloides elongate (Fig. 4) ; all tibiae and tarsi light yellowish-brown (Georgia to Connecticut and Kansas). 2). . : =... similis musp: SIEROLOMORPHA BICOLOR new species Holotype. 9, ARIZONA: Southwestern Research Station, 5 mi. W. of Portal, Cochise Co., 9 August 1959, 5400 feet eleva- tion (H. H. Evans) [MCZ]. Description of type female. — Length 5.8 mm., length of fore wing 4.3 mm. Head black; pronotum, mesoscutum, and seutellum wholly bright rufo-castaneous; remainder of thorax and pro- podeum black; abdomen very dark brown, approaching black basally; mandibles light brown, darker basally and apically; elypeus suffused with reddish-brown apically; scape black, flag- ellam dark brown, outer side of apical segments paler; legs dark brown except front and middle tibiae and all tarsi light brown; fore wing lightly, uniformly infuseated, veins and stigma dark brown. First four antennal segments in a ratio of about 12:5:7:9, segment three 1.4 X as long as thick. Front strongly polished, punctures small but rather strong; spacing of punctures rather irregular, those on the sides being mostly rather close, often not much more than their own diameters apart, those on the middle of the front rather sparse; temples also with distinct punctures. Head subcircular in anterior view, vertex evenly rounded off a short distance above eye tops; inner orbits sub- parallel, minimum width of front about .9 the eye height. Ocelli small, in a broad triangle, the front angle greater than a right angle; postocellar line slightly exceeding ocello-ocular line. Thoracic dorsum strongly polished, non-alutaceous; pro- notum with strong, widely spaced punctures; mesoscutum with 1961 NEARCTIC SPECIES OF SIEROLOMORPHA 5 a very few punctures on the sides, impunctate medially. Pro- podeum strongly polished medio-basally, elsewhere slightly roughened by obscure punctures; median line weakly impressed. Mesopleurum polished and with well-defined punctures. First abdominal tergite without a subapical depressed and _ striate band, completely smooth; constriction between first and second tergites weak; abdomen rather strongly hirsute beyond segment three. Male (assigned here tentatively).— ARIZONA: Cochise Stronghold, Dragoon Mts., 4850 feet elevation, oak-juniper zone, 2 July 1947 (Werner & Nutting) [Univ. Arizona]. Description of male. — Length 5 mm., length of fore wing 4.8 mm. Head and thorax entirely black, abdomen dark brown; apical half of mandibles hght brown; antennae wholly dark brown; legs wholly dark brown except front tibiae and tarsi bright yellowish-brown; wings subhyaline. First four antennal segments in a ratio of about 20:8:13:20, segment three 1.5 X as long as thick, segment four 2.2 X as long as thick, this segment typical of the remaining segments except the last ; segments eight through ten each with a short, rather weak longitudinal polished ridge (Fig. 5). Front polished, wholly covered with small but well-defined punctures which are separated by scarcely more than their own diameters; vertex and temples more weakly punctate; front somewhat impressed along the inner orbits and beside the posterior ocelli, faintly impressed medially. Minimum width of front approximately equal to eye height; ocelli in a broad, flat triangle, postocellar and ocello-ocular lines subequal ; ocelli of moderate size, diameter of anterior ocellus .18 X mini- mum width of front. Pronotum short, shining and with dense, rather weak punctures. Mesonotum shining, rather sparsely and weakly punctate. Propodeum polished, impunctate, and non- alutaceous over most of its surface. Mesopleurum polished, densely but weakly punctate. First abdominal tergite smooth, without a subapical depressed and striate band. Remarks. — The type female was taken on the ground beneath oak trees in dry, open forest. The male associated with it tenta- tively was apparently taken in the same type of forest and at nearly the same altitude. SIEROLOMORPHA APACHE new species Holotype. — 6 ARIZONA: 3-5 mi. SW of Apache, Cochise Co., 8 August 1959, about 4300 feet elevation (H. E. Evans, on Baccharis glutinosa) [MCZ]. 6 BREVIORA No. 140 Description of type male.— Length 4.5 mm., length of fore wing 4.0 mm. Entire body dark brown, the head nearly black; mandibles light brown; antennae uniformly light brown; legs dark brown except tibiae and tarsi light brown, the front tibiae a rather bright yellowish-brown; wings hyaline. First four an- tennal segments in a ratio of about 15:9:13:17, segment three 1.6 X as lone as thick, segment four 2.0 X as long as thick, segment eleven 2.1 X as long as thick; segments seven through ten each with a weak longitudinal polished ridge (Fig. 6). Front polished, with weak, scattered punctures, with a very strong median groove extending downward from the anterior ocellus; eyes large and prominent, inner orbits subparallel below; head about 1.2 X as wide as high. Minimum width of front 1.07 X eye height; ocelli large, diameter of anterior ocellus .22 X mini- mum width of front; postocellar line 1.15 X ocello-ocular line. Pronotum short, shining, weakly punctate. Mesonotum strongly shining, obscurely punctate. Propodeum in large part strongly polished, median area without ridges or other sculpturing except for a simple carina on the posterior third. Mesopleurum shining, obscurely punctate. Abdomen with scarcely any indication of a constriction between the first two segments either dorsally or ventrally ; first tergite without an apical striate depression. Remarks. — This striking specimen was taken on vegetation in the daytime, although the large ocelli suggest that the species may be nocturnal or crepuscular. The locality was a dry wash in an area of desert grassland. SIEROLOMORPHA SIMILIS new species Holotype.— 6, MARYLAND: Takoma Park, 22 Sept. 1945 (H. & M. Townes) [HKT]. Description of type male. — Length 4.4 mm., leneth of fore wing 4.1 mm. Body dark brown, head and thorax almost black ; mandibles light brown; antennae uniformly dark brown; coxae dark brown, femora medium brown, paler apically, tibiae and tarsi yellowish-brown; wings lightly tinged with fuscous. First four antennal segments in a ratio of about 19:10:13 :18, segment three almost twice as long as thick, segments four and eleven about 2.1 X as long as thick; tyloides in the form of low carinae on segments eight through ten, the carina on eight extending for much of the leneth of the segment, the others shghtly shorter (Fig. 4). Front strongly polished, with small, rather evenly distributed punctures which are separated by 1-2 X their own be | 1961 NEARCTIC SPECIES OF SIEROLOMORPHA diameters; median line of front weakly impressed. Head 1.12 X as wide as high, subcircular in anterior view, temples only moderately developed, contracted immediately behind eyes. Inner orbits convergent below, minimum width of front subequal to eye height ; ocelli of moderate size, diameter of anterior ocellus .17 X minimum width of front; postocellar line 1.1 X ocello-ocular line. Pronotum strongly shining, with a great many minute punctures. Mesoscutum strongly shining, obscurely punctate, notauli strong, nearly reaching anterior margin. Dorsal surface of propodeum shining, with a linear median groove. Mesopleurum strongly polished and nearly impunctate. First abdominal tergite polished, smooth, with no evidence of a transverse apical de- pression; second tergite with a narrow transverse basal impres- sion, so that there is vague evidence of a constriction between the first two tergites. Allotype.— °, Kearny, New Jersey, 9 Sept. 1935 (C. W. Funaro) [AMNH]. Description of allotype female. — Length 5.1 mm.; length of fore wing 3.6 mm. Head and thorax dark brown, abdomen medium brown; mandibles and apical half of clypeus lght yellowish-brown ; antennae dark brown except scape and pedicel suffused with yellowish-brown and flagellum lght brown be- neath; coxae brownish but legs otherwise wholly bright yellow- ish-brown ; wings lightly tinged with fuscous. First four antennal segments in a ratio of about 23:10:11:15, segment three 1.5 X as long as thick, segment eleven 1.7 X as long as thick. Front strongly polished, punctures small and widely separated Head subcireular in anterior view, about as wide as high; inner orbits subparallel, minimum width of front subequal to eye height. Ocelli small, in a broad triangle; postocellar line subequal to ocello-ocular line. Pronotum polished, with scattered small punctures. Mesonotum impunctate, notauli strong on posterior 8 of mesoscutum, diverging and attenuate anteriorly. Pro- podeum mostly smooth and shining, with a very thin median eroove which posteriorly is paralleled by some irregular carinae. Mesopleurum convex, smooth and shining. Abdomen fusiform, depressed ; first tergite smooth and polished, with no evidence of a transverse apical impression; second tergite barely depressed basally. Paratypes. — CONNECTICUT: 1 ¢@, Bank of Conn. River, East Hartford, 2 Sept. 1947 (H. E. Evans) [MCZ]; NEW YORK: 1 2, Sea Cliff, Long Island [MCZ]; MARYLAND: 3 8 ¢, Takoma Park, same data as type [MCZ, HKT]; WEST 8 BREVIORA No. 140 VIRGINIA: 1 ¢, Shaver’s Fork, Tucker Co., Oct. 1988 (G. H. Wallace) [Carnegie Mus.]; SOUTH CAROLINA: 1 ¢, Green- ville, Oct. 1952 (L. & G. Townes) [HKT]; GEORGIA: 1 ¢, Macon, 1 Dee. 1923 (T. H. Hubbell) [USNM]; KANSAS: 3 6 6, Manhattan, Sept., Nov. (D. A. Wilbur, T. F. Winburn) | Kansas State Univ. ]. Variation. — The two female paratypes are very similar to the allotype. The Connecticut specimen has the basal two antennal segments bright amber-colored, contrasting strongly to the flagel- lum; the Long Island specimen is without a head. The males show a small amount of size variation; the smallest specimen (Manhattan, Kansas) has a fore wing measuring 3.6 mm., the largest (Takoma Park, Md.) has a fore wing measuring 4.4 mm. The Kansas specimens tend to have the body (especially the abdomen) slightly paler in color, but otherwise little variation in color or body sculpture can be noted. In most specimens the postocellar line is subequal to the ocello-ocular line, and in one specimen it is somewhat shorter. SIEROLOMORPHA BREVICORNIS new species Holotype.-— 6, SOUTH CAROLINA: Greenville, 21 Sept. 1952, Gus Ge Downes) || EEK |: Description of type male.— Leneth 3.4 mm., length of fore wing 2.7 mm. Body dark brown, head almost black; mandibles light brown on apical half; antennae dark brown, very slightly paler beneath; legs dark brown except front tibiae and tarsi light yellowish-brown; wings hyaline. First four antennal seg- ments in a ratio of about 12:6:7:8, sezment three 1.1 X as long as thick, apical segment unusually short and thick, about 1.5 X as long as thick; tyloides present on segments nine through twelve, but rather small (Fig. 3). Front strongly polished, punctures minute, shallow; median line strongly impressed in front of anterior ocellus. Head very broad, 1.25 X as wide as high: front broad, its minimum width .61 X width of head, 1.18 X height of eye; ocelli small, diameter of anterior ocellus 14 X minimum width of front; postocellar Ine very shghtly greater than ocello-ocular line. Vertex forming a broad, even are above the eye tops; temples moderately developed, in dorsal view about two-thirds as wide as eye. Pro- and mesonota shining, obscurely punctate; notauli strong, complete, diverging ant°rior- ly. Propodeum mostly smooth and polished, med’an line weakly grooved, ecarinate. Mesopleurum strongly shining, obscurely 1961 NEARCTIC SPECIES OF SIEROLOMORPHA 9 punctate. Venation differing from that of other species only in having the second recurrent and second transverse cubital veins very weakly indicated and the margin cell somewhat more rounded apically. First two abdominal segments without a con- striction between them, first tergite smooth and polished, without a striate depression along its apical margin. STEROLOMORPHA NIGRESCENS new species Holotype. — 6, WASHINGTON: Olympia [USNM]. Description of type male. — Length 4.4 mm., length of fore wing 3.7 mm. Body dark brown, almost black; mandibles light brown; antennae uniformly dark brown; legs dark brown except front tibiae bright yellowish-brown; wings lightly tinged with fuscous. First four antennal segments in a ratio of about 20:8 :11:16, seement three only 1.3 X as lone as thick, segment four about 1.7 X as long as thick; segments seven through ten each with a short but rather strong longitudinal ridge (Fig. 1). Front strongly polished, with minute punctures which are rather elose together below, much more widely scattered above; median line of front weakiy impressed. Head about 1.15 X as wide as high, rather thick, seen from above with the temples nearly as thick as the eyes, the head across the temples nearly as wide as across the eyes. Inner orbits subparallel below; minimum width of front 1.13 X height of eye; ocelli small, diameter of anterior ocellus .14 X minimum width of front; postocellar line subequal to ocello-ocular line. Pronotum strongly shining, obscurely punctate. Mesoscutum polished and nearly impunctate, with very strong notauh which diverge slightly anteriorly and do not quite reach the anterior margin. Propodeum with two median earinae between which it is somewhat grooved, disc otherwise with weak and irregular sculpturing, somewhat shining. Meso- pleurum shining and with very small punctures. Abdomen with a strong constriction between the first two segments; first tergite depressed along the posterior margin, second tergite along its anterior margin, both depressions with longitudinal striations. Allotype. — 2, WASHINGTON: Seattle [USNM]. Description of allotype female. — Length 4.5 mm., length of fore wing 3mm. Head and thorax dark brown, abdomen medium brown, somewhat darker apically; mandibles and apical half of elypeus light brown; antennae dark brown, suffused with lighter brown on the sides of the basal flagellar segments; legs dark brown except front tibiae bright yellowish-brown; wings lightly 10 BREVIORA No. 140 tinged with fuscous. First four antennal segments in a ratio of about 22:9:9:13, segment three 1.2 X as long as thick. Front strongly polished, punctures minute and widely scattered. Head subeircular in anterior view, very slightly wider than high; inner orbits subparallel, minimum width of front 1.16 X height of eye. Ocelli small, in a broad triangle; postocellar line 1.1 X ocello-ocular line. Pro- and mesonota polished, obscurely pune- tate; notaul deeply impressed, diverging only slightly anterior- ly, terminating well short of anterior margin of mesoscutum. Propodeum and mesopleurum as described for male. Abdomen fusiform, shining; first tergite with a transverse apical depres- sion which is strongly longitudinally striate. Paratypes. — WASHINGTON: 1 ¢, same data as type [USNM]; 1 ¢@, Almota, 24 June 1911 [MCZ]; IDAHO: 1 4, Harvard, 24 June 1935 (J. M. Beck) [USNM];2 ¢ 6, Moscow, June, July [USNM];1 4, Nezperce, 18 June 1935 (J. M. Beck) [USNM]; UTAH: 1 ¢, Uinta Co. (G. E. Wallace) [Carnegie Mus.|]; CALIFORNIA: 1 ¢, Sagehen, nr. Hobart Mills, 21 June 1954, on Phacclia hunulis (P. D. Hurd) [CIS]; 1 4, Carnelian Bay, Lake Tahoe, 17 June 1958 (R. M. Bohart) [UCD]; 4 ¢ 4, Pasadena, April 1944 (K. W. Cooper) [USNM];1 ¢, Rio Linda, Sacramento Co., 19 May 1958 (Light trap, Jack Fowler) [UCD] ; ts; Donner Pass, 1 Aug: 1948" (HS M. G. & D> Townes) [HRT]> 1 3, Cisco, 31 July 1948 (Hi, MG. & D: Townes) [HKT]; ARIZONA: 1 ¢, North Rim, Grand Canyon, 29 July 1954 (H. E. Evans) [MCZ]; COLORADO: 1 ¢? , Waldo Canyon, 30 June 1916 (W. D. Edmonston) [USNM] ; SASKATCHEWAN: 1 ¢, Saskatoon, 30 June 1950 (A. R. Brooks) [CNC] ; ALBERTA: 1 2,30 ¢ 6, Onefour, 3 June 1956 (O. Peck) [CNC] ; YUKON: 1 ¢, Whitehorse, 4 July 1948 (Mason and Hughes) [CNC]. Variation The four female paratypes are very similar to the allotype in color and in all important structural details; the length of the fore wing varies from 3 to 3.6 mm. The males exhibit much size variation, the smallest (Donner Pass, Calif.) having the fore wing only 2.2 mm. long, the largest (Rio Linda, Calif.) having the fore wing 4.5 mm. long. There is little color variation except that the front tibiae are dull brown in a few specimens, while in others the middle tibiae are light yellowish- brown like the front tibiae. In most specimens the postocellar and ocello-ocular lines are subequal. Remarks. — This species is very similar to canadensis and may represent no more than a western race of that species. However, there seems to be some overlap of the ranges in Arizona and in western Canada. 1961 NEARCTIC SPECIES OF SIEROLOMORPHA 11 SIEROLOMORPHA CANADENSIS (Provancher ) Photopsis canadensis Provancher, 1888, Add. Corr. Faune Ent. Canada, Hymen., p. 410 [Type: 6, Ottawa, Canada (Har- rington) |. Sierola (2?) ambigua Ashmead, 1893, Bull. U. 8. Nat. Mus., 45: 56 [Type: ¢, Brookings, So. Dakota (Coll. Ashmead) (USNM, no. 56018) |. Mutilla tertia Dalla Torre, 1897, Cat. Hymen., 8: 91 (new name for canadensis, preoccupied in Mutilla). Sierolomorpha ambigua Ashmead, 1905, Canad. Ent., 35: 42. Sicrolomorpha canadensis Krombein, 1951, U. S. Dept. Agri., Agri. Monoer., 2: 749. Remarks. — Provancher’s canadensis was transferred to this genus by Krombein on the advice of R. M. Schuster, who stated that a specimen in the University of Minnesota collection which had been compared with Provancher’s type was found to be the same as ambigua Ashmead. I have not seen Provancher’s type, but his description does indeed seem to fit this species rather well. The following statement in particular makes it clear that his name applies to a species (such as this one) with a rather marked constriction between the first two abdominal tergites: ‘‘la suture entre les segments 1 et 2 enfoncée et crénelée.’’? Un- fortunately, the abdomen of the type of ambigua is missing, but my interpretation of Ashmead’s species is based upon a topo- type which agrees closely in all details regarding the head and thorax, including the characteristic sculpture of the median line of the propodeum. It seems to me best to consider canadensis and ambigua synonyms, with the former having priority over the more widely used name ambigua. I admit the possibility that I may be confusing more than one species here. However, speci- mens from the northern tier of states and from Canada show no undue amount of variation, and I consider it very doubtful that Ashmead and Provancher had different species before them. Distribution. — This species occurs widely over eastern North America, ranging from Florida, Texas, and Arizona to Alberta, Ontario, and Massachusetts. I have seen specimens from the following localities: MASSACHUSETTS: Waltham, June; CON- NECTICUT: Lyme, June; NEW YORK: Oneonta, Ithaca, River- head, L. I., Northwest, L. I., June-Aug.; PENNSYLVANIA: Hummelstown, May; MARYLAND: Bowie, Takoma Park, May- July; VIRGINIA: Rosslyn, Great Falls, Fredricksbure, May- June; NORTH CAROLINA: Highlands, Wallace, Pink Beds, 12 BREVIORA No. 140 June-July; GEORGIA: Tifton, Valdosta, May; FLORIDA: Ormond, Osceola Co., Volusia Co., Brevard Co., Dee.-Jan. ; OHIO: Columbus, June; MICHIGAN: Ann Arbor, Erie, Liv- ingston Co., June-Aug.; ILLINOIS: Champaign; IOWA: Sioux City, Ames, June; ALBERTA: (no further data) ; NORTH DA- KOTA: Bottineau, Aug.; SOUTH DAKOTA: Brookings, June- July; KANSAS: Lawrence, Manhattan, Pottawatomie Co., Geary Co., May-July; LOUISIANA: Tallulah; TEXAS: Sealy, Victoria, May, Nov.; NEW MEXICO: Mesilla Park, Sept.; ARIZONA: Springerville, July. ; Figs. 1-6. Middle antennal segments of males of six species of Sierolo- morpha, In each case the first segment shown (at the bottom) is segment seven. These figures are somewhat diagrammatic, as the tyloides normally form a somewhat twisted series, here shown as a straight series; further- more, the tyloides are normally somewhat obscured by setulae, here omitted. Fig. 1 — 8. nigrescens n. sp., segments 7-10; Fig. 2 — S. canadensis (Prov.), segments 7-10; Fig. 3—S. brevicornis n. sp., segments 7-13; Fig. 4—S. similis n. sp., segments 7-10; Fig. 5—S. bicolor (?) n. sp., segments 7-10; Fig. 6 — S. apache n. sp., segments 7-10 and 13. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. JUNE 28, 1961 NUMBER 141 THREE NEW TOADS FROM SOUTH AMERICA: BUFO MANICORENSIS, BUFO SPINULOSUS ALTIPERUVIANUS AND BUFO QUECHUA By Jos& M. GALLARDO Museo Argentino de Ciencias Naturales Buenos Aires INTRODUCTION In the course of a general study of the Neotropical Bufonidae, I received some material from the British Museum (Natural History) (BMNH), the American Museum of Natural History (AMNH), the Carnegie Museum (CM) and the Museum of Zoology at the University of Michigan (MZUM) which in- eluded the new forms described below. One of these is a quite different form of Bufo from the State of Amazonas, Brasil; another is a subspecies of Bufo spinulosus from the Department of Oruro, Bolivia, and the last a new species of the Bufo ockendent group from the Department of Cochabamba, Bolivia. BUFO MANICORENSIS Sp. Nov. Type. BMNH 1898, 3.10.1, adult male, Manicoré, Rio Madeira, State of Amazonas, Brasil. Description, Head elongate and sharp. Rostrum nearly ver- tical. Nostrils on a prominence, elongate and oblique. Cephalic crests well marked, with smooth or somewhat rippled borders. Subnasal crests visible. Canthal crests nearly convergent. Mavxil- lary crests somewhat expanded. Preorbital crests slightly slop- ing posterolaterally. Postorbital crests sloping anterolaterally, close to the anterior border of the tympanum. Suborbital crests not expanded, rather distant from the lower border of the eye. 2 BREVIORA No. 141 Supraorbital crests somewhat raised; interorbital space very narrow and concave, with granules in the parietal region. Parie- tal crests elongate and oblique, forming an angle with the supra- orbital crests. Orbitotvmpanie crests short but distinct. Greatest diameter of eye, 6 mm. Vertical diameter of tympanum, 3 mm. Tympanum on an outward sloping plane, Paratoids subtriangu- lar with indistinet borders, with their long axis oblique medio- laterally, with flat dorsal granules. One subgular vocal sae. Dorsum with flat granules. Belly with the larger granules on abdominal region. Limbs with dorsal conical granules. Without interdigital membrane in the hand; Ist and 2nd fingers sub- equal; subarticular tubercles double on fingers 2 and 3; two carpal tubercles, the inner one smaller, elliptic and somewhat salient, the outer larger and rounded. Subarticular tubercle double on toe 4; interdigital membrane in the toes to near the tip of the digits, but in toe 4+ basal and prolonged as a serrated cutaneous fringe; two metatarsal tubercles small but elongate, the inner more salient; tarsal fringe absent. Dorsal coloration hight brown, with some darker spots, not well marked. Belly light. Dimensions. Head and body 50 mm. Head length 12.5 mm. Head width 17 mm. Head height 5.5 mm. Interorbital space 2.5 mm. Elbow to the third finger 21 mm. Femur 18 mm. Tibia 16 mm. Heel to the fourth tee 25.5 mm. Foot 17 mm. Diagnostic features. Bufo manicorcns’s differs from all other Neotropical toads; the shape of the head and cephalic crests recall certain Asiatic forms. In the Neotropical area the nearest species is Bufo intermedius Giinther, but this is distinguished by the well marked parietals and preorbitals, the interorbital space narrower than the upper eyelid, the distinct tympanum, and by the paratoids which are not elliptical and are separated from the eye by the orbitotympanic crests. The new species 1s not as close to Bufo valliceps Wiegmann and differs from that species in having less prominent cephalic crests, the interorbital space narrower, double subarticular tubercles on the foot and no lateral granules in a row continuing the paratoids posteriorly. Material studied. BMNH 1898, 3.10.1 (1 specimen), Mani- coré, Rio Madeira, Brasil, B. Piffard. BUFO SPINULOSUS ALTIPERUVIANUS subsp. nov. Type. AMNH 14418, adult female, Challapata, Department of Oruro, Bolivia. Description. Head very short and wide. Loreal region sloping 1961 NEW TOADS FROM SOUTH AMERICA 3 outward. Rostrum vertical. No cephalic crests, excepting maxil- laries; canthus rostralis thick. Interorbital space granular. Tym- panum sloping out. Paratoids well marked and rounded, con- tinued laterally by large granules (each granule with many horny points). Two types of dorsal granules: the larger with one central horny point and many others around it, the smaller with only one horny point. Larger granules on abdominal region. First finger longer than second; subarticular tubercles on the fingers, double or semidivided; palmar outer tubercle larger and rounded, inner smaller and elongate. Interdigital membrane of the foot basal but prolonged as a fringe on the toes: subarticular tubercles on the toes, simple or sometimes double; two metatarsal tubercles, the inner more salient; a thick tarsal fringe. Dimensions. Head and body 80 mm. Head length 17 mm. Head width 30 mm. Head height 11 mm. Interorbital space 6 mm. Upper evelid width 6 mm. Eve 7 mm. Tympanum 3.5 mm. Paratoid 9 mm. by 8.5 mm. Elbow to the third finger 38 mm. Femur 36 mm. Tibia 30 mm. Heel to the fourth toe 50 mm. Foot 35 mm. Paratype. AMNH 14417, Choro, Bolivia, adult female 82 mm. Distribution. The two localities of the material studied, Chal- lapata and Choro, are in the Department of Oruro, Bolivia; the type locality is at 3700 metres altitude. Diagnostic features. According to Vellard (1959), there are six subspecies of Bufo spinulosus: B. s. spinulosus, B, s. are- quipensis, B. s. limensis, B. s. trifolium, B. s. flavopictus and B. s. orientalis. B. s. altiperuvianus adds a seventh. Bufo s. altiperuvianus differs from B. s. spinulosus, the Bol- ivian subspecies structurally and geographically closest (De- partment of La Paz) in having the head shorter, not so distinet from the body; the loreal region sloping more laterally; tym- panum larger; paratoids larger and more rounded. Capurro (1950: 11) has cited B. spinulosus from Tarapaca Provinee, Chile (west of Oruro Department, on the other side of the Cordillera Occidental), but specimens of this provenance that I examined at the Chicago Natural History Museum are dif- ferent from the form here described. Remarks. 1 name this subspecies after the old Spanish name, Alto Pert, of the region from which it derives. Material studied. AMNH 14418 (1 specimen) Challapata, Bo- livia. AMNH 14417 (1 specimen) Choro, Bolivia. 4 BREVIORA No. 141 BUFO QUECHUA sp. nov. Type. CM 4225, adult female, Incachaea, 2500 m., Department of Cochabamba, Bolivia. Description. ead triangular, widest at the angle of the mouth ; loreal region sloping outward. Maxillary border marked ; eanthus rostralis thick; supraorbital crest absent; parietal crest visible; orbitotympanie crest thick; one rostral-internasal crest more or less marked. Tympanum not visible. Paratoids approximately elliptic, dorsally smooth. Body dorsally with sparse large granules, but with abundant small granules. One lateral row of granules, continuing the paratoids, each granule with a large central papilla and smaller papillae around it; below the row, lateral granules of the same type. Belly with abundant conical and simple granules. Elongate limbs, with conical granules dorsally. First finger longer than second; fingers free, borders with small conical granules; subarticular tubercles generally simple, but double on the third finger; outer palmar tubercle large and rounded, inner one smaller and elongate. Tarsal fringe absent; two metatarsal tubercles elon- gate and approximately of the same size, the outer one more salient; interdigital membrane near the toe tips, but on the fourth only a little more than half its length and prolonged as a cutaneous fringe; subarticular tubercles on toes, small and simple. Dorsum heht brown with three large darker triangular spots not well marked ; one interocular with base to the front and two others on the body with the base to the rear; a vertebral heht line divided the last two triangles. Limbs dorsally with transverse wide dark bands. Belly yellowish with dark spots shaped very irregularly. Dimensions. Head and body 50 mm. Head length 12 mm. Head width 17 mm. Head height 7 mm. Eye 5 mm. Upper eyelid width 4.5 mm. Interorbital space 5 mm. Paratoid length 7.5 mm. Elbow to the third finger 22 mm. Femur 20 mm. Tibia 17 mm. Heel to the fourth toe 28 mm. Foot 20 mm. Paratypes. CM 4223, 4224, 4226, Incachaca, Dept. Cocha- bamba, Bolivia. Head and body: 62 mm., 41.5 mm., 37 mm., respectively. Diagnostic features. Four other species related to Bufo que- chua have been previously described: B. ockendent Boulenger, B. inca Stejneger, B. leptoscelis Boulenger and B, fissipes Bou- lenger. In the table below their differential characters and the altitude at which they are found are shown. AMERICA SOUTH TOADS FROM NEW 1961 so.joul 0096-0066 [Boruog sqyurod a ‘ 6) 7 > 1 —_— Ea BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. Aueust 21, 1961 NuMBER 144 A LARGE OPHIACODONT PELYCOSAUR FROM THE PENNSYLVANIAN OF THE PITTSBURGH REGION By ALFRED SHERWOOD ROMER The presence in the early Permian of a varied series of pely- cosaurs indicates that the beginnings of this synapsid group occurred well back in Carboniferous times. Pennsylvanian re- mains of pelycosaurs are, however, very rare. Because of this paucity of data, the remains described below are worthy of record despite their incomplete and fragmentary nature. Along the course of McKnight Road, about 6 miles north of Pittsburgh, Pennsylvania, there has recently been considerable quarrying of sand and shale deposits of Conemaugh age in order to level the surface for building construction. The materials quarried were used as fill in North Park, several miles farther north. During the course of this work geology students at the University of Pittsburgh noticed three pieces of bone in freshly dumped park fill; through the courtesy of Mr. Martin Bender of the University’s geology department they were deposited in the Carnegie Museum. Inquiry made it rather certain that this material had been excavated from a locality on the east side of McKnight Road near the junction of Brown’s Lane. The sand- stone excavated here, les just above the Ames limestone, and hence pertains to the upper part of the Conemaugh group of the Pennsylvanian. It seems certain that all three of the fragments, pertaining to a right shoulder girdle and front leg, were parts of a single, presumably articulated, specimen, but the state of the quarrying operation precluded any attempt to find further remains. The specimens are entered as no. 13942 in the vertebrate paleontology collections of the Carnegie Museum, Pittsburgh; I am indebted to Curator Craig Black for placing them in my hands for description. Needed preparation was done at the Museum of Comparative Zoology under a grant from the National Science Foundation. ~ 2 BREVIORA No. 144 The most revealing specimen consists of the distal half of a right humerus. It is clearly pelycosaurian in nature and, it would seem, equally certainly ophiacodontid, the diagnostie features being the greatly expanded entepicondyle, the transversely di- rected supinator process and the strong development of the lateral marein of the ectepicondyle. In addition, the preserved portion of the shaft indicates that proximal and distal halves of the bone were sharply twisted on one another as in ophiacodonts and in contrast to the lesser torsion in other pelycosaur groups. The overall breadth is 96 mm. The bone is almost exactly super- posable on a mature Ophiacodon humerus in the M.C.Z. collee- tions (no. 1426) from the Archer City bonebed (Putnam formation). This Texas element is 120 mm. in length, and pertains to a small individual of O. retroversus or to an immediate pre- decessor of that species. If the proportions of the present form were similar to those of Ophiacodon, the animal in life would have had a length of head and trunk combined of about 109 em., and a total length (including the usual long pelyeosaurian tail) of about 204 em. or about 678”. Our Pennsylvanian specimen thus appears to be one of the largest of ophiacodonts, exceeded in size only by O. retroversus and O. major of the later Wichita formations and by Stereorhachis dominans of the European Permian. Post-mortem, the bone has lost the hemispherical ventral swelling with which the radius articulated; the supinator process is imperfectly preserved and there has been some slight damage to the ventral surface at the end of the entepicondyle. Otherwise, apart from crushing, the bone is well preserved. Notable and possibly significant is the fact that ossification had been completed at the time of death. This is in contrast to most specimens of Ophiacodon, in which but a small fraction of limb bones show complete ossification of their ends, even when the animal appears to have reached a ‘“mature’’ size. Although the general pattern of the bone is comparable to that of Ophiacodon, there are definite differences in detail. The present specimen has, dorsally, a prominent muscle scar proximo- medially situated near the end of the entepicondyle; this is not present in Ophiacodon. The ridge extending proximally along the lateral margin of the ectepicondyle is better developed in the present form than in Ophiacodon. The supinator process appears to be less developed than in that genus, but the apparent dif- ference is due to post-mortem damage in our specimen. 1961 LARGE PENNSYLVANIAN PELYCOSAUR 3 A second fragment preserved is the proximal end of the right ulna. This has been greatly crushed, so that the sigmoid articular surface for the humerus is unnaturally narrow, and identification of diagnostic features is difficult. As in the case of the humerus, ossification is practically complete so that, in contrast to most Ophiacodon specimens, the whole olecranon is represented by bone rather than cartilage. A third fragment is the upper end of a right scapula. The posterior margin is thickened and rounded in typical pelycosaur fashion. The lower posterior portion of the piece preserved is somewhat crushed and distorted, but there is an increase in thickness and a curvature of this border such as one would expect as the supraglenoid region is approached. In many pely- cosaur specimens, even when seemingly ‘‘mature,’’ the upper end of the scapular blade has a thickened distal edge obviously con- tinued in cartilage. The present specimen, which appears to be nearly intact in the posterior part of its upper end, shows no such terminal surface; ossification was apparently complete. More anteriorly the bone has been damaged so that for about 6 em. there is a thick broken margin on the preserved portion. Beyond and below this the margin as preserved, although im- perfect, is thinner where broken off. Probably the missing area here had about the extent indicated by the broken line in the figure. The anterior margin flares out laterally to a marked degree from the general plane of the scapular blade. It seems fairly certain that there was considerable constriction in the width of the blade toward the bottom of the preserved segment. This is in contrast with Ophiacodon and Varanosaurus in which the scapular blade is broad throughout its height (the scapula of Clepsydrops is incompletely preserved). Fragmentary as are the remains here described, they show definitely the presence at this horizon, fairly well down in the Upper Carboniferous, of a large pelycosaur which is surely an ophiacodontoid and nearly equally surely an ophiacodontid, ante- cedent although probably not ancestral to Ophiacodon of the Permian. Despite its incomplete nature, this fossil deserves taxonomic standing because of its importance in the story of pelycosaur evolution, and it is herewith made the holotype of Clepysdrops? magnus, sp. nov. It will not probably prove, when better known, to be generically distinet from Clepsydrops, but it appears to be related to that genus, likewise of late Carboniferous age, and it may be provisionally included in it. Because of imperfect knowledge of comparable anatomical features, we may 4 BREVIORA No. 144 for the present rely for diagnosis simply on the size, which is approximately half again that of specimens assigned to the genotype, C. collettu. Advantage may be taken of the present opportunity to mention a few fragmentary remains of Pennsylvanian pelycosaurs found in Ohio by Dr. Donald Baird and Mrs. Baird during a 1955 expedition of the Museum of Comparative Zoology. (1) M.C.Z. no. 2411 is from the Summerfield limestone of the Conemaugh group; the locality lies in Center Township, Noble County, Ohio, on the south side of state highway 78, just west of its junction with route 147. Found here were a crushed claw-shaped ungual phalanx 24 mm. long, comparable to the unguals of Clepsydrops, two complete pelycosaur centra with transverse diameters of 20 and 14 mm., and a fragmentary centrum with an estimated diameter of about 15 mm. (2) M.C.Z. no. 2295 from the Ewing limestone of the Conemaugh, in the central part of section 7, Noble Township, Noble County, a fragment of a centrum with an estimated diameter of 20 mm. or so. (3) M.C.Z. no. 2777, from the Lower Uniontown limestone of the Monongahela group, along Leith Run Road 3.8 miles from the junction with state route 7, Washington County, Ohio, an ungual phalanx compar- able to that mentioned under no. 2411. It may be noted that this is the only identified reptilian bone from the Monongahela. There is, of course, little in these fragments to allow us to give any positive generic determination. But since these specimens are within the size range of the Illinois materials of Clepsydrops they may be provisionally referred to that genus. As mentioned earlier, described pelycosaurian remains from the Carboniferous are very rare indeed. Their rarity is pre- sumably to be correlated with the fact that most Carboniferous fossil localities are from coal swamps in which, apart from am- phibious ophiacodonts, pelycosaurs would not be expected. Prev- iously described are: (1) Numerous isolated elements of a small ophiacodontid, Clepsydrops (Romer and Price, 1940, pp. 212- 216) from the McLeansboro formation near Danville, Illinois. (2) A fragmentary spine of a small Hdaphosaurus from the Round Knob formation of the Conemaugh group near Pitcairn, Pennsylvania (Case, 1908, pp. 237-238; Romer and Price, 1940, p. 388). (3) From the late Stephanian of Kounova, Bohemia, a single vertebra of a similarly small Edaphosaurus (Romer and Price, 1940, p. 388). (4) From the same locality, a number of bones which represent a sphenacodont of good size, to which the name Macromerion schwarzenbergu is applicable (Romer, 1945, pp. 429-431). 1961 LARGE PENNSYLVANIAN PELYCOSAUR 5 Little as we know of Pennsylvanian pelycosaurs, a few general conclusions can be reached. That our scanty data on Carboni- ferous pelycosaurs should include remains of ophiacodonts such as Clepsydrops is not surprising. It is believed that the ophiaco- dontoids are the most primitive of pelycosaur stocks and would hence be expected to appear at an early period in the record. Further, it is believed that many, at least, of the ophiacodontids were amphibious in habit — essentially piscivorous water-dwellers whose ancestors had never abandoned an aquatic existence. In consequence the chances of finding ophiacodonts in the coal swamp deposits which constitute most of the Carboniferous record is much greater than of finding representatives of the more terrestrial sphenacodontoids or edaphosauroids. Any idea that the finding of Clepsydrops in the Conemaugh takes us near in time to the point of origin of the pelycosaurs is an illusion. For fragments of edaphosaur spine have been recovered not only from the very late Kounova deposit but also from the Round Knob formation of the Conemaugh. Hdapho- saurus is a highly specialized end form. It is obvious that its evolution from an ancestral pelycosaur must have begun at a time much further back in the Carboniferous than the Cone- maugh. The assumption (not an unreasonable one) that the major pelycosaur groups evolved independently from a capto- rhinomorph stock only begs the question. In general, reptilian stocks of any sort begin with forms that are of small size (as well as of primitive nature) and become increasingly larger as their history progresses. Such a process seems clear in the Lower Permian history of the pelycosaurs. The earliest dimetrodonts of the Texas Wichita were relatively small, and giants developed in the Clear Fork; Hdaphosaurus species increase greatly in size in later beds; the caseids grew from small forms to enormous Cotylorhynchus species; even Ophiacodon (which became extinct before the typical Clear Fork) developed from small forms in the lower Wichita and New Mexican beds to the relatively gigantic O. major, in the Clyde Formation. Until the discovery of the present specimen of C. magnus the known Pennsylvanian pelycosaur material fell in line with the belief that all early pelycosaurs were small. The Clepsydrops specimens from Illinois and Ohio represent small animals, as do both Pennsylvanian scraps of Hdaphosaurus. To be sure, Jacro- merion of Kounova is a good-sized sphenacodont, but Kounova is close to the Permian boundary and Macromerion could be regarded as a slightly premature representative of the Permian 6 BREVIORA No. 144 trend to large size. The discovery of C. magnus, however, destroys the illusion, for, as noted, this animal was larger than most of his later Permian relatives. When and if we ever come upon Upper Carboniferous deposits with a representative terrestrial, rather than coal-swamp fauna, it can be confidently predicted that there will be included a considerable number of pelycosaurs which will vary widely in structure and size. It seems certain that for the origin of this fauna, of which so far we have but slight traces, we must look far back in the Carboniferous, to at least early Pottsville or Namurian levels. LITERATURE CITED Cass, E. C. 1908. Description of vertebrate fossils from the vicinity of Pittsburgh, Pennsylvania. Ann. Carnegie Mus., 4:234-241. Romer, A. 8. 1945. The late Carboniferous vertebrate fauna of Kounova (Bohemia) compared with that of the Texas redbeds. Amer. Jour. Sci., 243:417-442. Romer, A. 8S. and L. I. PRICE 1940. Review of the Pelycosauria. Spee. Pap. Geol. Soc. Amer., no. 28: 1-538. 1961 LARGE PENNSYLVANIAN PELYCOSAUR ‘i Figure 1. Clepsydrops magnus sp. noy. A, B, distal end of right humerus, dorsal and ventral surface. C, upper part of right scapular blade; probable outline when complete suggested by broken line. D, EF, lateral and proximal views of upper end of left ulna. All x ¥%. BREVIORA Museum of Comparative Zoology CAMBRIDGE, MASs. SEPTEMBER 5, 1961 NuMBER 145 A NEW SPECIES OF THE CETOMIMID GENUS GYRINOMIMUS FROM THE GULF OF MEXICO By Henry B. Bigelow GYRINOMIMUS PARRI new species Type. U.S. National Museum No. 196180, from north central part of Gulf of Mexico, 26° 52’ N, 89° 44” W, OREGON Station 2573. One specimen, 51 mm in standard length. Distinctive Characters. G. parri differs conspicuously from G. simplex Parr 1946,! from the Gulf of Mexico, in its relatively much longer dorsal and anal fins, from G. myersi Parr 1934,? also from the Gulf, in the simple structure of its lateral line pores, and from G. bruni, Rofen 1957-59,* from the southwestern Indian Ocean, in the number of lateral line pores. Description. Proportional dimensions in per cent of standard length of the type specimen, 51 mm in standard length. Snout in front of eye: 51. Diameter of eye: about 1. Length of head to pectoral fins: 33. Snout to origin of dorsal: 62. Base of dorsal: 24. Base of anal: 23. Rear end of base of dorsal to base of caudal: 10. Length of caudal: 12. Length of pectorals: 6. Height at pectorals: 24. Height at origin of dorsal: 17. Least height of caudal peduncle: 8. Height of base of caudal fin: 12 1 Copeia, 1946: 116, Pl. 1. 2 Bull. Bingham Oceanogr. Coll., # (6) : 29, Figs. 8, 9, 1934. 3 Galathea Rept., 1 : 257, 1957-d9. bo BREVIORA No. 145 Greatest width between rear end of rami of lower jaw: 18. Width between upper ends of gill slits: 14. Dorsal rays: 17. Anal rays: 17. Principal caudal rays: 17+ (damaged). Pectoral rays: 16. Dorsal profile sloping in nearly a straight line from nape to tip of snout, the latter narrowly rounded vertically but rather broadly rounded transversely ; eve minute, much nearer to upper jaw than to dorsal profile; upper jaw projecting a little beyond lower, margins of both jaws weakly concave, the jaws extending rearward about to a perpendicular at upper corner of gill open- ings. Trunk highest at nape; dorsal profile of body and tail sectors sloping in nearly a straight line from nape to caudal pedunele. Dorsal fin a little higher anteriorly than posteriorly, its margin very weakly convex; anal opposite dorsal and similar to dorsal in outline. Caudal (somewhat damaged) apparently truncate or nearly so; lateral line with 14 large oval pores from nape to base of caudal, the anterior margin of the pores without rearward extending flap. Teeth along entire leneth of each jaw in 3 continuous rows, those in the row next the gum much the shortest and those in the row farthest from the gum much the longest. Color. General ground tint sooty black, the eyes blue, the outer parts of the unpaired fins chocolate brown, the teeth ivory white, the inside of the mouth pale gray. Size. The type (and only known) specimen is 51 mm in stand- ype ( A I ard leneth. Range. So far known only from the north central part of the Gulf of Mexico, 26°52’ N, 89°44” W, OREGON St. 2573, from a trawl haul at 1350 fms. Gyrinomimus parri n. sp. Type specimen, USNM no. 196180. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 14, 1961 NUMBER 146 NEW RODENTS FROM THE EARLY MIOCENE DEPOSITS OF SIXTY-SLX MOUNTAIN, WYOMING By Craia C. BLACK Carnegie Museum, Pittsburgh, Pennsylvania During the summers of 1930, 1931, and 1933, Dr. E. M. Sehlaikjer working for the Museum of Comparative Zoology, Harvard University, made a large collection of early Miocene mammals in the southern portion of Goshen Hole, Wyoming. Most of these were obtained from the slopes of Sixty-Six and Sear Creek mountains, from beds that Schlaikjer (1935) in- terpreted as the Lower Harrison formation. On two occasions during July, 1959, Sabra B. Black, Laura MeGrew, Bryan Patterson, and I made a brief reconnaissance of the area on the western slope of Sixty-Six mountain. During this reconnaissance, it became evident that Schlaikjer’s Brule-Lower Harrison contact was actually a local channel conglomerate de- veloped within the early Miocene sediments. He states (1935, p. 112), ‘‘At the northwest end of Sixty-Six mountain in the N.E. 1/4, See. 7, T.20N., R.GOW. the typical Brule clay is overlain by a three-foot clay conglomerate above which are nineteen feet of clay sands that grade upward into the gray sands with pipy eoneretions.’’ The ‘‘typical Brule clay’’ below the conglomerate, however, is the unit from which the rodents described below were obtained together with a badly weathered Cyclopidius skull and jaws and the posterior portions of two rami of Wesoreodon cheek. These specimens were found in place and they clearly indicate an early Miocene, probably Gehring equivalent, age. Below the grey, tuffaceous sand and grading into it are buff clays with some sand which are probably equivalent to the Brule. In this area deposition appears to have been continuous from the Oligocene through the early Miocene with no sharp lithologic or erosional break within this sequence. The clay conglomerate is not ex- tensively developed either on Sixty-Six or Bear mountains and 2 BREVIORA No. 146 where it does appear on Sixty-Six mountain it les within early Miocene sediments as is clearly shown by the oreodonts and ro- dents described below. A full account of the relationship of the Oligocene and Miocene sediments in the southern part of Goshen Hole is still in progress and will be published at a later date. [ would like to take this opportunity to thank the Kellam family of Torrington, Wyoming, who helped us immeasurably during our stay in the Goshen Hole area. I would also like to thank Professor Bryan Patterson for his criticisms and sugges- tions. The drawings are by Mr. Clifford J. Morrow and were made possible by a grant from the Gulf Oil Corporation. Abbreviation used: M.C.Z.— Museum of Comparative Zool- ogy, Harvard University. Family CRICETIDAE ScCOTTIMUS KELLAMORUM' Nn. sp. Figure | Type: M.C.Z. No. 7342, a right maxillary with M!-M2. Hypodigm: Type only. Horizon and locality: Section 11, T.20N., R.61W., Goshen Co., Wyoming. Arikareean, from the supposed Gehring equivalent, early Miocene. Diagnosis: Smaller than Scottinus lophatus; teeth narrower in relation to length than in NScottimus eriguus; antero-posterior lophs more prominent than transverse; protoloph very weak on M!, absent on M?; metaloph very weak on M?, incomplete on Mt. Description: In general, the upper molars of Scottimus hel- lamorum, like those of S. lophatus, are longer and narrower than those of the various species of Eumys. The mure is as strongly developed as in SN. lophatus, but there are more accessory trans- verse crests than in that species. The cusps on M1!-M? are high, and the crests are at a lower level. The anterocone of M! is ex- tremely large. There is a connection between the paracone and ‘protocone on M}, but it is shifted posteriorly and thus does not form a transverse loph. A comparable crest is not present on M2. The paracone and metacone are joined by a low crest which in turn is joined by the mesolophid. There is a short crest pro- jecting lingually from the mure between the protocone and hypo- cone on both M! and M2. The anterior cingulum on M-? is ele- 1 Named for David and Jean Kellam of Torrington, Wyoming. 1961 NEW MIOCENE RODENTS 5) vated and strongly developed. The posterior metacone arm on M! is directed backward to fuse with the elevated posterior cingulum, whereas on M? it passes lingually to the hypocone to form a weak, transverse metaloph. Figure 1. Scottimus kellamorum n. sp., Type, M.C.Z. No. 7342, RM1-, anterior end to the right, X20. Discussion: Two species of Oligocene eumyines described by Wood (1937) seem referable to Scottimus. These are Lumys e.xr- iguus, from the Middle Oreodon beds of South Dakota, and Leidymys vetus, from the Middle Oligocene Cedar Creek beds of northeastern Colorado. Wood (op. cit., p. 254-255), in describing Eumys exiguus, stated that it was transitional between typical ELumys and Scottimus. Galbreath (1953, p. 72) stated that all the specimens referable to Humys exiguus in the University of Kansas collections from the Cedar Creek beds were also ‘* closely similar to the type of Leidymys vetus Wood. In fact there is no question in my mind but that they represent the same species.”’ However, he did not synonymize the two, stating, ‘‘I think this small species (Humys eriguus) is generically distinct from the species of Humys but am not prepared to say whether or not it should be referred to Leidymys or to another genus.’” Under the circumstances, it seems proper to place Humys exiguus in Scottimus and to refer Leidymys vetus to the synonymy of this species. This brings the number of recognized species of Scottimus to three; S. lophatus, S. eriguus, and S. kellamorum. 4 BREVIORA No. 146 In the development of antero-posterior rather than transverse lophs Scottimus kellamorum is more advanced than S. exiguus. Sinee NS. eriguus is of middle Oligocene age this is, of course, to be expected. S. kellamorum, however, is not as advanced a species as the earher SN. lophatus. The transverse crests seen on M! of S. hellamorum are also present on S. lophatus, but these crests are not seen on M? of S. lophatus. This would seem to exclude SN. lophatus from the ancestry of S. kellamorum; presumably a species similar to S. eriguus was ancestral to both. EuMYS sp. M.C.Z. No. 7334, a partial left ramus with the incisor and M,-Ms cannot be indentified specifically. It does not show any tendeney toward the formation of a strong central antero- posterior lophid at the expense of the transverse lophids, and henee is certainly not the lower dentition of Scottimus hella- morum. The posterior protoconid arm is extremely elongate on both M, and Ms. On M,, a short anterior metaconid arm just fails to reach the anteroconid whereas a longer protoconid arm does so. In size and general pattern the teeth closely resemble those of Humys elegans and Eumys obliquidens. Unfortunately the lower teeth of Leidymys nematodon and lockingtonianus are unknown. Family HETEROMYIDAE HELISCOMYS SCHLAIKJERI’ 1. sp. Figure 2 Type: M.C.Z. No. 7333, a right maxillary with P4-M?. Hypodigm: Type only. Horizon and locality: Section 11, T.20N., R.61W., Goshen Co., Wyoming. Arikareean, from the supposed Gehring equivalent, early Miocene. Diagnosis: Size larger than that of any other known species ot Heliscomys: internal cingulum undivided; posterior cingulum on P* extending from hypocone to metacone ; central transverse valley straight, directed somewhat posteriorly. Description: No trace of the paracone remains on the ante- roloph of the premolar. In this respect, the tooth is typically perognathine. The posteroloph bears three cusps, the metacone Ov 1961 NEW MIOCENE RODENTS and hypocone being of equal size with the protocone and the entostyle being smaller. The entostyle curves forward and joims the internal edge of the protocone at the same level as does the hypocone. There is a small posterior cingulum running from the hypocone to the base of the metacone, and separated from the posteroloph by a small pit. Figure 2. Heliscomys schlaikjeri n. sp., Type, M.C.Z. No. 7335, RP4-M?, anterior end to the left, X30. The molars are somewhat more lophate in appearance than are those of Heliscomys gregoryt (Wood, 1933) and H. hatcheri (Wood, 1935), although the principal cusps are still prominent. In this respect the teeth resemble more closely those of H. tenwiceps (Galbreath, 1948). The molars of both agree in having anterior and posterior cingula that rise rather steeply to joi the protostyle and entostyle. These cusps are closely appressed with no gap between them, a further point of resemblance to //, tenuiceps. The median valley is straight on both teeth, slanting somewhat posteriorly, not sinuous as in //. gregory: and H. hatchern. The anterior cingulum is stronger than the posterior on both M?! and M-. Discussion: Although it is somewhat larger and shows a few minor differences, Ieliscomys schlaikjeri is extremely close to IT. tenwiceps. Both species have an undivded internal cingulum 2 Named for IE. M. Schlaikjer in recognition of his extensive work in the Goshen Hole area, 6 BREVIORA No. 146 and a straight median valley, characters found neither in /H. hatcheri nor in IT. gregoryt. In H. schlaikjeri, in addition to the somewhat larger size, the median valley of the molars is directed somewhat more posteriorly and the posterior cingulum on P#4 has shifted position. The latter distinction seems unimportant since the presence or absence and the position of the posterior cmeulum are variable in fHeliscomys. In H. hatcheri this ein- gulum varies from absent to present, in some cases extending alone the entire posterior margin of the molars; in a specimen from Pipestone Springs described by MeGrew (1941), it appears to be completely absent, while in Hf. fenwiceps it connects the hypocone and the entostyle. Recently, Reeder (1960) has de- seribed two new genera of heteromyids from the White River formation and has emphasized their laree size and the quadri- cuspidate nature of Py in Apletotomeus and the quinquicuspidate P, of Akmaiomys. These genera are known only on their lower dentitions and hence can not be compared with HT. schlaikjert. H. schlaik jeri, however, is somewhat larger than either genus. Table of Measurements (in mm. ) pt M1 M? My Mo Scottimus kellamorwumn anteroposterior 2.50 1.85 transverse protoloph 1.60 1.43 transverse metaloph 1.53 1235 Kumys sp. anteroposterior |. 2.10 2.00 transverse metalophid ; 1.40 1.90 transverse hypolophid ; 1.70 1.90 Heliscomys schlaikjeri anteroposterior. se cunorrson MOY) 1.10 1.10 transverse -em He20 1.40 1.40 REFERENCES GALBREATH, E. C. 1948, A new species of heteromyid rodent from the Middle Oligocene of northeast Colorado with remarks on the skull. U. Kansas Publ., Mus. Nat. Hist., 1: 285-300, 2 pls. 1953. A contribution to the Tertiary geology and paleontology of northeastern Colorado. U. Kansas Publ. Paleon. Contrib. Verte- brata, 4: 1-120, 2 pls. ~] 1961 NEW MIOCENE RODENTS McGrew, P. O. 1941. Heteromyids from the Miocene and Lower Oligocene. Geol. Ser. Field Mus. Nat. Hist., 8: 55-57. REEDER, W. G. 1960. Two new rodent genera from the Oligocene White River forma- tion (Family Heteromyidae). Fieldiana: Geology, 10: 511-524 SCHLAIKJER, E. M. 1935. Contributions to the stratigraphy and paleontology of the Goshen Hole area, Wyoming. Part LV. New vertebrates and the stra- tigraphy of the Oligocene and early Miocene. Bull. Mus. Comp. Zool., 76: 95-189, 41 pls. Woop, A. E. 1933. A new heteromyid rodent from the Oligocene of Montana. Jour. Mam., 14: 134-141. 1935. Evolution and relationships of the heteromyid rodents with new forms from the Tertiary of western North America. Ann, Carnegie Mus., 24: 73-262. 1937. The mammalian fauna of the White River Oligocene. Part II. Rodentia. Trans. Amer. Philos. Soc., (m.s.) 28: 155-269, 11 pls. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DeEcEMBER 15, 1961 NuMBER 147 AUSTRALIAN CARABID BEETLES VIII. LEIRADIRA, ESPECIALLY THE TROPICAL SPECIES By P. J. DARLINGTON, JR. Museum of Comparative Zoology, Cambridge, Mass. This is the third part of this series to be devoted to flightless Carabidae of zoogeographiec importance from tropical eastern Australia. Pertinent earlier parts, including a locality list and a discussion of transition of carabid faunas in wet forests from New Guinea to Tasmania, are listed at the end of the present paper. Information on deposition of types and a brief note on methods will be found at the beginning of Part VI (1961c). The present paper is concerned with the pterostichine genus Lewiradira. Typical Leiradira are very distinetive Carabidae, dis- tinguished from (for example) Notonomus by : deep-bodied form ; heavy, sometimes more or less deflexed head, with mandibles somewhat longer and more acuminate than in Notonomus; gen- iculate antennae; mentum tooth absent ; and inner lobe of maxilla lined with what look like long slender teeth rather than bristles (see following key). These characters led Tschitschérine (1902, p. 506) to place Leiradira in the tribe Deliniini, far removed from Notonomus. However, some of my new North Queensland species reduce the gap between Leiradira and Notonomus and suggest that tribal separation of the two genera may not be justified and that, although the genera are distinct, they (and Delinius) may be derived from a common ancestor. This is something to be de- cided in the future, by the next reviser of the generic classifica- tion of Australian Pterostichini. As I now understand the genus, Lewradira (in a broad sense) includes about a dozen species and is discontinuously distributed, chiefly in rain forest, along the eastern edge of Australia from just below Cooktown in tropical North Queensland south at least to the lower edge of the Dorrigo plateau in warm temperate BREVIORA No. 147 north-central New South Wales. The genus divides into three well marked subgenera, which have geographical as well as structural bases. They are distinguished in the following key, which ineludes Notonomus for comparison. The tropical species of Notonomus will be treated in another paper. Co 6. Key to subgenera and some species of LEtRADIRA Mentum tooth normal, long, deeply emarginate; inner lobe of maxilla lined with many coarse setae not in single row; head not deflexed; mandibles shorter; antennae not geniculate, lst segment shorter than segments 2 + 3; form usually more depressed (but variable) ..... (Notonomus ) Mentum tooth broad and short, or absent; inner lobe of maxilla lined with single row of about 6 to 8 slender teeth; head often (not always) more or less deflexed, with mandibles usually (not always) longer and more acuminate than in Notonomus; antennae often (not always) geniculate, with Ist segment usually (not always) as long as or longer than segments 2 + 3; form relatively deep-bodied (Leiradira, sensu GRRE oe Ses weak AMM Fle peels 2 ra Ad Yop ieee PAC h ns tac earned CUA Be 8 2 Labrum moderately emarginate, 6-setose; posterior-lateral pronotal setae on thickened margin; (mentum tooth present but broad and short ) (North Queensland) (subgenus Metadira)........................ 4 Labrum either deeply, almost semicircularly emarginate or only 4- setose; posterior-lateral pronotal setae inside (not on) thickened margin) (mentum tooth present) or absent)... 90+ eeoe eee 3 Labrum moderately emarginate, 4-setose; mentum without tooth; man- dibles shorter, but not so short as in Notonomus (northern New South Wales and South Queensland) (Leiradira, sensu stricto) ........... 3 or more full species Labrum semicirecularly emarginate, 6-setose; mentum tooth present (but very broad and short); mandibles longer than usual (South Queens- land and Eungella Range) (subgenus Stomimorphus)..............8 Elytral intervals very unequal (odd several times wider than even ones) ; Ist antennal segment shorter than segments 2 + 3; (color green- DUTDPLE) epbat ls Says cece AS emaee ees ara iirc ree teers _.ignifer Elytral intervals less unequal, or equal; Ist antennal segment usually equalito.orlongerythan2'c-F Si4. Asaun Boomer sede ae ie eee 5 Elytral intervals subequal and striae normal, not widened except some- TIME Bet Cait? VOX | -hy.0 8 Gossett ES EA Oe ni ee 6 Hither odd elytral intervals wider than even ones or striae partly wide 9nd Opaque f 9.3 cae acpertawWe nee oe Aker moce eke A fabate np tipese of Wangers GS-19 enim) i idarkesblue-piin ple mer ar errr renee: alticola Simeuilcrom ClaleilPasy semen) O LCG Soosoucomncoedundsutaauwereycoce soror 1961 AUSTRALIAN CARABID BEETLES a Odd elytral intervals wider than even ones; elytral striae not much ba | widened; 3rd elytral intervals usually 3- or 4-punctate; scutellar striae weak or obsolete (variable) ; form more slender....... . alternans — Elytral intervals nearly equal; elytral striae widened and opaque espe- cially behind middle; 38rd elytral intervals always 2-punctate; scutellar striae short but deep; form less slender ......... _... opacistriatus G llamagre CGM iain, Or min) sleccestonoboavecenes _...... violaceus —— Shinmilee(G, Wa wai.) .s5cesce0eseane ee Mere re arty ed um 2.8 to) 9. Prothorax wider (W/L 1.14-1.19); duller Pes _. jacobi — Prothorax narrower (W/L 1.06-1.08); more shining............ tenwis Of the three subgenera, Metadira is most like Notonomus. Even the least modified Metadira, aurifer, differs from Notonomus in having a shorter mentum tooth and a regular row of about 8 slender tooth-like processes on inner edge of maxilla instead of more than 20 thickened setae less regularly arranged as in, for example, Notonomus doddi Sloane, but there are two indications besides the general similarity that suggest a real relationship. One is the position of the posterior-lateral prothoracic setae, on (not inside of) the thickened margin at basal angles in both Metadira and the tropical species of Notonomus. The other is the fact that some Metadira and some tropical Notonomus have elytral striae conspicuously widened and opaque. This is an unusual character, unlikely to have evolved independently in the two genera. METADIRA subgen. n. Form varying from that of a convex Notonomus to that of typical Leiradira. Head stout, sometimes somewhat deflexed ; eyes small, genae prominent (but variable) ; 2 supraocular setae each side; mandibles varying (in different species) in length and eurvature; labrum moderately emarginate, 6-setose; antennae with basal segment varying (in different species) from shorter to longer than next 2 segments together; frontal foveae short but well defined ; mentum tooth very short and broad, broadly emargin- ate; inner lobe of maxilla with single row of setae so thickened as to resemble long slender teeth; palpi slender in both sexes. Prothorax cordate, with narrow margins; baso-lateral impressions small but deeply impressed; usual 2 marginal setae each side, posterior ones on thickened margins at basal angles. EHlytra usu- ally with (sometimes almost without) basal margin; humeri + dentate; striae entire, narrow or widened (in different species) ; intervals equal or unequal (in different species) ; narrow 10th 4 BREVIORA No. 147 interval present or indicated posteriorly ; 3rd interval typically with 2 dorsal punctures (usually only 1 in alticola and soror, and usually 3 or 4 in alternans). Last 3 ventral segments weakly, variably sub-impressed across base but not suleate; mesosternum and prosternal process not setose. Male with anterior tarsi slightly dilated, 3 segments with squamae; ¢ with 1, 2 2 setae each side last ventral segment. Genotype: Leiradira aurifer Darlington (below). Five species of this subgenus are now known. All occur on (different parts of) the Atherton Tableland/or Mts. Bartle Frere and Bellenden Ker (see map) and the northernmost of the five species extends north nearly to Cooktown. Three of the species (alternans, soror, opacistriatus) may be (distinct) geographical forms of one original widely distributed stock (see notes under alternans). The other two species are very distinct, localized endemiecs. All the species are usually found (by day) under cover on the ground, in rain forest, but I got one opacistriatus in good savannah woodland a few hundred feet outside rain forest by the Davies Creek Road. LEIRADIRA (METADIRA) AURIFER N. sp. Form as figured (Fig. 1), of a rather convex Notonomus, but with characters of Metadira as given above; head and pronotum bright green, pronotum with variable purplish reflections, elytra dark copper-purple, lower surface and legs black, mouth parts and antennae brownish; surface shining, except bottoms of striae opaque. Head 4, or slightly less width prothorax; mandibles shorter than usual in Leiradira, rather strongly curved, acumi- nate apically; eyes moderate (in genus), genae ¢. wide as eyes, weakly rounded to neck; 1st antennal segment c. 45 (by measure- ment) length of 2nd + 3rd segments together ; middle segments nearly 2X long as wide. Prothorax c. 49 (+) wider than long at middle, moderately narrowed behind, more in front; base c. Yo wider than apex; apex subtruncate or very broadly emargi- nate with angles scarcely advanced, margined only near sides; base slightly sinuously subtruneate, deeply margined toward sides but not at middle; sides broadly rounded for much of length, sinuate near base; basal angles ec. right, secareely blunted; dise moderately convex with middle line distinct and reaching base, 1961 AUSTRALIAN CARABID BEETLES if | | e Daintree | | | | | | | | e Mossman Mt. Lewis\ NX ~~ SS NX S\ ~ SS ¢ Kuranda | e | g WE) | - Gdirns ./ Sure f xO rs 6 Q@ ia) : ar pe! AS ee .. \ M om is \ Mero Mt. B.K ) . e Lie Fa: \ Oe NE ° | Qv Mo Mt. B.F or ase ed One Z con KSEE 2 e \ ww, Ww be, : Ravenshoe : Known distribution of Leiradira, endemic subgenus Meladira, in North Queensland. The finely dotted line is the approximate eastern edge 9 Aly of high land (the Atherton Tableland ete.). No. 1, aurifer; alticola; 3, alternans; 4, soror; 5, opacistriatus, whichoceurs also north beyond the limits of the map almost to Cooktown. 6 BREVIORA No. 147 transverse impressions almost obsolete, and baso-lateral impres- sions short, linear, connected with side margins by deeply im- pressed marginal grooves. EHlytra % (or slightly more) wider than prothorax; basal margin entire, strongly raised, slightly scalloped, rectangular at subdentate humeri; striae very wide and opaque especially posteriorly; intervals convex, very un- equal, odd ones several times wider than even, 7th strongly con- vex but not sharply carinate at base; a narrow 10th interval present posteriorly; each 3rd interval usually 2-punctate just behind middle and near apex (an additional puncture present near middle on 1 side in 1 specimen). Length 15-19; width 5.0- 6.2 mm. Holotype ¢ (M.C.Z. Type No. 30,382) and 9 paratypes from mountains north of Kairi, Atherton Tableland, North Queens- land, 3000-4000 ft., Dee. 1957; and 3 paratypes from south end of Davies Creek road, Atherton Tableland, Dec. 1957: all speei- mens taken by myself, in rain forest. The two localities are not far apart in the mountain mass that lies on the Atherton Table- land between the Mareeba-Kuranda road on the north and Yungaburra-Lake Barrine road on the south. See preceding discussion and key for place of this species among other Leiradira. Although it has striking specific charac- ters (color, extreme alternation of elytral intervals), it is the least specialized member of the group in form, mandibles, and leneth of Ist antennal segment. LETRADIRA (METADIRA) ALTICOLA Nn. sp. Form as figured (Fig. 2), prothorax strongly cordate, elytra narrowed anteriorly ; rather strongly convex ; black, usually with purplish tinge, margins of elytra blue; moderately shining. /Tead c. 34 or slightly less width prothorax; mandibles shehtly longer and less curved than in aurfer; eyes small, genae usually longer and more prominent than eyes, strongly convex; antennae very short, Ist segment shehtly longer than next 2 together (by meas- urement), outer segments scarcely longer than wide. Prothorax slightly (less than 49) wider than long at middle; apex truncate with angles scarcely advanced, not margined; base virtually truncate, strongly mareined toward sides; sides arcuate for c. °4 of length (sometimes subparallel near middle), strongy sinuate c. ¥ or more of length before base, then parallel or diverging to base; basal angles well defined, right or acute; dise with middle line well impressed, very deep basally, transverse impressions 1961 AUSTRALIAN CARABID BEETLES Tf obsolete, baso-lateral impressions deep, sublinear, connected with side margins by deep basal grooves. Elytra c. 4 (or slightly less) wider than prothorax, narrowed basally; basal margins entire, scalloped, forming sharply defined right angles at sub- dentate humeri; striae narrow, slightly wider apically, with a little dull sculpture at bottom; intervals subequal, shghtly con- vex; 3rd interval with 1 puncture near or behind middle, usually without posterior puncture, but latter present on 1 side in 1 individual (see Fig. 2). Parts of lower surface (sides of meso- sternum, first ventral segment, and parts of other ventral seg- ments) more or less punctate or subpunctate. Length 15-19; width 5.2-6.5 mm. Holotype ¢ (M.C.Z. Type No. 30,383) and 3 paratypes from Mt. Bellenden Ker, E. side, 3000-4500 ft., Jan. 1958, and 1 para- type, Mt. Bartle Frere, W. slope, near 5000 ft., Dec. 1957. These 2 mountains are close together at the eastern edge of the Atherton Tableland south of Cairns, North Queensland. All specimens taken by myself, in mountain rain forest. Distinguished from other Leiradira as indicated in the preced- ing key; unique in form, and notable for very short antennae. Lerrapira (METADIRA) ALTERNANS Darlington This species (Fig. 3) was described by me in 1953 (p. 90) from Malanda ete. It is widely distributed on the central-southern Atherton Tableland, the limits of its known distribution being Atherton, Lake Barrine, the lower western slope of Mt. Bartle Frere, and Mt. Fisher southwest of Millaa Millaa. It may be represented by soror (below) on the eastern slope of Mt. Bellen- den Ker (and perhaps elsewhere on the eastern slope of the Tableland) and by opacistriatus on the northern edge of the Tableland and northward. LeIRADIRA (METADIRA) SOROR N. sp. Small, slender, convex; shining black with silky lustre but no distinct metallic color. Head *4 (+) width prothorax ; mandibles relatively weakly arcuate; eyes small, genae slightly more promi- nent than eyes, broadly convex; antennae with Ist segment longer than next 2 together. Prothorar narrowly subcordate, as long as or slightly longer than wide; apex subtruncate with angles scarcely advanced, not margined at middle; base sub- truncate except slightly rounded toward sides, not margined 8 BREVIORA No. 147 except near sides; sides broadly arcuate, subparallel before mid- dle, strongly sinuate about 149 of length before base; basal angles c. right, scarcely blunted; dise with middle line rather fine but reaching base, transverse impressions weak, baso-lateral impres- sions nearer sides than middle, linear, deep, joining lateral margins at base. EHlytra *49 (or less) wider than prothorax, subparallel, weakly narrowed anteriorly; basal margin deep, scalloped, rectangular at subdentate humeri; striae fine; intervals convex, subequal ; 3rd interval usually 1-punctate slightly behind middle, posterior punctures usually absent (1 present on 1 side in 1 specimen). Parts of lower surface (sides of mesosternum, Ist ventral segment, parts of other ventral segments) variably punctate. Length 11.0-12.5; width 3.3-3.7 mm. Holotype ¢ (M.C.Z. Type No. 30,384) and 2 paratypes all from E. side Mt. Bellenden Ker, ¢. 3000 ft., North Queensland, Jan. 1958, taken by myself in rain forest. This species most resembles alternans (above) but is smaller, narrower, with less alternation of elytral intervals, and fewer punctures on 3rd elytral interval. LEIRADIRA (METADIRA) OPACISTRIATUS (Sloane ) Sloane first (1902, p. 319) described this species in Notonomus, then (1913, p. 409) ruled it out of that genus. The types were said to be from Cairns, collected by Froggatt; they probably really came from the mountains near Cairns. I could not locate the types in Australia, but my specimens fit the deseription reasonably well. They are from the Davies Creek road on the northern Atherton Tableland; near Black Mt. about 20 miles north of Kuranda (these are probably virtual topotypes) ; Mt. Lewis southwest of Mossman; and Mt. Finnegan south of Cook- town. LEIRADIRA, sensu stricto Castelnau 1867, p. 72. Csiki 1929, p. 500 (see for additional references and list of previously described species ). Genotype, by present designation: Leiradira auricollis Castel- nau (Fig. 4) (genus originally based on this species and latreille: Castelnau ). Leiradia, sensu stricto apparently occurs only below the tropics, from the Blackall Range ete. in South Queensland south at least to the lower (eastern) edge of the Dorrigo plateau. The 1961 AUSTRALIAN CARABID BEETLES 9 subgenus includes at least three full species, perhaps more. I have not studied them and cannot establish synonymies. Subgenus STOMIMORPHUS Straneo Straneo 1953, p. 1. Leiradira-like Carabidae with mandibles long, acuminate; labrum deeply emarginate, 6-setose; mentum tooth present but short, broad; antennae with Ist segment longer than 2nd + 3rd together; posterior-lateral setae of pronotum inside thickened margin. Genotype: Stomimorphus violaceus Straneo. This subgenus is apparently confined to southern and central eastern Queensland. The type locality of violaceus is simply Queensland. I have what may be this species from Maleny, on the Blackall Range, and additional, smaller species are described here from Mt. Jaccb and the Eungella Range. LEIRADIRA (STOMIMORPHUS) VIOLACEUS (Straneo) Straneo 1953, p. 1. Described as violaceous; length 14.7 by 4.9 mm. The unique type isa é from ‘‘ Australia, Queensland’’ in the Straneo Collec- tion. Two specimens that I collected near Maleny, on the Blackall Range, South Queensland, in rain forest, are greenish violaceous and larger than the type, but I hesitate to describe them without more material to show extent of variation. LEIRADIRA (STOMIMORPHUS) JACOBI 0. sp. Form as figured (Fig. 5); labrum, mentum tooth, and other characters as described for Stomimorphus; head and prothorax greenish black, elytra purplish darker dorsally, lower surface and appendages dark; moderately shining, elytra slightly duller and with distinct, fine reticulate microsculpture. Head not quite 34 (ec. .72) width prothorax. Prothorax subcordate, slightly less than 15 wider than long at middle, width/length 1.14-1.19; apex subtruncate; base slightly emarginate at middle and rounded toward sides; base and apex not distinctly margined; sides broadly rounded for most of length, slightly sinuate before base ; basal angles slightly obtuse, slightly blunted ; disc convex, middle line fine, transverse impressions weak, baso-lateral impressions linear, moderately impressed, not punctate. Elytra %4 (or more) 10 BREVIORA No. 147 wider than prothorax, slightly narrowed anteriorly; basal margins entire, forming obtuse-right angles at humeri; latter finely bluntly subdentate ; striae moderately impressed, entire, not punctate; dorsal intervals equal, slightly convex; 10th interval present posteriorly; 3rd intervals 2-punctate in all specimens, near middle and posterior °4 (but position of punctures slightly variable). Secondary sexual characters normal, i.e. ¢ with an- terior tarsi shghtly dilated with 3 segments squamulose below ; and ¢ with 1, @ 2 setae each side apex last ventral segment. Length 9.8-12.0; width 3.3-4.1 mm. Holotype ¢ (M.C.Z. Type No. 30,385) and 9 paratypes all from Mt. Jacob, c. 45 miles south of Gladstone, South Queens- land, c. 2000 ft. altitude, Mar. 1958, taken by the Darlingtons, in rain forest. Superficially this species is deceptively similar to Leiradira, sensu stricto, but the species’ technical characters are as in Stom- morphus. LEIRADIRA (STOMIMORPHUS) TENUIS 0. sp. Form as figured (Fig. 6); with characters of Stomimorphus as here given; dark purplish, lower surfaces and appendages dark; shining, elytra with faint fine microreticulation. Head °4 or slightly less width prothorax. Prothorax narrow, width/length 1.06-1.08 (all specimens) ; apex subtruneate or very broadly emar- ginate; base broadly emarginate at middle, shghtly rounded toward sides; base and apex not distinctly maregined; sides weakly arcuate for much of leneth, slightly sinuate near base ; dise with fine middle line, very weak transverse impressions, moderate linear baso-lateral impressions, not punctate. Hlytra c. % wider than prothorax ; basal margin entire, forming slightly obtuse (nearly right) angles at humeri; latter bluntly subden- tate; striae entire, moderately impressed, not punctate; intervals slightly convex, subequal; 10th interval present posteriorly ; 3rd interval 2-punctate, near middle and apical °4. Length 10.0- 11.5; width 3.3-3.5 mm. Holotype ? (M.C.Z. Type No. 30,386) and 2 ? @ paratypes all from the Eungella Range, west of Mackay, Queensland, 2000- 3000 ft., Nov. 1957, taken by the Darlinetons, in rain forest. This resembles and may be related to yacobi (above) but the prothorax of the present species is narrower and the elytra more shining. 1961 AUSTRALIAN CARABID BEETLES 11 REFERENCES CASTELNAU, F. DE 1867. Notes on Australian Coleoptera: pp. 1-139 (Reprinted in Trans. R. Soe. Victoria, 8: 30-38, 95-225, 1868). CsIKI, E. 1929. Junk-Schenkling Coleoptera Catalogus, Pars 104, Carabidae, Harpalinae ITI. DARLINGTON, P. J., JR. 1953. Australian carabid beetles II. Some new Pterostichini. Psyche, 60: 90-101. 196la. Australian carabid beetles IV. List of localities, 1956-1958. Psyche, 67: 111-126. 1961b. Australian carabid beetles V. Transition of wet forest faunas from New Guinea to Tasmania. Psyche, 68: 1-24. 1961ce. Australian carabid beetles VI. The tropical and some subtropical species of Pamborus, Mystropomus, and Nurus. Breviora, No. 142: 1-13. 1961d. Australian carabid beetles VII. Trichosternus, especially the tropical species. Psyche (In press). SLOANE, T. G. 1902. A revision of the genus Notonomus ... Proc. Linn. Soc. New South Wales, 27: 252-324. 1913. Revisional notes on Australian Carabidae. Part IV. The genus Notonomus. Proce. Linn. Soe. New South Wales, 38: 404-449. STRANEO, S. L. 1953. Nuovi Pterostichini VII. Doriana, Suppl. Mus. Civico Storia Nat. ‘‘G. Doria,’’ Genoa, 1, no. 36, 1-12. TSCHITSCHERINE, T. 1902. Notes sur les Platysmatini de 1’Australie. Horae Soc. Ent. Rossicae, 35: 502-534. 12 BREVIORA No. 147 ; QUA: / Fig.1. lLeiradira (Metadira) aurifer n. sp. Fig. 2. Leiradira (Metadira) alticola n. sp. Fig.3. Leiradira (Metadira) alternans Darlington Fig. 4. Leiradira (sensu stricto) auricollis Castelnau Fig. 5. Leiradira (Stomimorphus) jacobi n. sp. Fig.6. Leiradira (Stomimorphus) tenuis n. sp. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 18, 1961 NuMBER 148 AUSTRALIAN CARABID BEETLES IX. THE TROPICAL NOTONOMUS By P. J. DARLINGTON, JR. Museum of Comparative Zoology, Cambridge, Mass. This is the last of four papers on flightless Carabidae of zoo- geographic importance from tropical eastern Australia — but these papers are part of my general series on Australian carabid beetles, which will be continued. Pertinent earlier parts of the series, including a locality list (with maps) and a discussion of the transition of wet forest carabid faunas from New Guinea to Tasmania, are listed with references at the end of the present paper. Information on deposition of types will be found at the beginning of Part VI (1961c). In this as in other parts of this series, proportions stated as simple fractions are based on actual measurements made with a ruled ocular in the microscope. The present paper concerns the tropical species of the dominant eastern Australian pterostichine genus Notonomus. The genus as a whole ranges along the eastern edge of Australia from the base of the Cape York Peninsula (somewhere between Daintree and Cooktown) to Victoria and Tasmania in the south, extending west into the eastern edge of South Australia, with one very distinct species geographically isolated in southwestern Australia and two species described from New Caledonia. I have dealt with some of the tropical Australian species before (1953) but several additional species are now to be described and a new key is necessary. The tropical species of Notonomus are all apparently related among themselves and form what may be called the doddi sub- group of Sloane’s (1913) kingi group of the genus. The doddi subgroup is characterized by posterior-lateral setigerous punc- tures of pronotum on (not inside) thickened margin at basal angles; elytra fully and strongly striate; 3rd interval not more than (but sometimes less than) 2-punctate (except that rarely 2 BREVIORA No. 148 individuals have extra adventitious punctures) and 5th and 7th intervals impunctate; 8th interval (10th in saepistriatus) nar- row and convex (least so in masculinus) ; metepisterna short; intercoxal declivity of prosternum flat, without setae; tarsi not striolate above; posterior tarsi with Ist seegment rather long (but not quite so long as next 2 together) and with claw segment glabrous below; and ¢ with 1, 2 usually 2 setae each side last ventral segment (1 seta each side in 2 montellus, and single anal setae may be either missing or duplicated in individuals of other species), and 6 with anterior tarsi dilated with 3 seg- ments squamulose below, 2 usually with tarsi unmodified (but some [not all] 22 of doddi, flos, montorum, and masculinus have front tarsi with Ist segment squamulose). Other characters common to all members of the doddi subgroup, and therefore not repeated in the following brief descriptions, are: prothorax with apex subtruncate or broadly emarginate with angles not or not much advanced beyond are of emargination, margined at sides but not at middle; base slightly sinuously subtruncate, margined only at sides; side margins rather narrow, only slightly wider posteriorly ; dise with middle line nearly entire, transverse im- pressions weak or obsolete, baso-lateral impressions long, linear, moderately impressed, not punctate; elytra with basal margin c. rectangular at finely subdentate humeri; narrow extra (10th, or 12th in saepistriatus) submarginal interval present at least posteriorly. Although they agree in these characters (with ex- ceptions noted), the species of the doddi subgroup differ re- markably among themselves in some other characters, given in the following key. EXPLANATION OF Map ON PAGE 3 Known distribution of Notonomus in tropical Queensland. The finely dot- ted line is the approximate eastern edge of high land (the Atherton Table- land ete.). No. 1 (arrow) indicates occurrence of Notonomus transitus endem- ic on the Eungella Range south of the limits of the map; 2, N. doddi in the Herberton-Atherton area, and also (arrow) south of the limits of the map on the Kirrama Range, the Mt. Fox plateau, and the Mt. Spee plateau; 3, montel- lus; 4, dimorphicus ; 5, spurgeoni; 6, flos; 7, montorum with subspecies azul on Mt. B(ellenden) K(er); 8 (arrow) ellioti, endemic on the Elliot Range south of Townsville, beyond limits of map; 9, masculinus; 10, saepistriatus. Note the wide gap between the Atherton Tableland species and those on the mountains of the Mossman-Daintree area; the 3 species on the latter moun- tains are apparently derived from one secondary ancestor. 1961 AUSTRALIAN CARABID BEETLES ~ <-------— Distribution of tropical Notonomus. 4 BREVIORA | No. 148 So far as I know, all the tropical Notonomus are confined to rain forest, and they may all be derived from a single primary ancestor that invaded tropical rain forest from the more south- ern part of eastern Australia. N. transitus, on the Eungella Range in east-central Queensland, may be a (presumably modi- fied) derivative of this ancestor. Of the more northern species (see map), doddi of the Dividing Range system from Mt. Spee to Atherton, and montellus of Mts. Bartle Frere and Bellenden Ker, may be related to each other; montorum, of Mts. Bartle Frere and Bellenden Ker, and elliott, of the Elliot Range south of Townsville, may be relicts of one more widely distributed secondary ancestor; masculinus and saepistriatus, with allopatric ranges on the Atherton Tableland, may be related to each other in spite of their structural differences; and the three species north of the Atherton Tableland (dimorphicus, flos, spurgeont) may be interrelated and may now be in process of radiation from one secondary aneestor. All this suggests the probable complex- ity of evolution of the group in the rain forests of North Queens- land, especially on and near the Atherton Tableland. The radia- tion of Trichosternus in the same area (Darlington 1961d) may have been even more complex. Incidentally, no species of either Notonomus or Trichosternus has yet been found on the heavily rain-forested mountains that le on the northern part of the Atherton Tableland between the Yungaburra-Lake Barrine road (on the south) and the Mareeba-Kuranda road (on the north), although these mountains are the home of a striking endemic Leiradira (aurifer Darlington 1961e). Key to tropical species of Noronomus (doddi subgroup) 1. Species with all following characters: elytral striae not widened on disc, and intervals normal in number and not much interrupted, and 3rd THRE yall Cloweseul {WNMCOTN 5.5 oop cadasuaccsedcounasncu vonaee wae — Either striae wide and opaque on dise (not just on declivity), and/or intervals more numerous or much interrupted, and/or 3rd interval with- Oty CASE Nl FOWUNEMOKS san ceccantcoosdcuaasnccce af 7 2. Sides of ventral segments 4-6 extensively punctate (more so than Ist ventral); (dull black, pronotum virtually smooth)......... _transitus — Ventral segments not punctate or, if punctate, lst most strongly so... .3 3. Pronotum deeply and closely transversely strigulose............... 4 — Pronotum only normally (lightly or faintly) transversely strigulose. ..5 4. Brownish black, usually larger (12.5-16 mm.); 2 with 2 setae each sidelast ventral sceoment* A070 Set pk ter neh atone ee ES oper doddi 1961 AUSTRALIAN CARABID BEETLES 5 — Bluish or purplish; usually smaller (10.5-13 mm.); 92 with 1 seta each sidewlastaventralyess eee ean tae ee boo en eva eset montellus 5. Male slender (Fig. 3) ( 2 less so); (bicolored) .......... dimorphicus =A OUDESOXE SA SLOULEE Mees sic UN et iL ree chen evEte cle ohn aicns oA Rae, set a elen ten 6 6. Wholly purplish or bluish; length c¢. 12-17 mm. ............ spurgeoni — Bicolored, head and pronotum purple or coppery, elytra black; broader cual, Ireaqere, Ikeradac , IMT-PAN) si; 5 6 aah aduacaseadaebenbsovonsoecus flos 7. Elytral intervals not much interrupted (3rd without dorsal punctures) 8 — Elytral intervals much interrupted (38rd with or without dorsal pune- TUTE SY lo st i Sb i ls Sek Ca car hae uae dean hep ear at irate. kates Pcie CR ta ag Bieta cache crt 9 See Normebroadern sdullee ep se nt detects | co as ttn cite oe ee montorum da TeOMISNeOrspUTplIS har ea ae een eee montorum, sensu stricto b. Bluish with elytral margins bright blue ................ subsp. azul ——) Horm) narrower; more shinies, purplish’ ]..55--2--c)o.se se cee: ellioti 9. Elytron with 9 intervals, without dorsal punctures........ masculinus — Elytron with 11 intervals plus narrow submarginal one posteriorly (7th interval tripled), and with dorsal punctures ............. saepistriatus NOTONOMUS TRANSITUS Ni. sp. With characters of doddi subgroup; form as figured (Fig. 1), rather broad and depressed (in group); black, upper surface sometimes slightly aeneous, marginal channels of elytra usually cupreous or dull greenish; most of upper surface with fine iso- diametrie microsculpture. Head 7% or slightly less width pro- thorax, without noticeable unusual characters. Prothorax c. 4 or less wider than long; sides weakly arcuate for most of length, slightly sinuate almost at base; basal angles slightly obtuse, not much blunted; disc smoother than usual, with transverse strigu- lation very faint or absent. Hlytra slightly less than 44 wider than prothorax; striae rather fine, not punctate, not widened even on declivity; intervals nearly flat on disc, more convex laterally, the discal ones subequal; 3rd interval 2-punctate, be- hind middle and behind apical “4. Lower surface nearly impune- tate anteriorly but much of abdomen finely and closely but rather irregularly punctate. Secondary sexual characters nor- mal; all 22 (16) with simple anterior tarsi. Length c. 13-18; width 4.5-6.1 mm. Holotype ¢ (M.C.Z. Type No. 30,387) and 26 paratypes all from the Eungella Range, west of Mackay, east-central Queens- land, 2000-3000 ft. altitude, Nov. 1957, taken by the Darlingtons, in rain forest. 6 BREVIORA No. 148 Superficially, transitus is similar to the common, variable N. nitidicollis Chaudoir of South Queensland. There may be a real relationship between these two species. However, transitus leads toward the North Queensland species of the genus in position of the posterior-lateral prothoracic setae, on the thickened margin (inside the margin in nitidicollis), and it differs from nitidicollis in other specific characters. For example, transitus is broader and more depressed, with flatter elytral intervals than nitidicollis, and the extensive punctation of the abdomen of transitus is lack- ing in nitidicollis. The northern limit of nitidicollis, incidentally, is probably Mt. Jacob, about 45 miles south of Gladstone, South Queensland. Of the North Queensland species, transitus is prob- ably nearest doddi but differs in being broader and more de- pressed, with flatter elytral intervals, without special pronotal sculpture, but with abdominal punctation. NOTONOMUS bDoppI Sloane The type locality is the Herberton District, Atherton Table- land, North Queensland (Sloane 1913, p. 489). Specimens that I collected on the mountains south and west of Atherton (be- tween Atherton and Herberton) are virtually topotypes. The species extends south (discontinuously) along the Dividing Range system to the Kirrama Range, the Mt. Fox plateau, and the Mt. Spee plateau not far north of Townsville. The exceptionally close and deep transverse strigulation of the pronotum is apparently always present in this species, but the other pronotal microsculpture is dimorphic. Most individuals have the head very finely and the elytra more coarsely (but still finely) isodiametrically reticulate and the pronotum longitudi- nally roughened between the deep transverse strigae. However, my series of 11 specimens from the Mt. Spec plateau includes 3 66 with pronotum without longitudinal roughening, although the 4 other ¢¢ and all 4 22 have the roughening present, as do all specimens from other localities. The length of this species is c. 12.5-16.5 mm., with the average size decreasing southward. NOTONOMUS MONTELLUS N. sp. With characters of doddi subgroup as given above; form as figured (Fig. 2), small, shghtly depressed; bluish or purplish black; head and elytra with fine isodiametric microsculpture, less distinet on pronotum. Head *% or shghtly less width pro- thorax, without obvious unusual characters. Prothorax VY (+) =~) 1961 AUSTRALIAN CARABID BEETLES wider than long; sides broadly arcuate for most of length, briefly sinuate before c. right, scarcely blunted basal angles; main (cen- tral) part of disc with many deeply impressed, close-spaced, slightly irregular, transverse strigae; surface of disc otherwise not distinctly punctate. Hlytra 4% (+) wider than prothorax ; striae well impressed, not punctate, not or scarcely widened even on declivity ; intervals moderately convex, subequal on disc; 3rd 2-punctate near middle and apical 14 (punctures slightly vari- able in position). Lower surface with a little scattered (variable) punctation especially on sides of mesosternum and Ist ventral segment. Secondary sexual characters normal except 2 with 1 seta (not 2) each side last ventral segment. Length ce. 10.5-13.0; width 3.5-4.4 mm. Holotype ¢ (M.C.Z. Type No. 30,388) and 5 ¢¢ 4 22 para- types from Mt. Bartle Frere, west slope, 3000-5000 ft., Dee. 1957 ; and 1 additional 2 (not a type) from Mt. Bellenden Ker, east side, about 4500 ft., Dec. 1957. These two mountains are close together at the east side of the Atherton Tableland, south of Cairns, North Queensland. Specimens all taken by the Darling- tons, In mountain rain forest. This small Notonomus is closest in technical characters to N. doddi Sloane, of the Dividing Range system, and may be related to it, but differs in smaller size, more depressed form, color, and presence of only 1 seta each side 2 last ventral segment. The pronotal sculpture resembles that of the exceptional doddi from Mt. Spee, with deep transverse strigulation but without longitud- inal roughening. NOTONOMUS DIMORPHICUS nh. sp. With characters of doddi subgroup as given above; ¢ (Fig. 3) exceptionally slender, 2 (Fig. 4) less so; head and prothorax aeneous, elytra purplish black; moderately shining with fine iso- diametric microsculpture on head and elytra, not distinct on pronotum. Head c. ?4 or more width prothorax, without notice- able unusual characters. Prothorax shghtly longer than wide in slender 6 6, slightly wider than long in stouter 2 2 (but slightly variable in both sexes); sides rather weakly rounded, in 4 4 nearly straight and converging both before and behind sub- median curve but more regularly arcuate in 2 @ ; sides briefly, variably sinuate just before base; basal angles right or slightly obtuse, not much blunted; surface of dise with faint transverse strigae, not punctate. Hlytra c. Y5 (=) wider than prothorax; 8 BREVIORA No. 148 striae not widened except slightly so (striae 1 and 2) at extreme apex, on declivity ; intervals moderately convex, not interrupted, subequal on disc; 3rd 2-punctate, near middle and posteriorly (middle puncture sometimes duplicated). Lower surface nearly impunctate. Secondary sexual characters (other than dimor- phism of form) normal. Length c. 12-15; width 3.6-4.6 mm. Holotype ¢ (M.C.Z. Type No. 30,389) and 7 66,7 22 para- types all from Mt. Lewis, near Mossman, North Queensland, c. 8000 ft., Dec. 1957, collected by the Darlingtons, in rain forest. Males of this species are unique in form at least among the tropical species of the genus, and at first I thought they might represent transition toward Leiradira, but the mandibles, an- tennae, and setae of the inner edge of the maxilla are as usual in Notonomus. The 2? are close to N. spurgeoni (below) but are more slender and bicolored. All specimens that I refer to the present species were taken within a comparatively small area of continuous mountain forest, and I feel sure they represent a single population. A single 2 that I refer to spurgeoni, although labeled from Mt. Lewis, may have been taken outside of and below the area occupied by dimorphicus. NOTONOMUS SPURGEON! Darlington This is the most northern known species of Notonomus. I described it (1953, p. 98) from a series from Mt. Spurgeon, about 12 miles northwest of Mt. Lewis in the same mountain system. Three specimens from Thornton Peak, northeast of Daintree, near 4000 ft., Dee. 1957, taken by the Darlingtons in mountain rain forest, seem to be the same species, although the color is bluish rather than purplish. A single 2 from Mt. Lewis is also apparently referable to this species (see note under dimorphicus, above). NOTONOMUS FLOS N. sp. With characters of doddi subgroup as given above; form as figured (Fig. 5), rather large and broad in group; black, head and pronotum bright violaceous or cupreous ; moderately shining, elytra shghtly duller; fine isodiametric microsculpture very dis- tinct on elytra, absent or indistinct on head and pronotum. Head %, (+) width prothorax, without obvious unusual characters. Prothorax large, ¥ to 44 wider than long; sides broadly arcuate for almost entire length, usually minutely sinuate at base (the sinuation involves hardly more than widening of the marginal 1961 AUSTRALIAN CARABID BEETLES 9 bead); basal angles obtuse (sometimes nearly right); surface of dise with weak transverse strigae. Elytra slightly wider than prothorax ; striae well impressed, not punctate, not or not much widened on dise but inner ones wider and dull on declivity ; intervals convex, subequal on disc; 3rd 2-punctate near middle and posteriorly (exact position of punctures variable as usual). Lower surface nearly impunctate. Secondary sexual characters normal except some 22 (2 of 4) with front tarsi with Ist seg- ment squamulose below, although not dilated. Length c. 16-20; width 5.5-6.4 mm. Holotype ¢ (M.C.Z. Type No. 30,390) and 17 paratypes (18 66, 4 22) all from Mt. Lewis, southwest of Mossman, North Queensland, c. 3000 ft., Dec. 1957, taken by the Darling- tons in mountain rain forest. The comparatively large, broad form, color, and characters given in the key should easily distinguish this species. It was found with dimorphicus on Mt. Lewis, and the occurrence of spurgeont too there or nearby raises an interesting problem of speciation. I think all 3 species have probably been derived from one ancestor, but I do not know how the divergence has come about. NOTONOMUS MONTORUM Nh. sp. With characters of doddi subgroup as given above; form as figured (Fig. 6), rather large, broad, and depressed in group; ereenish or sometimes vaguely purplish with elytral margins usually greenish and never bright blue; rather dull, entire upper surface with very fine, isodiametric reticulate microsculpture. Head c. % width prothorax, without noticeable unusual charac- ters. Prothorax 14 to % wider than long; sides broadly arcuate for much of length, usually broadly but shghtly sinuate pos- teriorly; basal angles usually subprominent, c. right, scarcely blunted ; surface of disc with weak transverse strigulae. Elytra c. 1% or more wider than prothorax; striae widened and opaque especially posteriorly ; intervals moderately convex, subequal on dise ; 3rd without dorsal punctures. Lower surface with sides of ventral segments (especially Ist) finely and irregularly punctate, but sterna nearly impunctate. Secondary sexual characters of 646 and most 22 normal, but exceptional 2 2 with Ist segment front tarsi with squamae below. Length c. 15.5-19.5; width 5.2- 6.9 mm. 10 BREVIORA No. 148 Holotype ¢ (M.C.Z. Type No. 30,391) and 76 paratypes from Mt. Bartle Frere, west slope, 3000-5000 ft., Dee. 1957, and 12 additional paratypes with the same data except 2000-3500 ft. altitude; all specimens taken by the Darlingtons, in mountain rain forest. Mt. Bartle Frere is at the eastern side of the Ather- ton Tableland south of Cairns, North Queensland. A weak sub- species (below) occurs on neighboring Mt. Bellenden Ker. See key for distinguishing characters of this species, and see also note under elliots (below). NOTONOMUS MONTORUM AZUL Nn. subsp. Structurally similar to typical montorum (above) but more shining and different in color, bluish black with elytral margins bright blue in all specimens. Of 5 22, 1 has and 4 have not squamae on Ist segment of front tarsi. Length c. 16-19; width 5.3-6.6 mm. Holotype ¢ (M.C.Z. Type No. 30,392) and 11 paratypes all from Mt. Bellenden Ker, east side, 3000-4500 ft., Jan. 1958, collected by myself in mountain rain forest. Mt. Bellenden Ker is about 10 miles north of Mt. Bartle Frere at the eastern edge of the Atherton Tableland south of Cairns, North Queensland. NOTONOMUS ELLIOTI nN. sp. With characters of doddi subgroup as given above; form as figured (Fig. 7), narrower than montorum, moderately de- pressed; purplish black, rather shining; fine reticulate micro- sculpture present but lightly impressed on head and pronotum, slightly more distinet on elytra. Head 7% or slightly less width prothorax, without noticeable unusual characters. Prothorax c. \. or shghtly less wider than long; sides broadly, rather weakly arcuate for most of length, then shgehtly sinuate near base; basal angles c. right, well defined ; dise faintly, transversely strigulose (as in most species of genus). Elytra c. YS or less wider than prothorax; striae narrow anteriorly but widening on posterior part of dise and especially on declivity; intervals moderately convex, subequal on dise (odd ones slightly wider than even) ; 3rd without dorsal punctures. Lower surface almost impunctate. Secondary sexual characters normal in ¢; 2 unknown. Length c. 16-18 ; width 5.0-6.0 mm. Holotype ¢ (M.C.Z. Type No. 30,393) and 5 paratypes (all 64) all from the Elhot Range (actually from near the summit 1961 AUSTRALIAN CARABID BEETLES Wk of ‘‘Sharp Elliot’’), c. 8000 ft., Mar. 1958, taken by my son and myself in mountain rain forest. This species may be related to montorum of Mts. Bartle Frere and Bellenden Ker. If so, the two mountain species are pre- sumably relics of a once more widely distributed stock. As com- pared with montorum, the present species is more slender, with elytral striae narrower, especially anteriorly. NoToNOMUS MASCULINUS Darlington N. masculinus (Fig. 8) is a relatively large, broad species with elytral intervals normal in number but heavily catenulate (much interrupted) and without dorsal elytral punctures. It has an extensive distribution on the southern part of the Atherton Table- land. I described it (1953, p. 99) from Millaa Millaa, and other known localities are indicated below. I have noted elsewhere (1961c, p. 7) the resemblance between this species and Pamborus punctatus, and have suggested that it may be a case of mimetic convergence. In ¢¢4 of masculinus the front tarsi are moderately dilated, with three segments squamulose below. In some 2 2 the front tarsi are normal (without squamae) but in others the front tarsi, though scarcely widened, have the 1st segment biseriately squam- ulose. My single 2 from Millaa Millaa has the front tarsi squamulose as described, and so does the single 2 from Herber- ton. A single 2 from between Millaa and Innisfail has the tarsi simple. In a series of specimens from mountains (including Mt. Fisher) between Millaa Millaa and Ravenshoe, 5 2 2 have tarsi simple, 3 squamulose. Three 22 from Longlands Gap all have simple tarsi. My single specimen from the western foot of Mt. Bartle Frere (the northeastern known limit of the species’ range ) isa 6, NoTONOMUS SAEPISTRIATUS Sloane <9 triplicatus Darlington 1953, p. 100 (new synonymy ). Sloane (1907, p. 364) described this species (Fig. 9) from Atherton, on the Atherton Tableland, North Queensland. I mis- interpreted Sloane’s description and re-described the species from Lake Barrine and Yungaburra, on the Tableland not far from the type locality. In 1957-1958 my wife, son, and I took a total of 8 more specimens of the species at Atherton, Yunga- burra, and Lake Eacham. At Atherton, we found it only in patches of rain forest on the flat part of the Tableland along No. 148 BREVIORA 12 1961 AUSTRALIAN CARABID BEETLES 13 Fig. 1. Notonomus transitus n. sp. Fig. 2. Notonomus montellus n. sp. Fig. 3. Notonomus dimorphicus n. sp. (slender ¢ ). Fig. 4. Notonomus dimorphicus n. sp. (@ ). Fig.5. Notonomus flos n. sp. Fig. 6. Notonomus montorum n. sp. Fig. 7. Notonomus ellioti n. sp. Fig. 8. Notonomus masculinus Darlington Fig. 9. Notonomus saepistriatus Sloane 14 BREVIORA No. 147 the road toward Mareeba. The species, therefore, seems to be uncommon, confined to a very limited area (a strip about 10 miles long) of the central Atherton Tableland. It apparently does not occur in the mountains west and south of Atherton, where it is replaced by doddi, and does not overlap the range of masculinus. In other words, these 3 strikingly different species seem to be allopatric. My 3 2 2 of saepistriatus all have narrow, unelothed front tarsi. REFERENCES DARLINGTON, P. J., JR. 1953. Australian earabid beetles II. Some new Pterostichini. Psyche, 60: 90-101. 196la. Australian earabid beetles IV. List of loealities, 1956-1958. Psyche, 67: 111-126. 1961b. Australian ecarabid beetles V. Transition of wet forest faunas from New Guinea to Tasmania. Psyche, 68: 1-24. 1961¢e. Australian carabid beetles VI. The tropical and some subtropical species of Pamborus, Mystropomus, and Nurus. Breviora, No. 142: 1-13. 1961d. Australian ecarabid beetles VII. Trichosternus, especially the tropical species. Psyche (In press). 196le. Australian carabid beetles VIII. Leiradira, especially the trop- ical species. Breviora, No. 147: 1-12. SLOANE, T. G. 1907. [Notonomus saepistriatus.] Proce. Linn. Soe. New South Wales, 32: 364-365. 1913. Revisional notes on Australian Carabidae. Part IV. The genus Notonomus. Proce. Linn. Soc. New South Wales, 38: 404-449. 7 i 3 i + : aan “ae # Fi i} 3 A oun) GF) Cpe Pegs \ & } \ ’ : i ; Nb t f i ae A * “= ? a Me ou Hi bok io at Pi I ie i er eee ee ey F | eff, 1 “ay TA) th pele bik 4 alte hin bond Ms ae vie h ady yo sacle 1° denc bigs: | *yF* — = or) . iyi pal oi hy = t ~ n HO « i xe’ r 7 j=e © iii BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. DECEMBER 19, 1961 NuMBER 149 A PRELIMINARY STUDY OF THE SILURIAN CERATIOCARIDIDS (CRUSTACEA: PHYLLOCARIDA) OF LESMAHAGOW, SCOTLAND By W. D. IAN ROLFE Museum of Comparative Zoology AND T. P. BURNABY University College of North Staffordshire, Keele, England The following ten species of Ceratiocaris have been recorded by Peach (1901, pp. 450-451) and Jones and Woodward (1888b, p. 72) from beds of uppermost Valentian age in the Lesmaha- gow inher, Lanarkshire, Scotland : C. inornata M’Coy, 1851 *C. angusta Etheridge, Woodward and Jones, 1886b *C. lara Etheridge, Woodward and Jones, 1886b C. longa Jones and Woodward, 1885 C. murchisoni (Agassiz), 1837 =O. papilio Salter, 1859 *C. stygia Salter, 1860 C. patula Etheridge, Woodward and Jones, 1888 C. robusta Salter, 1860 C. attenuata Etheridge, Woodward and Jones, 1886b [=C. tyrannus Salter in Etheridge and New- ton, 1878, nom. nud. | Species indicated by an asterisk are based on type specimens from Lesmahagow. It is unlikely that ten sympatric species of Cerati- vcaris would be ecologically compatible (Gause’s principle — see also Simpson, 1961, p. 74), and examination of the supposedly diagnostic characters of these species suggests that the majority 2, BREVIORA No. 149 are mere variants. Thus C. inornata is stated by Jones and Wood- ward (1885, p. 394; 1886a, p. 343; 1888b, p. 37) to ‘‘agree per- feetly in form and proportions [of the carapace] with C. papilio from Lesmahago, also in ornament, except that the postero-dorsal convergence of the striae is not present.’’ Yet elsewhere they state that the C. inornata from Lesmahagow (BM 59648) is ‘‘near to C. papilio in form’’ but that its ‘‘proportions are different from those of C. papilio’’ (1885, pp. 460-461; 1886a, p. 346; 1888b, p. 49). My own measurements (W.D.I.R.) on this speci- men differ from those given by Jones and Woodward, but even if their values are taken the proportions are found to be identical with the figured specimen of C. papilio provided by Jones and Woodward (1885, pl. 10, fig. 1; 1888b, pl. 12, fig. 1). The speci- men is too poorly preserved to distinguish whether the diagnostic striae convergence is present or not. C. lara is a juvenile instar of C. papilio, as Jones and Wood- ward first thought (1885, p. 396). C. murchisoni is used to de- note moderately large styles of several species, in this case of C. papilio. In Etheridge, Jones and Woodward’s own words ‘‘C. robusta, being based on some small caudal appendages without carapaces, is troublesome and unsatisfactory to deal with. We find some equivalent styles and .. . stylets in C. papilio, stygia, acuminata ete., but none of these seem small enough for the several little sets of trifid appendages, more or less perfect, which we have met with. C. robusta takes in some of these; but Oxford Mus. T is relatively broad, and might be termed lata [= patula 1888]; BM 58878 from Muirkirk has very nar- row members (angusta).’’ C. longa was first regarded as a variety of C. robusta (1885, p. 464) although some specimens ‘‘may well belong to C. papilio or C. stygia’’ (1886b, p. 458). This ‘variety’ was raised to species level later as the style was considered too long for either C. papilio or C. stygia (1888b, p. 43). C. attenuata differs in having ‘‘narrower and smaller’’ abdominal segments and style and stylets shorter than C. gigas or C. murchisom (1886b, p. 456-457). Style length is unsuitable as a character for specific differ- entiation since only when cameo and intaglio are available is it possible to know if the long, needle-like, distal portion of the style is preserved. Furthermore, the style is a hollow structure and appears ‘‘relatively broad’’ simply due to flat- tening during burial. It is therefore suggested that all the above mentioned ‘species’ belong to either C. papilio or C. 1961 PRELIMINARY STUDY OF CERATIOCARIDIDS 3 stygia, which form the subject of the present preliminary study. An account of the morphology of the species is given elsewhere (Rolfe, in press). Although Salter (1860, p. 156) stated that C. stygia had its ‘‘margine ventrali plus minusve angulato,’’ the diagnostic char- acter of C. papilio was ‘‘the much shorter body — searcely longer (tail included) than the great carapace — [which] easily distin- guishes it.’’ Jones and Woodward (1885, p. 392; 1886a, p. 341; 1888b, p. 36) misrepresented Salter’s diagnosis of these two spe- cies, thus ‘‘as mentioned by Salter, one (C. papilio) has the eara- pace more oblong than the other (C. stygia).’’ This revised diag- nosis led them to suggest (1885, p. 393; 1886a, pp. 341-342 ; 1888b, p. 36) that only the first of the three figures ‘‘termed C. papilio, evidently from oversight’’ by Salter (1860, p. 154) was in fact that species, the remaining two being C. stygia. Salter had al- ready suggested (1860, p. 156) that there were ‘‘at least two varieties of carapace in C. stygius itself.’? Subsequent workers have found it difficult to distinguish the two species, and it has been suggested that they are identical (Stormer, 1935, p. 294). This difficulty was also encountered in the present study and hence the museum collections of material available to Salter and Jones and Woodward (i.e. their hypodigm) have been re- studied to see if the species distinction could be maintained. One complicating factor in the use of the older collections is the separation of parts from counterparts. Specimens of iden- tical dimensions occur in different institutions, and without bringing all the material together it is impossible to be certain that part and counterpart of the same individual are not treated as two individuals. This does not affect the present study, how- ever, since it is arguable that such duplication will be distributed evenly throughout the sample. Two new collections have been examined to determine if, for example, C. papilio and C. stygia were allopatrie or even succes- sional species (= chrono- or palaeospecies). The first of these, the A. Ritchie collection in the Grant Institute of Geology, Edin- burgh University, came from the ‘Jamoytius Beds’, half a mile up the Logan Water from Logan House (Ritchie, 1960, p. 647). The other was collected by J. S. Jennings from a horizon ca. 700 feet higher in the suecession at the locality near Logan Reser- voir known as Shank’s Castle (Peach and Horne, 1899, p. 573). This collection is now deposited in the Geology Department, University College of North Staffordshire. 4 BREVIORA No. 149 The older material is housed in the several museums listed on Table 1. Few of the specimens have accurate localities indicated, but the majority probably came from the Shank’s Castle region. Several other localities are known, however, and these have been detailed by Etheridge (1873b, p. 49). The authors wish to thank the several curators for access to collections, and Professor H. B. Whittington for reading the manuscript of this paper. Salter’s criterion for distinguishing C. papilio from C. stygia by the number of segments protruding from the carapace is arti- ficial, since it depends solely on the degree to which the thorax and abdomen are impacted into, or drawn out from the carapace after death or exuviation. Jones and Woodward suggested that ““C. stygia was rather larger than C. papilio; its telson was larger; the carapace was markedly distinet by its trapezoidal outline, deep ventral region, and mucronate antero-dorsal angle, which was not nearly so often lost in fossilization as the front angle of C. papilio’’ (1885, p. 395; 1886a, p. 344; 1888b, p. 40). Style leneth has already been criticised as a specific character ; thus of 202 styles measured only 64 were complete and associated with the last abdominal segment. The ratio of style length/last abdominal segment length ranges from 1.5 to 2.9, and a seatter diagram of this ratio showed normal uncorrelated variation when plotted against the last abdominal segment length. This does not confirm Jones and Woodward’s assertion that ‘‘in C. stygia the style is usually rather more than twice, and in C. papilio only about twice as long as the ultimate segment’’ (1888), p. 39). The remaining two characters utilised by Jones and Wood- ward are carapace size and shape. We here define a ‘size factor’ P such that P = log (LH) and a ‘shape factor’ Q such that Q = log (L/H) where L and H are the overall length and height of the cara- pace in millimeters. Figure 1 shows the relationship between the size and shape factors for the 128 intact carapaces listed in Table I. The distribution of points is the same as would have been ob- tained by plotting L against H on double-log paper and rotat- ing the diagram through 45°. The object of performing the logarithmic transformation and rotation analytically is two-fold : to normalise the distribution of the shape factor Q, and to sim- plify investigation of the extent to which growth in Ceratiocaris is allometrie. 1961 PRELIMINARY STUDY OF CERATIOCARIDIDS 5 The overall mean value of the shape factor and standard de- viation were found to be Q = 0.2775 = 0.05917 corresponding to a geometric mean value of the ratio L/H of 1.894. The regression of Q on P was calculated by the first-mo- ment method of Wald and Bartlett. The observed P values were divided into five groups and the mean values of Q and P were found for each group, the number of points in each group being 20, 20, 48, 20, and 20. The five mean points are shown on Figure 1, (Q,-Q;). Their coordinates are (2.8109, 0.3097), (3.0779, 0.2787), (3.2100, 0.2717), (3.8350, 0.2698), and (3.4830, 0.2646). The regression is thus non-linear, although as may be seen from Figure 1, the four points @5-6- le on a practically straight line. The deviation of Q: from this line is scarcely significant, and the line itself does not depart significantly from a direction parallel to the P axis, if the first group of points is ignored. If we interpret the assemblage as a growth series, it follows that growth is isometric except in the range of size for which P is less than 3.07 (LH less than 1175 mm.?). In this latter size range, growth is not isometric, though the degree of allometry is small and is barely significant. Changes in the allometric constant during growth are well known in crustaceans, and commonly separate two distinct in- stars or mark a more critical eecdysis such as the prepuberty moult (Teissier, 1960; Simpson, Roe, and Lewontin, 1960, pp. 412-415). If we discard the 20 smallest specimens and test the marginal distribution of the remaining 108 Q values by plotting on prob- ability paper, we find that the distribution is an almost perfect unimodal normal curve (a straight line plot). There is thus no evidence to justify splitting up the assemblage on the basis of carapace shape. If, moreover, we regard the assemblage as a single homogeneous sample of a single species, we can readily calculate confidence limits for Q values, to test whether a given specimen may reasonably be regarded as a member of the as- semblage. (It will be as well to withhold judgment in the case of very small specimens, but otherwise there should be no need to worry about allometry. ) The first writer (W.D.I.R.) has been unable to trace either of the syntypes of C. papilio (Salter, 1859, p. 262) and hence speci- men GSM 7479, the original of Salter’s 1860, p. 154, fig. 1, is here treated as the neotype of the species. Jones and Wood- ward accepted this specimen as a genuine C. papilio (= M.P.G. x 45 in 1885, p. 393; 1886a, p. 342; 1888b, p. 36). Caleula- BREVIORA TABLE | Length (L) and height (H) of 128 museum specimens of ceratiocaridid Specimen number GSM-165-2 BM-24161 BM-24163 BM-59648 BM-16483 H=-no number Ke-09 123 fu Ke-09 123 cg Ke-Airdrie Coll. J-LW8 R2 Ke-09 123 dy Ke-09 123 cb Ke-R22 GSE -6650 Ke-09 123 cd 2 BM-45161 J-27 H-Stark Coll. BM-24150 AR 59-152 Ke-09 123 ed GSM-X 1/13 Ke-R29 J-LW8 Q2 GSM-256 AR 59-146 BM-24151 Ke-R23 BM-41895 BM-24160 H-A1901 Ke-09 123 ex BM-24158 AR 59-149 Ke-R8 Ke-Airdrie Coll. E-1902/30/11 J-LW8-L H-Macnair Coll. H-A1902 H-66 AR 59-108 BM-16495 E Specimen number Ke-09 123 gb BM -24164 H-A1908 GSM-7479 GSE -6647-1 BM-16501 K-5-3 Ke-R28 BM-24157 Ke-09 123 dn GSM-87338 AR 59-148 E-1902/30/9 K-6-1 GSE -6647-2 H-no number Ke-09 123 cn K-5-2 J-LW9 E Ke-R1 GSM-272 H-A1903 BM-no number AR 59-154 K-5-1 H-A1904-1 AR 59-Q BM-41894 Ke-R2 E-1902/30/7 H-A1908-3 BM-16517 Ke-09 123 dz GSM-no number H-A1908-2 H-T .Wlse Coll. GSM-X 1/22 Ke-09 123 ej Ke-R3 H-A1904-2 K-8 Ke-07 123 ey E-1902/30/4 E 53 H 26 Carapaces from the Lesmahagow area, in millimeters Specimen number AR 59-153 AR 59-144 AR 59-104 K-6-2 E-1902/30/10 E-1902/30/1 K-1 BM-16513 Ke-R33 AR 59-M K-7 AR 59-P BM-41894 J-LW8 B H-A1900-1 Ke-R4 BM-16482-] BM-16482-2 H-A1900-2 Ke-09 123 ez BM-24153 AR 59-147 BM-24155 Ke-R9 AR 59-150 AR 59-151 BM-58669 Ke-R25 BM-24149 E-1891/92/6 Ke-09 123 ea Ke-R24 BM-41898 K-3 BM-16479 GSM-X 1/21 E-1865/11/17 Ke-09 123 ce BM-45154 GSM-X 1/19 Ke-09 123 dr H-Macnalr Coll. 90 105 96 100 . 149 Ranked according to Increasing size factor P = log (LH); the five groups delineated by the heavy IInes are those referred to in the text. Repositorles: AR 59 - Ritchie Collection, Grant Institute of Geology, Edinburgh University; BM - British Museum (Natural History), London; E - Royal Scottlsh Museum, Edinburgh; GSE - H. M. Geological Survey, Edinburgh; GSM - H. M. Geological Survey Museum, London; H - Hunterlan Museum, Glasgow University; J - Jennings Collection, Geology Department, University College of North Staf- fordshire; K - Kilmarnock Public Museum; Ke - Kelvingrove Museum, Glasgow. | PRELIMINARY STUDY OF CERATIOCARIDIDS 1961 ‘ozIS puv odvys SULUYsp UL PEesN SUOTSUOUIIP YIM OATVA odedeied JYSLI JO oUuTpJNO SMOYS }YSIL waddn je ueiseiq “preMpoom pue sou0r Aq DJYULOUL *Q SB poytyuapt Usauroeds — gFZggG Wa ‘'090 ‘zc “By ‘OT ‘[d ‘Cggl ‘prvapooA, pue souor Aq poinsy ,,pIOhys ‘9,,— FSTSh WA ‘ (A10}SIV [RANQeN ‘uMesn YSIAg ur suout -1oads SULAMOT[OF puv sty) “ojo ‘T “sy ‘OT ‘Id ‘eggT ‘pavaMpoom pue souor Aq poinsy ovwndnd +g Jo wouttdeds , [eu -10uqe, — 6998¢ WA ‘(uopuoyT ‘unesnyy AdAING [BOISOTOay) 10};eg orpdpd *D Jo od£joou pojsods3ns — E/F, WSD :01B poT[oqul sjoq ‘sojsuvisy uryytm syutod Aq poyrvur S&H — 1% sonyea uwvow say oy} JO WOTZR[NOTed OY} UL pepnypoUt jou 4nq UostIVduI0d IOF poJLOSUI ‘AOT[UT ST[TFZ MBYSSe]T Sulmnoqgysrou oy} wory suowtoeds JYSIo o}voTPUT SossoIg “SUOISUOUIp [BoTJUNPL JO S[ENpLAIpul Jo JaquinU MOYS Sy} puv SuoutVods MOZLYVUSET QgT o}BOIPUT 8JOG ‘10}[BG oupndvd siupv01jv1aQ JO sodedeivod 1OJ S1OJOVVJ ozIS puev odeys usoajoqd Arysuoryepes oy} JO weIserp 10},Bdg “T “BLT (HVb9 =A HOLIOVS IZIS GE SSE ZE GE SE rE SE - es Kgs OFS AGc = Cice Vac, BIIce eSici % s Ke) m e O e ‘ Rs ab . , peas e x a es e Tan oye os e co 9 : Oe kee ais a emma ts re ie 5 rsispwe- * , s¥,- PaO "Lore Var ae See y od ae LY te ° e im (7 < J) e ” e ° =sie0° e-1, oe. . lo . ; ‘ 9) oer te ‘3 ° a = e " Q a SPr96ES WY > W F071 Q R¢ Rr \ Cals A so-s NX © 8 BREVIORA No. 149 tion shows that this specimen lies within the 95 percent confidence limits for group 3. The ‘primary’ types of C. stygia are unknown but one of the ‘‘good specimens of C. stygia’’ figured by Jones and Woodward (1885, pl. 10, fig. 2; 1888b, pl. 12, fig. 2) BM 45154, is similarly within the 95 percent confidence limits for group 5. C. stygia is thus not separable from C. papilio on statistical or indeed any other present evidence, and the former junior synonym should be suppressed. The morphospecies C. papilio and C. stygia form the one biospecies (or transient spe- cies) C. papilio. It is worth noting that the specimen of C. pap- ilio figured by Jones and Woodward (1885, pl. 10, fig. 1; 1888b, pl. 12, fig. 1) BM 58669 is ‘abnormal,’ as can be seen from its position on Figure 1. No significant differences in P or Q ean be detected in the stratigraphically separated material of the Ritchie and Jennings collections. There is a notable lack of clustering in terms of the size factor P corresponding to the mean size of successive moult stages or instars. Attempts to divide the Ritchie or Jennings specimens alone into instars were also unsuccessful. Intra-instar variation may obscure the limits of successive instars if sufficiently great, but not all crustaceans obey Brooks’ Law (Needham, 1950, pp. 10-11), and C. papilio may be another exception. At least part of this difficulty is due to sampling and preservation. Thus small individuals are only rarely collected (see Fig. 1) and all large individuals found have been incomplete. Relatively gigantic specimens up to two feet in total length occur both in the Les- mahagow and Haeshaw Hills inlers, but they can only be recon- structed from fragments and hence do not appear on Figure 1. A further complicating feature of the few large specimens avail- able is their distinctive dendritic carapace ornament. It can be ergued, however, that this ornament is characteristic of adult mstars (Rolfe, in press). Such giant individuals must have had younger growth stages coincident in size with the specimens of Figure 1, and no such dendritic ornament has been observed in that size range. It seems preferable to extend the name C. papilio to include these large individuals, at the risk of ‘lumping,’ until better sampling has been made. 1961 PRELIMINARY STUDY OF CERATIOCARIDIDS ) SUMMARY Of ten species of Ceratiocaris recorded from the Lesmahagow inlier, only C. papilio and C. stygia are sufficiently well founded to demand preliminary investigation. Both Salter’s diagnosis of these two species and Jones and Woodward’s subsequent de- finitions are artificial. Analysis of 128 carapaces in museum col- lections shows that C. papilio is indistinguishable from C. stygia, and the latter should be suppressed as a junior synonym. Carapaces of C. papilio show isometric growth except in the smallest individuals. The material cannot be resolved into a series of distinct instars. REFERENCES Only those references not listed by Van Straelen and Schmitz, 1934, are given here. NEEDHAM, A. E. 1950. Growth and regeneration rates in relation to age in the Crus- tacea with special reference to the isopod, Asellus aquaticus (Linn.). Jour. Gerontology, 5: 5-16, RivtcHig, A. 1960. A new interpretation of Jamoytius kerwoodi White. Nature, 188: 647-649. RourFr, W. D. I. (In press) Grosser morphology of the Scottish Silurian phyllocarid crus- tacean, Ceratiocaris papilio Salter. Jour. Paleont. SIMPSON, G. G. 1961. Principles of animal taxonomy. Columbia Univ. Press, New York. Srupeson, G. G., A. RoE AND R. C, LEWONTIN 1960. Quantitative Zoology. Revised ed. Harcourt, Brace and Co., New York. ST@GRMER, L. 1935. Dictyocaris, Salter, a large crustacean from the Upper Silurian and Downtonian. Norsk geol. tiddskr., 15: 267-298. TEISSIER, G. 1960. Relative growth. Jn The Physiology of Crustacea, Waterman, T. H. ed., vol. 1, pp. 537-560. Academic Press, New York and London. VAN STRAELEN, V., AND G. SCHMITZ 1934. Crustacea Phyllocarida (=Archaeostraca). Fossilium Catalogus; pars 64, Berlin. Bean ra spelt rah Chae ivie ed plore tne iniin: Or ae viele pti pert Mnivol alin 9611 eortsi tery anes | rh heal araaptlee det foe, abe is ogee, & vv 4 axTe ibe ities ponp tales ' tat AE iebyag iow ity webiig Aveall: Dien tae (| 1 ie seae ‘ rou) bei hivrdvriad ingens abe 9? ar duls pile, a sie es neem rit.) i ; 7 ; r } U baste ipa <= » A Ph spuuiaa a qi = } it) al ; oi¥. feel Wie ve ara ety || viet! | i i ry mii! iv (pi Gr. _ ' ie (as , i if F rd : j u nA Kero ie ’ TP a) al te a a sp Alst 2] he r wats ' edi poren i Th alah ais on T, ’ aa ' Lia is Cony ay i f ¢ Pe “kis, | j i° ai u 7 : i a) oe — i? ; ; is28e | i re? Mids Lie U mave : ; / a oe : = .4 J y ‘ ‘ca - | } Y ACER Ee Rafi ; ; 7 tree ay Bed PTA, eae } nie A y) COO AP Sotehht) siwiy, a $ a Mave ia ee ot ue a > a = oe uly b Weep lates Ole OV oP RT) tobe r sae 4 : iy pepe i >, One" b4 ‘ ‘o : 7 a jay 7 ‘ sOTQEE E La ; thé an a Te it é aig 17 bE) Cl alee i ph all jm! , be - 4 mihy 9g AP 01h Oty : - a ! i us pe 9 ip Crhine iis | of » BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. JANUARY 5, 1962 NuMBER 150 THE GENUS BETHYLUS IN NORTH AMERICA (HYMENOPTERA: BETHYLIDAE) By Howarp E. Evans Bethylid wasps are predominantly tropical and subtropical in distribution, with only a few species of diverse genera penetrat- ing temperate regions and virtually none entering arctic or sub- arctic regions. The sole exception to this statement, illogically, is the type genus of the family, Bethylus Latreille. This rather highly evolved genus is circumpolar in distribution. In North America, specimens have been taken close to the Arctic Circle, but none have been taken south of New York, Illinois, Colorado, and central California. In the Old World there are several species of northerly distribution and several others from the Mediterranean region. The genus is not known from the South- ern Hemisphere. In North America, four species have been described in the genus ; these are: castaneus Kieffer, amocnus Fouts, brachypterus Whittaker, and flavicornis Whittaker. A fifth species, decipiens Provancher, has recently been transferred to the genus by Krombein (1958, U.S. Dept. Agri., Monogr. no. 2, first suppl., p. 98). Examination of types in the U.S. National Museum re- veals that two additional species, Arysepyris californicus Brid- well and Perisemus oregonensis Ashmead, properly belong in the genus. Since the latter species is the type of the genus Digoniozus Kieffer (1905, In André, Spee. Hymen. Eur. Alger., v. 9, p. 245), this name can be added to the synonymy of Bethylus. One of these seven names can be removed from further con- sideration here. I have recently had an opportunity to study the type and only known specimen of Bethylus castaneus Kieffer (1907, Berlin. Ent. Zeitsehr., 51: 295). The wings of this speci- men are in poor condition, but enough remains to be sure that this species belongs not to Bethylus but to the related genus Goniozus (new combination). Be BREVIORA No. 150 Thus there are six specifie names available for the North American Bethylus, three of them newly assigned to the genus. The question naturally arises as to how much synonymy is in- volved and how many species, in fact, are there? The present paper is an attempt to answer that question. ANALYSIS OF THE PROBLEM Specimens of this genus are not common in collections, but by borrowing material from many sources I was able to obtain about 80 specimens. One’s first impression, on scanning this material, is the remarkable uniformity of the specimens in size, color, and structure. The only notable color differences are sex- ual: the males have yellow mandibles and wholly yellow an- tennae, while the females have dark mandibles and the antennae more or less infuseated apically. There are no noticeable differ- ences in the structure of the mandibles and elypeus, in the sculp- turing of the head or thorax, or in the male genitalia. There is, however, one character which varies strikingly, and that is wing length. The wings vary all the way from small pads scarcely larger than the tegulae to wings of normal size. This is not unusual in the genus, as several brachypterous species have been described and the European fuscicornis is known to exhibit much variation in wing length. In the case of the North American Bethylus, it was of interest to know whether wing length varied in a continuous spectrum or whether there were certain wing-length types which might represent different species. Following O. W. Richards (1939, Trans. R. Ent. Soc. London, 89: 185-344) in his revision of the British species, I first deter- mined the relative wing length of each specimen by dividing the length of the fore wing by the length of the hind tibia (which is much easier to measure accurately than total body length). I then plotted the number of individuals exhibiting a given relative wing length (Fig. 1). The males fell into two distinct groups. Those of the first group (Type A) might be termed subapterous, since the wings are exceedingly small, barely surpassing the anterior margin of the propodeum. Males of the second group (Type B) might be termed micropterous, since the wings are still very small, extending about to the beginning of the pro- podeal declivity. When one plots the females on this same scale he obtains a somewhat different picture (lower half of Fig. 1). The subapterous forms (Type A) tend to have shehtly longer wings, the micropterous forms (Type B) shghtly shorter wings, BETHYLUS NORTH AMERICAN L962 UL0jyOq 4B sopeuas ‘doz ye sopeur (| LET] Vlqry pury JO Yysuo] Aq poptatp [MAT] Sura o10g JO TYSueT) SYYSUO], SULA OATPL[IA WOATS SUTIGTYXo (9}VULpPAO) STENPTATpUL FO Stoq uM N aly AN / 662 68¢ GE 69E 69T 6LT 69T OST GL 6CT 62T 611 soT 66 68 6b 69° 65 6 62 62 -6@ -@e -L'e -9f “On 2s - 9 Si srk =81 =2i) =tn =01t 6: UWA GG; WA ”YZZ | Z ZA sadrFwyntmrmrnr tol LH /m a7 662 68S HLE 69¢ 69I 6LT 69T 6ST OFT GET 62T GLI 60T 66° 69 61° 67 65 br 6 -62 VC LE -98 Sic) msde Gun aomorwowoenMnes De Seats sejowe, $0 ‘ON se|OWw $0 ‘ON 4 BREVIORA No. 150 so that the two curves overlap slightly. Furthermore, there is a class of individuals with wings of moderate length, reaching about to the posterior margin of the first abdominal tergite, which might be termed brachypterous (Type C), as well as a few in- dividuals with wines of normal length (macropterous, Type D). The fact that wing length varies discontinuously suggests the possibility that several species may be involved, each exhibiting a different wing leneth. Presumably there would be four such species, with males of the two less common ones still to be dis- covered. However, in the absence of other characters one cannot rule out the possibility of polymorphism. I once again turned to Richards’ study of the British species, and discovered that the most useful character for separating the three forms occurring in Britain is the ratio between the distance separating the hind oeelli and the distance separating the hind ocelli from the occiput. I determined this ratio for all speci- mens available to me but obtained a unimodal curve, with the mean 1.7, the range of variation from 1.3 to 2.2 (close to the range for the European fuscicornis). Thus these measurements failed to support the possibility of more than one species. How- ever, in the course of making the measurements I found myself able to recognize ‘‘long-headed’’ and ‘‘short-headed’’ individ- uals. The difference was slight, but sufficient to induce me to measure the heads and determine the width/length ratio. In the case of the males I again obtained two separate curves (Fig. 2, top). For the females I obtained a bimodal curve (Fig. 2, bot- tom). It was at once apparent that all the subapterous individ- uals (Type A) were ‘‘long-headed”’ (left hand curves in Fig. 2), all the micropterous individuals (Type B) ‘‘short-headed’’ (right hand curves in Fig. 2). The brachypterous females (Type ©) were all ‘‘short-headed’’, while the fully winged females (Type D) were of both types. Here was a suggestion that two species might be involved, with both species being polymorphic for wing length in the female sex. Upon sorting the specimens into two lots represent- ing probable species, several other differences previously over- looked or discounted in importance were discovered. The most important of these involved the sculpture of the propodeum, the shape of the male subgenital plate (Fig. 3), and the wing vena- tion of the few available fully winged females. Thus I am now convineed that two polymorphic species are involved. The name amocnus Fouts is applicable to the ‘‘long-headed’’ species, while 1962 NORTH AMERICAN BETHYLUS 5 decipiens (Provancher) is the earlhest name for the ‘‘short- headed’’ species. The two species are widely sympatric east of the Rockies, but amoenus is not known to occur west of the Rockies. The characters separating the two species are summar- ized below, as are their synonymy and distribution. No. of Males / Z “LLL / 70) 9) 0081 G2, 85) 04 65 6 67 66 89 90 91 92 WH/LH 1Z 11 10 9 6 Y No. of if Y Females 6 y Y 5 Uy : | LL 3 wy GZ Y; 77 2 ZA le TI YY 1 WW Cit. ike) eee 61 Ze 85 64 85 66 87 68 89 90 91 92 wH/LH Fig. 2. Numbers of individuals (ordinate) exhibiting given relative head lengths (width of head [WH] divided by length of head [LH]), males at top, females at bottom. The only good series of decipiens from one locality is the series of 16 females and 4 males from Chilliwack, British Colum- bia, on which Whittaker based his descriptions of brachypterus and flavicormis. As discussed further below, I have studied or obtained the necessary information on this entire series. Twelve of the females are micropterous (Type B), three are brachyp- terous (Type C), and one is macropterous (Type D). This 12 :3:1 ratio, obtained in a series from one locality, is approached rather closely by the ratio for the species throughout its range, 6 BREVIORA No. 150 which is 41:9:4. Unfortunately, only one reared series of this species is available. That is a series of five females and three males in the U.S. National Museum reared from Vicia angusti- folia at the Lummi Indian Reservation, Washington. All indi- viduals in this series are micropterous. Unfortunately, no good series of amoenus is available; the longest series consists of three females and a male taken on different dates at Bar Harbor, Maine. All of these individuals are subapterous, and in fact only one fully winged individual of this species is known. This is a female taken by O. W. Richards on the window of an automobile at Buffalo, N. Y., 19 Sept. 1928. The ratio of subaptery: brachyptery : macroptery in the females of this species is 23:0:1. It is, of course, entirely possible that brachypterous individuals of amoenus may some day be discov- ered. It is also quite possible that polymorphism for wing length may occur in the male sex. At present only seven males of amoenus are known, only thirteen of decipiens. Clearly any hy- potheses on the genetics of polymorphism in these wasps will have to await the day when much more material has accumulated in museums. At present it appears that only the females are polymorphie for wing length and that the polymorphism arises from a very simple genetic mechanism. The Nearctic decipiens is undoubtedly closely related to the Palaearetic fuscicornis and may well be derived from it. Not only are the ocellar measurements similar, as noted earher, but the sculpturing of the propodeum is similar and the male sub- genital plate virtually identical. However, there is no doubt in my mind that they are specifically distinct. The antennae of fusci- corms are shorter and the scape is black at the base, yellowish apically (the scape is wholly yellowish-brown in both Nearctic species). Richards has found that the frequency distribution of relative wing length in fuscicormis is more or less trimodal or quadrimodal, but less distinctly so than in decipiens and with a much larger proportion of longer-winged individuals. Further- more, the males of fuscicornis are typically macropterous rather than micropterous as in decipiens. TAXONOMIC TREATMENT Key to North American Species of BEeTHYLUS Propodeum with a median polished ridge, remainder of dise con- trastingly alutaceous; head rather short (width/length ratio ~] 1962 NORTH AMERICAN BETHYLUS .85-.91 in female, .89-.92 in male) ; wings of micropterous in- dividuals reaching at least nearly to middle of propodeal disc (relative wing length .57-.91) ; fully winged individuals with radial vein curved upward sharply apically, vein arising from basal vein barely indicated; fore tibiae clear yellow; male subgenital plate strongly emarginate, but the side-pieces relatively broad and blunt (ig. 3, a) ..decipiens (Provancher ) Propodeum somewhat convex dorsally but without a median ridge which is set off from the remainder of the disc; head slightly longer (width/length ratio .79-.84 in female, .85-.86 in male); wings of most individuals extremely small, reaching barely beyond anterior margin of propodeum (relative wing length .26-.54) ; fully winged individuals with radial vein not curved upward sharply at apex, vein arising from basal vein nearly as long as transverse median vein; fore tibiae of female usually at least weakly suffused with brownish; male subgenital plate with a strong emargination, the side-pieces reduced to slender, acuminate processes (Fig. 3, b)...... amoenus Fouts a ) Fig. 3. Subgenital plates of (a) Bethylus decipiens and (b) Bethylus amoenus. BETHYLUS DECIPIENS (Provancher ) Gonatopus decipiens Provancher, 1887, Add. Corr. Faune Ent. Canada, Hymen., p. i179 [Type: 2, Cap Rouge, Quebee (Que. Proy. Mus., yellow label no. 1332) |.— Muesebeck and Walkley, Wal! U.S: Dept. Acri. Monogr. 2. p. 1038: Perisemus oregonensis Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 70 [Type: 2, Portland, Oregon (U. S. Nat. Mus. no. 40422)]. New synonymy. 8 BREVIORA No. 150 Digoniozus oregonensis Kieffer, 1905, Spec. Hymen. Eur. Alger., 9: 245 [Made type of new genus Digoniozus|.— Muesebeck and Walkley, 1951, U. S. Dept. Agri. Monogr. 2, p. 732. Arysepyris californicus Bridwell, 1919, Proc. Hawaiian Ent. Soc., 4: 34 [Type: °, Parkside, San Francisco Co., Calif. (U.S. Nat. Mus. no. 64124) |. New synonymy. Bethylus brachypterus Whittaker, 1929, Trans. R. Ent. Soc. Lon- don, 76: 385 {| Type: 2 (not ¢ as stated), Chilliwack, Br. Col. (British Museum) |. — Muesebeck and Walkley, 1951, U.S. Dept. Agri. Monogr. 2, p. 732. New synonymy. Bethylus flavicornis Whittaker, 1929, Trans. R. Ent. Soc. London, 76: 386 [Type: 36, Chilliwack, Br. Col. (British Museum) |. — Muesebeck and Walkley, 1951, U. S. Dept. Agri. Monogr. 2, p. 732. New synonymy. Glenosema californicus Muesebeck and Walkley, 1951, U.S. Dept. Agri. Monogr. 2, p. 727. Bethylus decipiens Krombein, 1958, U. S. Dept. Agri. Monogr. 2 Kirsh Suppl p: 98: Remarks on types. — Provancher’s decipiens was transferred to Bethylus by Krombein upon his examination of the type. Dr. Krombein has kindly placed his notes at my disposal, and they leave no doubt that Provanecher’s name apples to this species. The propodeum is alutaceous but with a median polished ridge, and the wings extend almost to the posterior slope of the propod- eum. The type is in good condition. The type of Ashmead’s oregonensis is also in good condition and is a fully winged female of this species. The type of Brid- well’s californicus is unfortunately in poor condition, the head, abdomen, and legs all being missing. However, the wings and propodeum are typical of the micropterous form of decipiens. A topotypie female in the collection of the California Academy of Sciences is very similar to the type and is in good condition. Whittaker’s two names require special discussion. The types and most of the paratypes are in the British Museum and I| have not seen them. However, Mr. G. E. J. Nixon has been good enough to examine these specimens and send me the eritical in- formation on them. I have studied one paratype of brachypterus in the collection of Cornell University as well as two of this species and one of flavicornis in the collection of Robert M. Fouts of Laredo, Texas. The characters Whittaker used for separating the two species are color characters which happen to be those which separate the sexes, and it happens that all the specimens 1962 NORTH AMERICAN BETHYLUS 9 of brachypterus are females and all of flavicornis are males — Whittaker’s statements to the contrary notwithstanding. The entire series is from Chilliwack, British Columbia; the type of brachypterus is a fully winged female, that of flavicornis a micropterous male. As indicated earlier, three of the paratypes of brachypterus are brachypterous, the remaining twelve microp- terous. Specumens exanuned. —40 22,10 6é. ALASKA: 1 2, Fair- banks, 25 June 1948 [USNM]; 1 2, ean 2 July 1958 (C. Lindroth) [CNC]; 1 2, Nenana, 17 June 1953 (R. I. Sailer) [USNM]; 1 2,2 64, Mile 1476, Alaska Highway (C. Lindroth) [CNC]. BRITISH COLUMBIA: 1 2, Mile 290, Alaska High- way, 19 June 1951 (W. Mason) [CNC]; 1 2, Smithers, 12 June 1958 (C. Lindroth) [CNC]; 1 2, Cranbrook, 12 May 1922 (C. Garrett) [CNC]; 1 2, Victoria, 28 Aug. 1923 (K. F. Auden) [CNC]; 3 22,1 ¢, Chilliwack, May-June, Sept. 1927 (O. Whit- taker) (CU,, Coll, Ke" M. Wouts|+ 2 o,aGahano, 2° Aue. 1929 [Coll. Fouts]; 1 2, Kaslo (A. N. Caudell) [USNM]; 2 22, Terrace [MCZ]. WASHINGTON: 1 &, Olympia [USNM]; 0 22,4 66, Red River Rd., Lummi Ind. Res., 1 Aug. 1944 (Vicia angustifolia, W. W. Baker) [USNM]. OREGON: 1 2, Portland [USNM]; 1 ?, Forest Grove, 1 Apr. 1919 (A. C. Bur- rill) [USNM]; 1 2, Ashland Loop, Siskiyou Mts., Jackson Co., 6 Aug. 1950 (Malkin & Thatcher) [CAS]. CALIFORNIA: 1 2°, Land’s End, San Francisco, 11 July 1922 (F. X. Williams) [CAS]; 1 2, Parkside, San Francisco Co., 8 Sept. 1910 (J. C. Bridwell) [USNM]. UTAH: 1 2, Logan [MCZ]. COLORADO: Zoo lad. Hort Collis, June, Sept. 1895 (Cy BE) Baker) [USNM]. IDAHO: 1 2, Coeur d’Alene (H. J. Rust) [USNM]. ALBERTA: 1 2, Edmonton, June 1917 [USNM]; 1 ¢, Elk- water Lake, 19 July 1956 (O. Peck) [CNC]. ONTARIO: 1 2, Sudbury, 1892 [CNC]. QUEBEC: 1 2, Anticosti Island, 9 Sept. [MCZ]. NEW BRUNSWICK: 1 2, Penobsquis, Dee. 1927 (C. A. Frost) [MCZ]. NOVA SCOTIA: 1 2, Portapique, 23 July 1929 (C. A. Frost) [MCZ]. MAINE: 4 2 2, Bar Harbor, July- Oct. (A. E. Brower) [USNM]. NEW YORK: 1 2, Grand Island, 11 Oct 1922 [USNM]; 1 2, North Fairhaven, 1 Sept. 1918 [CU]. 1 The following abbreviations have been employed for the museums involved : CAS, California Academy of Sciences, San Francisco; CNC, Canadian National Cc ollections, Ottawa: CU, Cornell University, Ithaca ; MCZ, Museum of Compara- tive Zoology, Cambridge ; USNM, U. S. N: ational Muse eum, Washington. 10 BREVIORA No. 150 Map showing distribution of North American Bethylus. Solid triangles: Bethylus amoenus, subapterous form; hollow triangle: macropterous form of amoenus. Solid circles: B. decipiens, micropterous form; half-solid circles: brachypterous form of decipiens; hollow circles: macropterous form of decipiens, BETHYLUS AMOENUS Fouts Bethylus amoenus Fouts, 1928, Proce. Ent. Soc. Wash., 30: 127 [Type: 2, Slaterville-Caroline, Tompkins Co., N.Y., 14 June 1904 (Cornell Univ. no. 934)].— Muesebeck and Walkley, 1951, U. S. Dept. Agri. Monogr. 2, p. 732. Remarks on types.— The type is in good condition. I have also studied a male allotype, bearing the same data, in the collec- tion of Robert M. Fouts. Specumens examined. — 24 22,7 66. NORTHWEST TER- RITORIES: 2 22, Norman Wells, 3-13 July 1949 (W. Mason) [CNC]. ALBERTA: 1 2, Aspen Beach, 23 Aug. 1944 (O. Peck) [CNC]; 1 2, Elkwater Lake, 19 July 1956 (O. Peck) [CNC]. SASKATCHEWAN: 1 ¢, White Fox, 10 July 1944 (O. Peck) [CNC]; 1 46, Holdfast, June 1946 (W. A. Nelson) [CNC]; 1 2, 1962 NORTH AMERICAN BETHYLUS 1! Assiniboia, June 1955 (J. R. Vockeroth) [CNC]; 1 ¢, Saska- toon, 15 Sept. 1924 (K. M. King) [CNC]. MINNESOTA: 1 2, Eaglesnest, 26 Aug. 1959 (W. V. Balduf) [USNM]. WISCON- SIN: 1 2, Cranmoor, 20 May 1910 (C. W. Hooker) [USNM]. ILLINOIS: 1 2, Palos Park, 17 March 1933 (Frison & Mohr) (Ill. Nat. Hist. Survey]. MICHIGAN: 1 2, Wexford Co., 4 July 1952 (R. R. Dreisbach) [Coll. Dreisbach]; 1 2, Presque Isle Co., 28 July 1952 (P. B. Kannowski) [Coll. Dreisbach] ;1 2, Midland Co., 20 June 1945 (R. R. Dreisbach) [Coll. Dreisbach]. ONTARIO: 1 2, Jordan, 25 Sept. 1916 (W. A. Ross) [CNC]; 1 2, Rondeau Park, Kent Co., 28 June 1936 (G. Steyskal) [Coll. Dreisbach] ; 1 2, Prince Edward Co., 10 July 1950 (J. F. Brim- ley) [CNC]; 1 2, Belleville, 2 Oct. 1956 (J. M. Smith) [CNC]. NEW YORK: 1 2, Buffalo, 19 Aug. 1928 (O. W. Richards) [MCZ]; 2 22, Ithaca, 28 May, 23 June (Babiy, Evans) [CU, MCZ];1 2,1 2, Slaterville-Caroline, 14 June 1904 [CU, Coll. R. M. Fouts]; 1 ¢, Caroline-Harford, Tompkins Co., 15 June 1904 [CU]; 1 4, Gannett Hill, 30 Aug. 1925, 2000 feet [CU]. MAINE 93° 2°. 1 6. Bar Harbor, July, Sept. Oct. (Av E: Brower) [USNM]; 1 2, Southwest Harbor, 6 Sept. 1922 [CU]. NOVA SCOTIA: 1 2, Portapique, 22 July 1929 (C. A. Frost) [MCZ]. BIOLOGY OF THE GENUS The only specimen of this genus which I have collected was taken walking over the ground in a small sand pit. Several speci- mens in collections are labeled as having been taken sweeping, one while ‘‘sweeping Carex,’’ another ‘‘while beating for ants.”’ Several specimens of both species were taken by A. E. Brower at Bar Harbor, Maine, on ‘‘Great Heath,’’ one of the female decipiens ‘‘on flowers of Ilex verticillata.’’ A female amoenus from Palos Park, [llinois, is labeled ‘‘in wet peat sample,’’ while a series of decipiens from the Lummi Indian Reservation, Washington, is labeled ‘‘Rd Vicia angustifolia.’’ Apparently these insects occur in a variety of situations. B. amoenus has been collected in every month from March to October, decipiens from April to October and also in December. Two specimens of amoenus bear host data. One is the female listed above from Cranmoor, Wisconsin, which is indicated as a probable parasite of Eudemis vacciniana. This name is now re- varded as a synonym of Rhopobata naevana (Hbn.), an oleu- threutid moth known as the black-headed fireworm. The other 12 BREVIORA No. 150 specimen is the female listed from Belleville, Ontario, which is labeled as a parasite of Brachypterolus pulicarius L. This is a nitidulid beetle introduced from Europe to the United States about 1918. If the latter record is correct, it is the only known instance of a Bethylus attacking a beetle. The European cephalotes Forster and fuscicornis (Jurine) attack various caterpillars, chiefly Microlepidoptera but occasionally Noctuidae. The fe- male wasps sting and malaxate their rather large prey, then drag it to a place of concealment, such as a hollow stem. Several eggs are laid on the prey and several larvae develop on a single host. Further details regarding the biology of these two species may be found in the papers of Richards (1932, Trans. Ent. Soe. So. England, 8: 35-40; 19389, Trans. R. Ent. Soe. London, 89: 185- 344). BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. JANUARY 12, 1962 NuMBER 151 A NEW PHYLLOCARID CRUSTACEAN FROM THE UPPER DEVONIAN OF OHIO By W. D. Jan ROLFE INTRODUCTION When curating the collections of non-trilobite arthropods in the Museum of Comparative Zoology, the writer recently found a fossil crustacean which had been sent to Professor P. E. Ray- mond for determination. The specimen was received from the Cleveland Museum of Natural History through Dr. D. H. Dunkle in January 1944; the late Professor Raymond published no description of the specimen and left no manuscript notes with it. Recent collecting in the same area by Mr. G. Lammers of the Cleveland Museum failed to find further specimens, although a fragment of a second specimen from a different locality was recently donated to the Museum of Comparative Zoology by Mr. R. Pritschan of Cleveland. This specimen will be referred to as the MCZ specimen to distinguish it from the original Cleveland Museum specimen. During the preparation of this description Mr. Lammers called the writer’s attention to the fact that H. K. Brooks of the Uni- versity of Florida had collected and studied the echinocaridids of this region. In correspondence, Mr. Brooks informed the writer that he had photographed this specimen some years ago, but was kind enough to allow the writer to submit this account for publication. The writer is indebted to Dr. G. A. Cooper, Professor A. La Roeque and Professor F. G. Stehli for searching through the collections at the U. S. National Museum, the Ohio State Univer- sity and Western Reserve University, Cleveland, for additional material, and to Mr. W. E. Scheele, Director of the Cleveland Museum of Natural History, for allowing the specimen to be retained for description. Professor H. B. Whittington kindly took the photograph for Plate 1 and offered helpful criticism of the manuscript. 2 BREVIORA No. 151 SYSTEMATIC DESCRIPTION Subclass MALACOSTRACA Latreille, 1806 Superorder PHYLLOCARIDA Packard, 1879 Order ARCHAHOSTRACA Claus, 1888 Suborder RHINOCARINA Clarke in Zittel-Eastman, 1900 Family OHIOCARIDIDAE fam. nov. Diagnosis. Carapace valves deep, with anterodorsal-medio- ventral fold and broad median dorsal plate. Rostral plate and number of thoracic and abdominal segments unknown. Remarks. The family Rhinocarididae comprises five genera which form a compact group characterised by elongate carapace valves and a narrow median dorsal plate. The present genus is so distinct from the previously described Rhinocarina as to war- rant the erection of a second family. Genus OHIOCARIS gen. nov. Type species. Ohiocaris wycoffi sp. nov. Diagnosis. As for the family. OHIOCARIS WYCOFFI Sp. Nov. Plate 1; Figure 1 Description. The Cleveland specimen is exposed with the dorsal surface of the carapace uppermost in one half of a concretion. It is preserved as a very thin film of golden brown ?cuticular material, but this has been destroyed over much of the specimen so that an internal mould is revealed. The concretion has been split apart so that the two carapace valves and median dorsal plate are separated by matrix from two complete abdominal seg- ments, a fragment of a third, and stylet fragments. The where- abouts of the counterpart of the concretion are unknown. As may be seen from Figure 1 and Plate 1, the carapace valves are deep; a well-defined fold, semicircular in cross section, runs from the anterodorsal region of each carapace valve, immedi- ately posterior of the strong earapace horn, and dies out ventrad of the centre of the valve. The ventral or free margin of each valve is bordered by a narrow reflexed rim except in the mid- ventral region, where it continues as a marginal ridge inside the ventral margin. 1962 NEW DEVONIAN CRUSTACEAN FROM OHIO 3 The broad median dorsal plate is separated anteriorly from the cephalic region of the carapace by a shallow transverse groove. The grooves laterally separating the plate from the main area of the carapace valves are narrower and deeper than the anterior groove, and are confluent posteriorly with the carapace mandibles anterior carapace horn : 4 perelfopods oblique fold Se BQ left valve BONS ae 2 of carapace marginal postero median ridge groove median dorsal plate ' ventra/ process of style head last abdominal segment (?=7th) stylet Figure 1. Outline drawing of Ohiocaris wycoffi gen. et sp. nov. showing structures visible on Plate 1. Cleveland Museum of Natural History 33241. x¢ Ill rim and a groove marking the posterior edge of the median dorsal plate. These grooves doubtless mark the position of marginal rims analogous in structure to those at the ventral edge of the carapace valves. The plate bears a faint posteromedian groove which extends anteriorly for 1.4 mm. The carapace valves and median dorsal plate are crazed by 4 BREVIORA No. 151 veinlets of a brown mineral ( ?collophane), whereas the abdomi- nal segments and the matrix are unaffected, indicating differen- tial chemical desiccation. The fragments of test preserved are smooth and free from ornament and the few wrinkles present are clearly secondary. The abdominal segments are inverted relative to the carapace and thus the style and stylets are exposed ventral side upper- most. The last segment (?7th) is 1.3 times the length of the preceding segment and has a concave posteroventral margin. After the photograph for Plate 1 was taken, the fragmentary style and stylets were broken from the matrix. The dorsal head of the style thus exposed was found to be of the echinocearidid type illustrated by Echinocaris sublevis Whitfield, 1880, figure 6 (=Hall and Clarke, 1888, pl. 29, fig. 13). Only the bases of style and stylets are preserved but the former is triangular im cross section and much shorter than the stylets. As in all the known archaeostracans, the head of the style embraces the proxi- mal portions of the stylets laterally and ventrally. A small denticle, 0.2 mm. long by 0.4 mm. broad, projects posterolaterally from the right lateral edge of the ventral style process or plat- form. The cuticle of the style head, stylets and abdominal segments lacks ornament. Only the inflated coxal parts of the mandibles are preserved, and excavation has failed to reveal the toothed gnathal lobes which were probably broken off at burial. The mandibles have been impressed through the anterodorsal region of the carapace valves, and the plane of section shows the left mandible to have had a wall thickness of 0.7 mm. At least four recurved ridges on the anteroventral region of the carapace fairly certainly mark the position of simple pereio- pods. As they are only seen as impressions through the thin cuti- cle of the carapace no detail of their structure can be discerned. The MCZ specimen is a fragment showing the median dorsal plate only. Dimensions, in millimeters Cleveland Museum of Natural History 33241 Maximum length of undistorted right carapace valve 34.0 Maximum height of undistorted right carapace valve, to right edge of groove bordering median dorsal plate 25.0 1962 NEW DEVONIAN CRUSTACEAN FROM OHIO 5 Length of median dorsal plate, along mid-line 21.0 Maximum width of median dorsal plate, at a point 7 mm. posterior from transverse groove 7.3 Length of penultimate abdominal segment 4.3 Length of last abdominal seement 5.0 Maximum dorsal width of style head 4.5 Length of style head to base of style 3:3 Width of style at base Tell Cross-sectional diameter of stylet 1.6 Museum of Comparative Zoology 6556 Maximum length of median dorsal plate ca. 21 Maximum width of median dorsal plate T.2 Holotype. Cleveland Museum of Natural History 33241. Col- lected by Dale Wycoff, 25th May, 1934, from the Chagrin Shale, Upper Devonian. Locality —shore of Lake Erie at mouth of Porter Creek, 12 miles west of Cleveland, Cuyahoga County Ohio. Other material. Museum of Comparative Zoology 6556. Col- lected by Raymond Pritschan and donated to the Museum via G. Lammers, July 31, 1961; found as float from Chagrin Shale. Locality — Painesville, 25 miles northeast of Cleveland, Lake County, Ohio (? Whitfield’s 1880, p. 37, Leroy locality). Remarks. The spread out carapace valves of the Cleveland specimen recall the condition in Dithyrocaris, which possibly lived with the valves in this attitude. The marginal rim fore- shadows the well-developed submarginal wall and doublure strue- ture of Dithyrocaris, and is similar to the condition in the ceratio- caridids Caryocaris curvilata (Gurley) and Callizoe bohemica Barrande. The anterodorsal-medioventral fold is more anteriorly situated than that of Pephricaris horripilata Clarke, whereas the ‘Schragrippe’’? and ‘‘Schnabelfurche’’ of Silesicaris nasuta Gurich (1929, p. 29) run closer to the ventral margin. As men- tioned above, the style resembles that in Echinocaris rather than that in any rhinocaridid, but the long smooth last abdominal segment is different from that of every species of that genus. The posteromedian groove on the median dorsal plate may prove to be of phylogenetic significance as a vestige of the non- rhinocaridid simple dorsal hinge, and homologous with the me- dian fold of other members of the Rhinoecarina. Ohiocaris shows the greatest development of the median dorsal plate and suggests a derivation from the Middle and early Upper Devonian rhino- earidids. Thus the plate width/carapace width ratio is 0.146 in ? 6 BREVIORA No..151 Ohiocaris, but only 0.096 in the specimen of Elymocaris siliqua figured by Beecher (1902, pl. 19, fig. 8; Yale Peabody Museum 22410). Hall and Clarke’s reconstruction of Mesothyra oceani (1888, pl. 32, fig. 1) is inaccurate in showing the hypothetical median dorsal plate too broad. Measurement of the specimen upon which this reconstruction was based (New York State Museum 4576) shows that the ratio is 0.073, not 0.150 as figured. Two other specimens of J. oceant (NYSM 4577, 4581) give com- parable ratios of 0.093 and 0.082, and in the Rhinocaris columbina figured by Clarke (1893, fig. 4, NYSM 4786) the ratio is only 0.074. The coneretion in which the Cleveland specimen occurs has the characteristic orange colour of oxidised Chagrin material described by Cushing, Leverett and Van Horn (1931, p. 34). Fossils do not seem to have been recorded previously from the Chagrin west of Cleveland (Cushing e¢ al., 1931, p. 35). The MCZ specimen forms part of a small collection comprising many of the same species of brachiopods and peleeypods as those listed by Cushing et al. (1931, p. 35), and in addition the crustaceans Echinocaris multinodosa Whitfield, EF. sublevis Whitf. and Palacopalaemon newberry: Whitf.. REFERENCES BEECHER, C. E. 1902. Revision of the Phyllocarida from the Chemung and Waverly groups of Pennsylvania. Quart. J. Geol. Soc. London, vol. 58, pp. 441-449. CLARKE, J. M. 1893. On the structure of the carapace in the Devonian crustacean Rhinocaris; and the relation of the genus to Mesothyra and the Phyllocarida. Amer. Nat., vol. 27, pp. 793-801. CUSHING, H. P., LEVERETT, F., and F. R. VAN Horn 1931. Geology and mineral resources of the Cleveland district, Ohio. Bull. U. 8. Geol. Surv., 818. GURICH, G. 1929. Silesicaris von Leipe und die Phyllokariden tiberhaupt. Mitt. Min.-Geol. Staatsinst. Hamburg, Heft 11, pp. 21-90. Haut, J. and J. M. CLARKE 1888. Trilobites and other Crustacea. Natural History of New York, Palaeontology, vol. 7. WHITFIELD, R. P. 1880. Notice of new forms of fossil crustaceans from the Upper Devon- ian rocks of Ohio, with descriptions of new genera and species. Amer. J. Sci., ser. 3, vol. 19, pp. 33-42. 1962 NEW DEVONIAN CRUSTACEAN FROM OHIO Ti Plate 1. Ohiocaris wycoffi gen. et sp. noy., Cleveland Museum of Natural History 33241, x 1.7. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. JANUARY 15, 1962 NUMBER 152 NEW AUSTRALIAN DACETINE ANTS OF THE GENERA MESOSTRUMA BROWN AND CODIOMYRMEX WHEELER (Hymenoptera — Formicidae) By Ropert W. TAYLOR Biological Laboratories, Harvard University The two new species described below are of considerable in- terest as members of the Australian ant fauna. The rare genus Mesostruma, to which Mesostruma browni n. sp. is added, in- cludes two previously described species which have recently been revised by W. L. Brown, Jr. (1952). Mesostruma is known only from eastern Australia and occupies an important phylogenetic position in the subtribe Epopostrumiti, being almost exactly in- termediate in character between the major Australian genera Epopostruma Forel and Colobostruma Wheeler. Codiomyrmex flagellatus n. sp. is the second member of its genus to be de- seribed from northern Queensland, and as such is the second representative of the important short-mandibulate stock of the subtribe Strumigeniti to be recorded from Australia. The de- ficiency of this element in the Australian fauna is of considerable zoogeographic interest. Its historical absence or scarcity on the continent has perhaps been important in allowing adaptive radia- tion of the short-mandibulate Epopostrumiti of the genus Colo- bostruma (Brown, 1952, 1959; Brown and Wilson, 1959). MeEsSOSTRUMA BROWNI new species (Figs. 1-5) Holotype worker. Synthetic aggregate length (TL)* 3.6; head length (HL) 0.78; head width (HW) 0.71; mandibular exten- sion (ML) 0.37; Weber’s length of alitrunk (WL) 0.84; cephalic 1 All measurements stated in the present paper are given in millimeters, with indices in units. The conventions followed in measurements are those established for the Dacetini by Brown (1953a, 1953b). An ocular scale with units of 0.0168 mm. was used for measuring, with correction to the nearest unit, the maximum error being estimated as +£0.01 mm. Pe BREVIORA No. 152 index (CI) 91; mandibulo-cephalic index (MI) 47. Correspond- ing precisely with the generic characteristics cited by Brown (1948, 1952) for head shape, mandibular and antennal structure, petiole and postpetiole form, and body sculpturing. Head shape as in Figure 1, similar to large WM. laevigata workers, but clypeus and anterior part of head somewhat more transverse. Eyes small, strongly convex as in M. turnert. Humeral angles rounded; dorsum of alitrunk in profile strongly and evenly convex. Pro- podeal lamellae as in Figure 2, more extensive than in either previously described species. Petiolar node as in Figure 2, similar to that of M. turneri, but more massive and less acute above in side view. =n ogo” > ge Mesostruma browni new species, Figs. 1 and 2, worker (Holotype). Fig. 1. Full-face view of head. Fig. 2. Alitrunk, nodes and base of gaster in side view. Figs. 3-5, male (Allotype). Fig. 3. Full-face view of head. Fig. 4. Alitrunk, nodes and base of gaster in side view. Fig. 5. Forewing. Pilosity and sculpture omitted from Figs. 2-4. Scale line: 1 mm, 1962 NEW AUSTRALIAN DACETINE ANTS 3 Seulpture of head, alitrunk and petiole consisting of large circular umbilicate foveae, more widely spaced than in the other described species, rarely separated by distances less than their maximum diameter; the surfaces between them smooth and strongly shining. Dorsum of alitrunk less densely sculptured than head, with a median longitudinal area almost devoid of foveae. Posterior parts of sides of alitrunk opaque, the foveae indistinct and mixed with coarse punctures. Foveae of petiolar node smaller than those of head and alitrunk; those of postpetiole indistinct, postpetiolar dorsum finely and irregularly sculptured and feebly shining. Dorsum of first gastric segment smooth and strongly shining, with no trace of longitudinal striae. Color rich golden brown; petiole, base and apex of gaster, legs, mandibles and antennae lighter. Type locality. Two miles east of Berry, New South Wales (B. B. Lowery). Worker variation. Thirty-two paranidotype workers, collected with the holotype, have the following dimensions: TL 3.2-3.9; HL 0.72-0.83 (mean 0.77) ; HW 0.66-0.74 (mean 0.71) ; ML 0.32-0.38 (mean 0.35) ; WL 0.77-0.92; CI 89-96; MI 42-49. No significant structural variation is indicated in the series. The workers are monomorphie with no bimodality in the frequency distributions of the dimensions listed, and no perceptible allometric differentia- tion between the head and mandibular dimensions, within the sample. The measurements and indices of ten paratype specimens from Riverview College, Sydney (B. B. Lowery), fall within the above ranges and have almost identical means, but an eleventh specimen of the same series is much smaller: TL 3.1; HL 0.69; HW 0.62; ML 0.34; WL 0.74; CI 90; MI 49. The HL of this individual is 2.80 standard deviation units smaller than the overall mean HL of all 45 specimens examined (76.7 = SD 2.75), its HW is 3.0 standard deviation units smaller than the overall mean HW (70.4 = SD 2.80). The specimen is thus of extremely small size when compared with workers from mature colonies, and is perhaps an old nanitie which had survived into the mature colony with which it was collected. A single paratype worker from Barrington Tops, New South Wales (T. E. Woodward), has the following dimensions: TL 3.2; HL 0.72; HW 0.66; ML 0.32; WL 0.77; CI 92; MI 44. Paratype queens. The first series of dimensions are those of an alate from Burns Bay, Sydney (B. B. Lowery) ; the second 4 BREVIORA No. 152 series those of a dealate from Pymble, New South Wales (C. Mer- eovich). TL 4.0, 4.3; HL 0.80, 0.86; HW 0.76, 0.82; ML 0.37, 0.40; WL 1.07, 1.16; CI 95, 95; MI 46, 46. Differing from the workers in the usual characters of full sexuality: larger size, presence of ocelli and wings, and unreduced structure of ali- trunk. Coloration as in worker, the ocellar area dark brown. Wings clear with pale yellow veins, venational pattern similar to male. The same diagnostic features as those of the worker serve to distinguish queens from those of M. turneri. Allotype male. TL 3.0; HL 0.54; HW including compound eyes 0.63; WL 0.88; forewing length ca. 2.2 mm. Head as in Figure 3. Compound eyes large, elliptical, strongly convex, their longest diameters about 0.28 mm. Mandibles slender, acute, probably not opposable. Antennae robust, thirteen segmented. Maxillary and labial palpi well developed; palpal formula ap- parently maxillary 5, labial 3, as in the worker (the mouthparts have not been dissected from the unique specimen). Body profile as in Figure 4. Mesonotum with well developed notauli, their posterior portion, forming the stem of the ‘‘Y,’’ indistinct. Propodeal lamellae smaller than in the female castes. Petiole sub-clavate, the node low, sloping gradually back from the anterior peduncle; antero-ventral tooth obsolete. Lateral edges of post-petiole lacking aliform appendages, but each with a distinet, low, obtuse, longitudinal carina. Gaster broad, some- what flattened basally ; the basal edges of its first segment feebly earinate longitudinally, as in many epopostrumite workers. Geni- talia exposed, enfolded by the parameres. The latter similar to those of Orectognathus (Brown, 1953b): broad in dorsal view, with convex lateral outlines and strongly concave inner faces, the apices rounded with their tips turned inwards and opposed mesally. Apex of sub-genital plate acute. Cerci short and stout. The penis valves and volsellae have not been dissected from the specimen. Forewing venation (Fig. 5) of the Solenopsis type, as in Orectognathus, the apical elements (Rsf 5, Mf 4 and Cu-A) feebly developed, and the radial cell open. Hindwing narrow, with a broad posterior fringe of microtrichiae, and four well developed subapical hamuli; venation as in Orectognathus. Head and most of alitrunk coarsely and closely punctate. Punetures of the pronotal dorsum and prescutellar area similar to the foveae of the workers, but almost contiguous. Sides of alitrunk with quite extensive shining areas, especially on the 1962 NEW AUSTRALIAN DACETINE ANTS 5 median parts of the larger sclerites. Petiole and postpetiole with coarse punctures, those on the latter shallow and irregular. First gastric tergite semi-opaque, with very irregular and _ shallow large, flat, piligerous punctures. Color blackish-brown ; antennae, mandibles, under-mouthparts, and legs yellowish-brown. Wings clear, their veins pale yellow. Described from a unique specimen collected with the holotype and its associated paranidotype worker series. Material examined. Northeastern New South Wales: 2 miles east of Berry (type locality) December 28, 1959, holotype and 32 paratype workers, allotype male (B. B. Lowery). Burns Bay, Sydney, February 2, 1959, ex leaf litter, a single alate queen (paratype) (B. B. Lowery). Riverview College, Burns Bay, Syd- ney, April 19, 1959, eleven paratype workers ( B. B. Lowery). Pymble, Sydney, March 18, 1956, a single dealate queen (para- type) (C. Mereovich). Barrington Tops, ex leaf mould (Berlese funnel sample), a single paratype worker (T. E. Woodward). The holotype, with paratypes, has been returned to Father Lowery for eventual deposition in the Commonwealth Scientific and Industrial Research Organization collection at Canberra; the allotype, with paratypes, is in the Museum of Comparative Zoology, Harvard University; the remaining paratypes are in the Queensland Museum. Biology. The following information regarding the biology and ecology of M. browni has been provided by Father Lowery. The type locality is about two miles inland from Seven Mile Beach, in low hill country behind scrubby alluvial coastal flats. The collection was made in a grassed clearing in a heavily tim- bered area, with Hucalyptus and turpentine growing in black non-sandy soil. The Riverview College locality overlooks Burns Bay, Lance River Cove, Sydney. The colony taken there was found nesting in damp yellow sand beneath a cover of moss and a little grass. The site was in a clearing in low scrub about 15 to 20 feet high, with Hucalyptus corymbosa, Grevillea, Lantana and Leptosper- mum. The soil near the nest contained a few small termite gal- leries and a large nest of the locally dominant ant Acropyga australis. The alate paratype queen was collected nearer the Burns Bay foreshore, wandering on leaf litter in warm sunshine, during the late afternoon. Father Lowery has collected three further colonies of MW. brownt within 200 meters of the type nest site. One of these 6 BREVIORA No. 152 colonies was nesting about four inches below the surface of coarse black sandy soil, under a small rock, but probably not in direct contact with it. A maze of termite galleries was located immediately beneath the rock, and permeated the surrounding soil. The new species is dedicated to Dr. W. L. Brown, Jr. of Cornell University, a leading authority in ant taxonomy who has worked particularly with the Dacetini and has devoted much study to the Australian ants in general. The addition of M. brownt to Mesostruma requires no change in the basic concept of the genus, as formulated by Brown (1952). Indeed Mesostruma retains its appearance as a compact and distinctive genus, with its species abundantly distinet from each other. Considering the characters of the worker and queen, M. browni seems to be most closely related to M. turnert Forel, which it resembles in the structure of the alitrunk, propodeal lamellae and petiole. It has a pecan ers narrower head, however (see Brown and Wilson 1959, fig. 7), and lacks the longitudinal striation of the basal gastric anki seen in turneri. The ab- sence of humeral denticles, and the general form of the alitrunk, propodeal lamellae and petiole distinguish M. brown from M. laevigata Brown. The new species differs from both the previ- ously described species in its less dense sculpturing and overall elossiness, and its more extensive propodeal lamellae and more massive petiolar node. The three known species of Mesostruma may be separated by the following key (based on the workers). 1. Humeri rounded; dorsum of alitrunk strongly and evenly convex in profile; eyes protruding and very convex to Humeri acutely subdentate; dorsum of alitrunk not so markedly convex in profile; eyes less convex and protruding only slightly. (Victorian mallee) _......M. laevigata Brown 2. Head broad, CI 98- 100; head cameale opaque, the foveae almost con- tiguous; gaster finely longitudinally striate over basal half or more of segment I (vicinity of Cairns, Queensland) ........ M. twrneri Forel Head narrower, CI 88-96; head capsule strongly shining, the foveae separated by smooth areas at least as wide as their maximum diameter ; basal segment of gaster smooth and strongly shining (northern New Sousa WAGE) 2 .ob25n0cn00008 ee ge M. brownt n. sp. In the discussion above I have not considered the enigmatic species ?Mesostruma monstrosa (Viehmeyer), 1925 (Brown, 1948). The unfortunate circumstances surrounding the original ~] 1962 NEW AUSTRALIAN DACETINE ANTS selection of this species, which was based on an apparently ab- normal specimen, have been discussed by Brown (1952). Dr. Brown now believes (personal communication) that Viehmeyer’s species was most likely based on a defective Hpopostruma speci- men. In any case monstrosa seems best ignored, pending the location and competent re-examination of the type. CODIOMYRMEX FLAGELLATUS new species (Figs. 6-9) Holotype worker. TL 1.9; HL 0.48; HW 0.32; scape length (SL) 0.23; ML 0.08; WL 0.47; CI 68; MI 19. General form much as in C. semicomptus Brown (1959) (Fig. 9), but smaller and more lightly constructed. Shape of head as in Figure 6. Dorsal surface of cranium convex, sloping towards occiput and elypeus as in C. semicomptus, the convexity less pronounced, however, and the occipital lobes more broadly rounded when viewed from the side (cf. Figs. 7, 9). Mandibles strongly convex, rising above the anterior clypeal border ; with five strong, acute, conical teeth, each slightly smaller than the one posterior to it. Basal lamella normally hidden at full mandibular closure, set at a slightly lower level than the teeth and oblique to them; its shape as in Figure 8 — roughly right-triangular, the posterior edge almost perpendicular to the mandibular axis, and the anterior edge diagonal, forming the hypotenuse. The anterior lamellar edge rises almost immediately from the base of the proximal mandibular tooth so that the diastema is very brief. Clypeus almost perfectly plane. Body profile somewhat as in C. senucomptus, with which it is compared in Figures 7 and 9. Alitrunk narrow, its maximum width 0.59 x the HW;; its dorsum almost perfectly plane, with- out sutures or a median longitudinal carinula. Alitrunkal dor- sum in side view evenly arched between the pronotum and the propodeal teeth. In dorsal view the sides of the pronotal disc are evenly rounded and those of the remaining alitrunkal dorsum almost parallel, with the distance between them, at the base of the propodeal teeth, slightly more than half the pronotal width. Dorsum of alitrunk margined with a fine carina, less distinct than in C. semicomptus, enclosing the pronotal dise anteriorly, and continuous with the upper edges of the propodeal spines posteriorly. The latter with their infradental lamellae similar to those of semicomptus. Propodeal spiracle minute, circular, its margin not appreciably expanded; it contrasts with that of C. 8 BREVIORA No. 152 semicomptus, which is larger and has a wide and very conspicu- ous rim-like margin (Fig. 9). Profile of petiolar node as in Figure 7, shorter than that of semicomptus, only about as long as high in side view; seen from above the node is slightly longer than broad with rounded sides and a truncate anterior border. A full complement of areolate spongiform appendages is de- veloped, distributed normally as in Figure 7. Gaster depressed ; basigastric costulae reduced to about five feeble lines on either side of segment one, the two groups of costulae separated by a wide median shining area; the costulae extend back about 149 the length of the segment. Mandibles shining, with a few scattered punctures. Head capsule with a close cover of large, flat, irregular, shallow pune- tures, which are almost effaced in a small region in the center of the frons. Antennae very finely punctate, their scrobes pune- tate-granulose. Alitrunk, both nodes, gaster, legs and anterior parts of frons and clypeus smooth and strongly shining. Petiolar pedunele coarsely granulate. | | Codiomyrmex flagellatus new species, Figs. 6-8, holotype worker. Fig. 6. Full-face view of head. Fig. 7. Alitrunk nodes and base of gaster in side view. Fig. 8. Mandible. Codiomyrmex semicomptus Brown, paratype worker. Fig. 9. Alitrunk, nodes and base of gaster in side view. Scale line: 0.16 mm., for Fig. 8; 0.25 mm., for Figs. 6, 7 and 9. 1962 NEW AUSTRALIAN DACETINE ANTS 9 Occiput with a number of very fine long (0.05-0.08 mm.) hairs with clavate tips, somewhat finer than in C. semicomptus. Hairs of elypeus shorter and more distinctly clavate. A very few simi- lar hairs are present on the anterior part of the pronotal dise and the dorsal surfaces of both nodes. The pilosity otherwise consists of exceedingly long (0.13-0.36 mm.), fine, tapering hairs, which are distributed symmetrically. On the head, five pairs in the following positions (see Fig. 6): on the occipital border, about halfway between its midpoint and its lateral extremity on either side; on either side and a little anterior to the median part of the frons; at the edge of the cephalic dorsum just anterior to its widest point; at the edge of the cephalic dorsum at about mid head length; on either side of the posterior extension of the cly- peus, above the antennal insertions. On the body, six pairs: two pronotal; one on each node; two at the base of the gaster — distributed as shown in Figure 7. On the forelegs: a single flagellum on the outer side of the tibia, near its apex. On the middle and hind legs: single hairs on the outer sides of the limbs near the bases of the tibiae and basitarsi. These long flagella curve upward and inward on the head capsule; those of the body are erect with their apices turned posteriorly; the anterior pronotal and postpetiolar pairs are more tangential, and inclined laterally. Type locality. Clump Point (near Mourilyan) Queensland, June 3, 1953 (T. E. Woodward), collected from a Berlese fun- nel sample of leaf mould. Worker paratype variation. Seventeen worker paratypes, col- lected with the holotype, show no significant variation in size or structure. The specimens have at some time been subjected to drying and fungal attack while in alcohol storage, with the result that some are fragmentary. A few fungal hyphae are still attached to several of the specimens, and a number have had the pubescence, particularly the elongate flagella, damaged dur- ing the consequent cleaning. Among the known Codiomyrmez, C. flagellatus is most closely related to C. semicomptus Brown, the only other known Aus- tralian species. The general resemblances between these forms are indicated in the accompanying figures. The two species may be readily separated by the characters discussed in the above comparative dsecription; the differences in size, pilosity, and propodeal spiracular structure are especially characteristic. The adaptive significane of the peculiarly sparse and elongate body pilosity of C. flagellatus is not understood, and deserves examination should live material become available for future study. 10 BREVIORA No. 152 The holotype, with paratypes, has been placed in the collection of the Queensland Museum. Paratypes are deposited in the Museum of Comparative Zoology, Harvard University, and the Commonwealth Scientific and Industrial Research Organization Collection, Canberra. I wish to acknowledge the generous assistance of Father B. B. Lowery of Sydney, and Dr. T. E. Woodward, of the University of Queensland, who collected the bulk of the material cited above and made it available to me for study. I wish also to thank Dr. E. O. Wilson, of Harvard University, for his assistance and advice during the preparation of this paper. REFERENCES Brown, W. L., JR. 1948. A preliminary generic revision of the higher Dacetini (Hymen- optera: Formicidae). Trans. Amer. Ent. Soc., 74:101-129. 1952. The dacetine ant genus Mesostrwma Brown. Trans. Roy. Soe. S. Aust., 75:9-13. 1953a. Revisionary studies in the ant tribe Dacetini. Amer. Midl. Nat., 50:1-137. 1953b. A revision of the dacetine ant genus Orectognathus. Mem. Queensland Mus., 13:84-104. 1959. Some new species of dacetine ants. Breviora Mus. Comp. Zool. Harvard, 108:1-11. Brown, W. L., JR. and E. O. WILSON 1959. The evolution of the dacetine ants. Quart. Rev. Biol., 34(4): 278-294. . i Pe ie ey lik ay "ne _ ©) Was tet ey 5 hse ibe-Q oe oe ii ; i ausines pride dame lj 7 es ¥ Abbe tbekeait rn EN Canscck Chwiain , a) s rah eters “hi ; 4 J j ad - ? o*\" Fi i] ¥ - ’ _ = " ad BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. Frspruary 15, 1962 NuMBER 153 Z — ANOLIS SCRIPTUS GARMAN 1887, AN EARLIER NAME FOR ANOLIS LEUCOPHAEUS GARMAN 1888 By A. STANLEY RAND Garman in 1887 described Anolis scriptus on the basis of five specimens in the Museum of Comparative Zoology, giving the type locality as ‘‘Silver and Lena Keys, Fla.’’ Barbour in 1914 re-examined Garman’s type series and decided that they were identical with Anolis cristatellus from Puerto Rico and the Vir- gin Islands and therefore placed A. scriptus in the synonymy of A. cristatellus. In the course of an examination of the Museum of Comparative Zoology anoles referred to A. cristatellus I had oceasion to study the type series of A. scriptus. I find that the series is mixed and none is cristatellus. One, a juvenile, is A. homolechis quadrocclifer of Cuba; the other four are conspecific with the form from the southeastern Bahamas described by Gar- man (1888) as Anolis leuwcophaeus, and apparently subspecifically identical with the form from the Turks and Caicos Islands now called albipalpebralis Barbour 1916. Clearly the name scriptus can no longer be kept as a synonym of Anolis cristatellus, but correction of its status raises certain problems. Since the series is mixed a lectotype must be selected to fix the name. The type series, three adult males and two juveniles, are all somewhat faded from their long period of preservation. One of the juveniles possesses the scale characters of Anolis homolechis and the color pattern, dark spots over the shoulders, is still suf- ficiently evident to identify it as Anolis homolechis quadrocelifer. This is the specimen labeled as coming from Lena Key, which thus would appear to be Cayos de la Lena, near Cabo San An- tonio, Cuba.’ I have arbitrarily excluded this juvenile from the 1 For further information on this form see Ruibal and Williams (1961). 2 BREVIORA No. 153 concept of A. scriptus and it therefore needs no further dis- cussion. The remaining four specimens seem to belong to a single species and I herewith designate M.C.Z. No. 65950 as the lectotype of Anolis scriptus Garman. The labels accompanying these specimens say ‘‘Silver Key Florida.’’ I, like Barbour, have been unable to locate a Silver Key anywhere in the West Indies. There is a Silver Bank near the islands from which the types must have come in the south- eastern Bahamas but it is completely submerged. These specimens are very like cristatellus as both Garman and Barbour agreed. Garman distinguished them from cristatellus on the basis of the greater size of the two paravertebral scale rows. Barbour (1914, p. 274) said, ‘‘I can not see, however, that these are at all enlarged; and there is no other character in which they vary from true A. cristatellus.’’ An examination of the type series helps to explain this contradiction. Two of the males have the two paravertebral scale rows enlarged more than is usual in cristatellus, but the third male has the paravertebral scale rows searcely enlarged at all and it is undoubtedly this specimen that Barbour examined. However, a close comparison shows certain other and more constant differences between the type series of scriptus and the many specimens of cristatellus examined. In scriptus the dorsal seales lateral to the paravertebral rows are larger than they are in specimens of cristatellus of similar size. In cristatellus also, the frontal ridges are higher and sharper and the frontal depres- sion correspondingly deeper than in the type series of scriptus. Finally, in cristatellus, there are only 1-3 scales behind the inter- parietal and these are abruptly larger than the very small dorsal scales. In scriptus there are many more rows of enlarged scales in this position and they grade more gradually into the dorsal seales. In all of these characters the type series of scriptus differ from cristatellus and agree with specimens of the species now called leucophaeus. So far as I ean find, the types of scriptus do not show any seale differences from leucophaeus, nor does leucophaeus show any additional differences from cristatellus. From this it appears that scriptus and leucophaeus are synon- ymous and scriptus as the older name must be substituted for leucophaeus. 1962 ANOLIS SCRIPTUS 3 The species ‘‘leucophaeus’’ is quite widely distributed in the southeastern Bahamas and has been divided into four subspecies. These races have been described primarily on the basis of color pattern, and they are all very similar in scalation. They are diagnosed in Table I. The types of scriptus lack the many dark spots characteristic of leucophaeus and the lectotype has a well-developed tail crest which is lacking in sularum. Thus the name scriptus definitely does not apply to the populations called leucophaeus and sularum. Distinguishing between albipalpebralis and mariguanae is more difficult. The diagnostic difference between them is the pres- ence of a broad dark lateral band in mariguanae. This is absent in the type series of scriptus but it is also absent in many of the adult males of mariguanae and best developed only in the juve- niles and females. Even the small ‘‘type’’ of scriptus lacks this band but this specimen is so faded that one cannot be positive that the band was never present. Many of the females of albipal- pebralis have dark middorsal blotches which are lacking in the small ‘‘type’’ of scriptus but, since they are absent in many albi- palpebralis, this is not conclusive. The male scriptus have a com- plex mottling along the sides in addition to a light narrow lateral line. The light lateral line is found in both albipalpebralis and mariguanae but the mottling in the types of scriptus is most like that found in albipalpebralis. Finally, the lectotype of scriptus has a dark line running posteriorly from the eye onto the neck. This marking is found in some of the males of albipalpebralis but in none of the mariguanae examined. So far as can be deter- mined there are no useful scale differences between mariguanae and albipalpebralis. From this it appears that the ‘‘type’’ series of scriptus, while not indisputably assignable to either of these races, is most like albipalpebralis and the lectotype most clearly so. For this reason it seems necessary to replace the name albi- palpebralis by the name scriptus. In accordance with this change the type locality of scriptus is restricted from ‘‘Silver and Lena Keys’’ to ‘‘Silver Key,’’ Turks and Caicos Islands. Further restriction seems pointless at this time. The correct names for the races of this species now stand as follows: Anolis scriptus scriptus Garman 1887 = Anolis albipalpebralis Barbour 1916 Anolis scriptus leucophaeus Garman 1888 Anolis scriptus mariguanae Cochran 1931 Anolis scriptus sularum Barbour and Shreve 1935 No. 153 BREVIORA jtlosqe two} ped aoddad pue 4[es Y}IM Yep GQ) vueld ISOM pues Avo SpooOMyYy wimnae[ns juosoi1d pueq ylvp peroaq £q DAOGB poa..lap.Log AyTJuonbeay OULL [B19] BT VEST MOLUEU BUBNSVAVIN OvVUBNSTLIBUL snjdiws syoup Jo sotoedsqns ot J, juosaad SUIPZ JOU SOUT, [V104B] }UESTL MO.LIVU SPURIST SOoIvD puB SAN, stperqodpedrqy[e = sn}dts jyuosaid sjyods yaep j[vus AuvU vVNSvUy of} UY] pue yBVotyH snovydoonoy IT S4eL 4So.10 Tre} eLBIN uio}yvd [BS.1o(, asuvy 1962 ANOLIS SCRIPTUS 5 REFERENCES CITED CocHRAN, D. 1931. New Bahaman reptiles. Jour. Washington Acad. Sci., 21: 39-40. BaRBouR, T. 1914. A contribution to the zoogeography of the West Indies, with especial reference to amphibians and reptiles. Mem. Mus. Comp. Zool., 44: 209-359. 1916. Additional notes on West Indian reptiles and amphibians. Proe. Biol. Soe. Washington, 29: 215-220. Barpour, T. AND B. SHREVE 1935. Concerning some Bahaman reptiles, with notes on the fauna. Proe. Boston Soe. Nat. Hist., 40: 347-366. GARMAN, S. 1887. On West Indian Iguanidae and West Indian Scincidae in the collection of the Museum of Comparative Zoology, Cambridge, Mass., U.S.A. Bull. Essex Inst., 19: 25-53. 1888. Reptiles and batrachians from the Caymans and Bahamas. Bull. Essex Inst., 20: 103-116. RvIBAL, R. AND E. E. WILLIAMS 1961. The taxonomy of the Anolis homolechis complex of Cuba. Bull. Mus. Comp. Zool., 125: 211-246, me Pat Seren 7 > ee © at le pa a > Mi % 7 = coe ee ry, raya Wh tt km ad “ea eli momatl yeh 51 mel ay | = Atty saelbnd a We th dae ere oath GA ors thd ante ak i ib ‘4 eR ant ciel” fallin jen inemtifaa al eee’: ile) | gene | 2 : Wty? th A aa y ° Aarts pattern Hag Cine aL OM wal PiU Ws iWbkh wD» | J i uy pei? ae depuiAal - oF Lol ad ; Gare 4 sae ee v6 acne aly an reins Oh etivis adit wives cuineeidh BeBy 7 nf et ; Ob tee ae Ree ee onciots dna f ie [ . B i? ~ Cnn a i CPoub Sieh GET? Gay bed ef thbim? Oval ty a i 7 = folds Hori eGivunS arhle- eQeae) Bo Gas, 40) % voiedide uv ' AOay ary cael, SVEOR ET Ae.) . Qaalé ait Meld Sewer? Crk eotierd? od! (anh) GPERFS! boo ating aT. PEL OT is eel ee ny 4 Aaa sures Hp! a 7 7, “ ra (ine ) a ni ) 5i8 j i hile iv vt i Bh y to Jf ad BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. APRIL 4, 1962 NuMBER 154 NOTES ON HISPANIOLAN HERPETOLOGY o.. THE NATURAL HISTORY OF THREE SYMPATRIC SPECIES OF ANOLIS By A. 8S. Ranp The lizard genus Anolis is abundant in species and individuals in the Greater Antilles. Quite a number of the common species oceur together over wide areas. There is thus a special oppor- tunity to study the ecological relations of sympatric species of a single genus under conditions genuinely favorable for field observation. Several papers have now been published which have begun to exploit this fortunate opportunity: Ruibal (1961) and Collette (1961) working on the anoles of Cuba; Oliver (1948) on those of Bimini in the Bahamas; Williams and Rand (1961) on the semilineatus group in Hispaniola. These and earlier workers — Grant (1940) for Jamaica; Stejneger (1904) and Schmidt (1928) for Puerto Rico and especially Mertens (1939) for His- paniola — have established that in each well studied case there are small but definite differences in microhabitat among the sympatric Anolis. In this paper I document the same point for the three com- monest species of Anolis in Hispaniola. I describe also a number of behavioral differences which seem to be associated with the ecological differences and attempt to assess the adaptive sig- nificance of these behavior patterns. This study was made during a two-month field trip to the Dominican Republic and supple- mented with observations on captive lizards kept in the lab- oratory in Cambridge. ACKNOWLEDGMENTS I wish to thank the government of the Dominican Republic that, through the Universidad de Santo Domingo, provided us y BREVIORA No. 154 with transportation and other assistance in that country; Dr. Kugenio de Jesus Marcano who helped us in the field and who sent me live specimens on our return. I wish also to thank Mr. Clayton Ray for inviting me to accompany him on this trip, and for his assistance in the field; Dr. E. E. Williams for helping to arrange the trip and for his advice and assistance throughout, and Dr. A. L. Rand for critically reading the manuscript. Also I wish to thank Sigma Xi for providing some of the funds that made this trip possible. MATERIAL AND METHODS The field observations during the summer of 1958 were made incidentally to the main project, a survey of Dominican caves for vertebrate fossils. Consequently, they include no lengthy ob- servations in any one locality but are a synthesis of data col- lected from widely separated places. Though I noticed no geo- graphical variation in behavior, obviously this is a possible source of error that runs throughout the present study. This is par- ticularly true as observations are lumped for different subspecies in two cases: Anolis chlorocyanus chlorocyanus and A. chloro- cyanus cyanostictus; and Anolis distichus ignigularis and A. distichus dominicensis. The main areas, where observations were made, were: Santo Domingo and vicinity, Santiago and the nearby Sierra Septen- trional, Padre Las Casas, Sabana de la Mar and the mountains south of there, and the Sierra de Neiba. These field observations are least complete on A. chlorocyanus and uneven on the other two species. For example, I found eggs only of cybotes, while I observed copulation only in distichus. The laboratory observations made at the Harvard Biological Laboratories involved only two species, A. cybotes and A. chloro- cyanus chlorocyanus. These animals, ten of each species, all adults and about half of them males, were sent to me by Pro- fessor Eugenio de Jesus Mareano. The lizards were released in a room 12 x 20 ft. and 9 ft. high. The room was a constant 80° F. and lighted by a south window. Eight small tree trunks 7 feet tall, mounted on bases so that they stood upright, an 8 foot rub- ber tree, a potted pine bush, and some other potted plants were arranged around the room. The lizards were fed meal worms on the floor and in dishes taped at various heights above the floor, on the tree trunks. Fruit flies were also released in the room. The whole room was watered daily. 1962 THREE HISPANIOLAN ANOLES 3 For a period of about three weeks, daily observations of vary- ing lengths were made. The lizards quickly came to ignore me as I sat quietly in one corner of the room and watched them with low power binoculars. This was obviously not a natural situation but the observations that could be checked against the field notes show a gratifyingly close agreement. This method seems to be a useful adjunct to field observations though obviously no substitute for it. All three of these species are small to medium sized lizards possessing the characteristic specializations of the genus: en- larged subdigital lamellae and a throat fan or dewlap that is best developed in the male. In each species the males grow to a larger size than do the females. Anolis distichus varies in dorsal coloration from green to gray to brown, frequently with mottling on the back. The dewlap is yellow in A. d. dominicensis, and red with a yellow border in A. d.ignigularis. The males measure 51 mm in snout-vent length (Cochran 1941). Anolis cybotes is dorsally gray or brown, sometimes reddish, frequently with two indistinct greenish lateral stripes. The dew- lap is whitish or yellowish in color. The males measure 67 mm in snout-vent length (Cochran 1941). Anolis chlorocyanus is usually bright green with the ability to change to brown. A. c. cyanostictus also has a rust-red spot in front of the shoulder and one on the head. A. c. chlorocyanus has a dewlap that is hght blue anteriorly and dark, almost black posteriorly. A. c. cyanostictus has the dewlap proximally cad- mium yellow, distally sky-blue. The males measure 71 mm. in snout-vent length (Cochran 1941). OBSERVATIONS The observations from both field and laboratory have been combined and arranged according to topics. Under each topic heading all three species are compared. Geographical Distribution. All three species are widespread in the Dominican Republic. Apparently they avoid only the high, wet pine forests of the Cordillera Central and are less common in the dry regions on the south coast near Asua and in the northwest near Puerto Plata. However, A. chlorocyanus is replaced by A. coelestinus in the Barahona Peninsula. Geo- graphical variation in each of these species has been described by Mertens (1939). 4 BREVIORA No. 154 Habitat. I found these three species at almost every locality I visited in the Dominican Republic. Mertens called distichus and cybotes eurytopic forms and chlorocyanus only slightly less so. He records both distichus and cybotes from the mangrove areas along the coast to the lower pine forests around Constanza at some 1200 meters altitude. The most significant environmental factor for all three species seems to be some sort of vertical perch, a tree trunk, fence post or cliff face, but A. chlorocyanus seems to avoid the most open areas. Anolis distichus lives primarily on isolated trees and fence posts and along the edges of woods and trails and in open woods. A. cybotes lives in these situations but also occurs in the deeply shaded interiors of densely wooded areas that distichus avoids. A. chlorocyanus certainly occurs in the edge situations that dis- tichus prefers and probably in the heavier woods as well. Though there are differences such as these in the extremes tolerated, these animals live in the same habitats over most of the Dominican Republic and part, at least, of Haiti. Mertens notes that in separated localities he observed individuals of all three species living in the same tree. My observations agree with this. Microhabitat. These three species, though living in the same habitat, have differences in their microhabitat preferences. A. distichus lives almost exclusively on exposed (i.e. not closely surrounded by vegetation) tree trunks, fence posts and similar structures, within 10 to 15 feet of the ground. It is seldom seen on the ground or in the smaller branches of bushes or trees. Though it must descend to the ground to reach isolated trees, it does not spend much time on the ground. A. distichus is often very common on the palisade fences in small villages. A. cybotes is also primarily an animal of exposed tree trunks and fence posts within 10 feet of the ground. However, it also frequents rocks and fallen logs and smaller individuals are frequently seen on the ground. It also avoids, during the day at least (see Sleeping), small twigs and foliage. These field ob- servations are confirmed by the laboratory data which show cybotes spending most of its waking time on perches less than five feet from the floor. A. chlorocyanus lives also on tree trunks and fences but unlike the other two, frequently ranges high in the trees and out among the smaller branches. I saw a female fall from the crown of a 30 foot palm tree. I suspect that this species is one that lives primarily up among the branches and ranges down the trunk 1962 THREE HISPANIOLAN ANOLES 5 rather than the reverse. However, individuals were seen also on fences and once I saw a dozen individuals on a pole frame- work nine feet high that with a few bits of palm thatch was all that was left of a shed in the middle of a treeless pasture. Like distichus, chlorocyanus must occasionally descend to the ground to reach isolated trees. In the laboratory, chlorocyanus spent most of its time high on the perches and ventured out among the leaves of the rubber plant much more than did the cybotes. Territoriality. As Mertens noted, all three species appear to be territorial. He says that for A. cybotes and chlorocyanus he found only a single adult male and one or more females of each species on any one tree. When he placed an additional male on one of the occupied trees it was immediately attacked. A. dis- tichus, he says, defends a smaller territory and the large trees may have several males, each with its own territory, as well as a number of females. These observations agree with mine, although it was only in areas where distichus were extremely common that I found more than one adult male distichus on the same tree, and then only on large trees. Certainly in the laboratory no male of either cybotes or chloro- cyanus would tolerate another male of the same species in his immediate vicinity for long, and even females were chased away if they approached too closely. Occasionally a male of one species would display to an individual of the other species. In addition to this horizontal effect a vertical stratification on individual trees or posts is evident in at least two of the species, distichus and cybotes. On the smaller trees that were occupied by two individuals of distichus, the lizards were usually of opposite sex and the male was usually closer to the ground than was the female. On one tree there were four lizards of this species: a large male near the base, a female above him, a slightly smaller female above her and, highest of all, a juvenile who was about 7 feet above the ground. During the hour that I watched this tree all of the lizards moved a number of times but this stratification remained the same for it was actively enforced. When one of the higher individuals moved in sight of and within 2 or 3 feet of a lower one, the lower animal immediately reacted to it. The male did this by bobbing his head and pumping his dewlap, the female by a short charge in the direction of the intruder. In every case the intruder retreated immediately, around the tree and up. 6 BREVIORA No. 154 In cybotes almost all of the individuals on the trees were large and most of them were males. The individuals that were seen on the ground were almost always juveniles or females; the females are much smaller than the males in this species. This spatial distribution seems to be a real phenomenon but I have no field evidence to support the hypothesis that it is the result of territorial defense. However, there are certain laboratory observations that support this contention. In the laboratory I saw no vertical stratification but I also saw no evidence that the females tended to spend more time on the ground than did the males. This may be because the perches were small and nu- merous enough so that each lizard could occupy its own, as they usually did. However, there was a definite stratification horizontally with the large males on the perches near the window and the smaller males and females on those farther away. When a smaller individual invaded a perch near the window, and this happened quite frequently, the resident male would display to it and the intruder would retreat. Thus, though there is no direct evidence that the distribution seen in the field is due to territorial defense, there is evidence that defense could be at least a contributing factor: the males taking the most de- sirable positions, the elevated perches, and chasing away any smaller individual that attempted to move in, with the result that the juveniles and females would spend most of their time on the ground. It is of interest to note that this vertical stratification in cybotes and distichus results in greater difference in size between the individuals that occur together. Adult cybotes are larger, and the adult males much larger, than adult distichus. This size difference could be reflected in the size of the prey items taken and so reduce the competition for food between the two species. The young of cybotes are of course no bigger than adult dis- tichus but these are the individuals that live primarily on the ground and so do not occur on the tree trunks with the distichus. The importance of this in reducing competition for food, if it acts in this way at all, is of course unknown. It would be inter- esting to have an analysis of the stomach contents of various sized individuals of each species. In chlorocyanus, I have no clear evidence for vertical strati- fication either in the field or in the laboratory. However, the fact that most of the individuals caught on fences and tree trunks were adult males suggests that the females and the juveniles may 1962 THREE HISPANIOLAN ANOLES Tf stay higher in the trees. This phenomenon may also give maxi- mal emphasis to the size difference between this species and the distichus with which it comes in contact for it is the large indi- viduals of chlorocyanus that descend the tree trunks and they meet first and presumably would compete most for food with the smallest distichus. Posture. Seen in silhouette these three species are usually im- mediately recognizable. This is partly because of their different proportions but even more because of their very different postures. A. distichus rests facing either up or down or angling across the tree with its head and at least the anterior part of its body well off the substrate and with its neck bent so that its head is parallel to the substrate but further away from it than are its shoulders (Cf. figure 4, plate 2, Mertens). A. cybotes typically rests facing down the tree with the fore part of the body off the substrate and the neck bent dorsally so that the head is nearly parallel to the ground. In one individual seen resting on the underside of a log which slanted at about 45°, the neck was bent back well over 90°. This posture is true in the laboratory as well as in the field. On the ground the posture is much like that of distichus with the neck bent so that the head is parallel to the ground and raised above it. A. chlorocyanus usually rests with both its head and body quite close to the substrate and its neck bent only a little if at all, both in the field and in the laboratory. While these postures are typical of the normal resting posi- tion, both cybotes and distichus when mildly alarmed flatten against the substrate, and chlorocyanus, when about to display, raises liself up on its legs. I believe these differences in posture can be correlated with feeding behavior as discussed below. Mertens says that most Anolis rest with their heads pointing toward the ground. I[ noticed this most commonly in cybotes and less so in the other species. Activity. A. distichus appears to be a much more restless lizard than either of the other two species. Like them it spends most of its time resting quietly but an individual seldom remains in one place for more than a few minutes. It then moves quickly a few inches away on its tree trunk and rests quietly again for another few minutes. A. cybotes, on the other hand, seems to spend much longer periods of time resting in one spot. Again, when a change is 8 BREVIORA No. 154 made, it is made quickly. In the laboratory when a cybotes moved from one tree to another, the lizard frequently ran down to the base of the tree, stopped for a few moments, left the tree with a jump and ran part way across the floor, paused, ran the rest of the way and with a jump started up the new tree, usually pausing again before settling down. A. chlorocyanus, though also spending periods of time im- mobile, moved quite frequently, going slowly and deliberately about in the trees, in the laboratory as well as in the field. In the laboratory, moving from one tree to another was a single process. The lizard moved slowly down to the base of the tree, jumped off, dashed across the floor and with a jump started up the new tree. As little time as possible was spent on the floor. Climbing. I have no observations on the climbing ability of distichus. In the laboratory A. cybotes did not appear to be as sure a climber as A. chlorocyanus. I saw several of the former fall to the ground from the smooth leaves of the rubber plant while I noted that only one of the latter did so even though chlorocyanus spent much more time on these leaves than did cybotes. While both these species started their climbs in the lab with a jump up on to the vertical surface, and I saw a large male cybotes make a 6 or 7 inch vertical jump to reach a hanging branch, chloro- cyanus made a great many more horizontal jumps. Particularly common was one used to cross the 10 inches that separated the two closest perches. A. cybotes is apparently a quickly moving lizard, quite at home on the ground but not so much so in the more treacherous footing of smooth green leaves and small twigs. A. chlorocyanus, on the other hand, is shy of the ground, moving across it only occa- sionally and then with as much speed as it can manage, while in the less secure footing of more arboreal situations, its de- liberate movements help keep it from falling. Feeding. On several occasions a distichus was seen to move from its resting position on a tree trunk or fence post and snap up something small from the bark. Twice I saw a male interrupt his displaying to do this. Once I saw a male move up to an ant about an inch away, follow it up the tree a couple of inches and then apparently lose interest and turn away. Thus there seems to be some selection of food and not everything small that moves within range is eaten. 1962 THREE HISPANIOLAN ANOLES 9 A. cybotes was never seen to catch anything in the field, though on two occasions I saw an individual struggling with a large dragonfly ; each time the lizard had the head and thorax in his mouth and the wings and abdomen still protruding. In the lab- oratory, cybotes came willingly to the floor to take meal worms. Sometimes a lizard would return to a tree to eat the meal worm but usually it remained for several additional minutes before returning either to its old tree or a new one. I never saw a cybotes moving around on the floor looking for food. On one occasion a male left his perch and ran about 10 feet across the floor to make an unsuccessful attempt to capture a 2-inch cock- roach. I have no data on chlorocyanus feeding in the field. In the laboratory several individuals were seen snapping at small ob- jects on the leaves and twigs. This species only rarely came down to the floor to capture meal worms and each time that one did so it moved slowly to the base of the tree, then rushed out, seized the meal worm, ran back and climbed up the tree before stopping to eat its captured prey. On the basis of this limited evidence, some tentative generali- zations about the relations of feeding, posture and movement can be made. A. distichus seems to feed primarily on smaller insects that it catches on the tree trunk. The posture with the head held high above the substrate would enable it to see more ef the tree trunk than it could if the head was held low. The frequent move- ments are necessary if the lizard is to take advantage of the in- sects that happen to be on the opposite side of the tree trunk. The head is held parallel to the surface in which the lizard is most interested. A. cybotes males get at least part of their food from the ground after sighting it from their elevated perch. The posture of this species keeps the head roughly parallel to the surface in which the lizard is most interested, in this case the ground. Since the lizard can see a large sector of the ground around him at all times, he need not change his position frequently to maintain a careful scrutiny of a considerable area. A. chlorocyanus almost certainly gets most of its food from the trees in which it usually lives and its slow movements sug- vest that at least part of the time, unlike the other two species, it goes looking for it instead of lying in wait. The head is held parallel to the surface in which it is most interested. 10 BREVIORA No. 154 We need more observations on all of these points and par- ticularly on the feeding of A. chlorocyanus. But it is interesting that the two species, distichus and cybotes, that occur in the same microhabitat, on the tree trunks, seem to differ so markedly in feeding behavior. Mertens remarks, in passing, that most Anolis get their food on the ground and among the roots of the trees and that their usual posture, as he records it, oriented toward the ground, may be related to this. I believe that this is true of cybotes but not of the other species. Escape. Each of the arboreal species has a noticeably differ- ent method for evading herpetologists and presumably other predators. A. distichus, when approached, quickly moves around to the other side of its tree trunk, usually moving up or down at the same time. If followed, the lizard may continue this maneuver- ing to keep the tree trunk between itself and the pursuer for some minutes. Soon, however, the lizard will either run up the tree out of reach or run down it to the base where it is concealed by the surrounding vegetation. Only when very hard pressed will one leave his tree or post and run out into the grass. Occasionally, and this was noted especially early in the morning, distichus would hide under a bit of bark or in a hole in the tree. Mertens says that distichus, when approached, runs around to the other side of its tree or post and then down to its base to hide. He does not mention any of the other behavior described here. A. cybotes showed the same tactics in the field and in the lab- oratory. When approached, an individual would remain still until I came very close, then suddenly it would dart around to the other side of the tree; it might stop there but only until I moved into view again, then, instead of employing evasive ac- tion on the tree, it would usually run down and, unlike distichus, frequently leave the tree completely. In the laboratory the lizard usually ran a few feet away, where it might remain on the floor for some moments or might immediately climb a new tree. Mer- tens records these same flight reactions for this species. An A. cybotes, discovered on the ground or on a fallen log, frequently hid under whatever cover was available. Sometimes a large male, when first approached, would display his dewlap to me before fleeing. This was observed occasionally in the wild and became very common with one male in the laboratory after he became accustomed to me. 1962 THREE HISPANIOLAN ANOLES Wal Usually in the field, chlorocyanus, when approached, imme- diately began to climb up the tree. This action was slower and more deliberate than that of the other two lizards but, since it was started sooner, the lizard was usually carried safely out of my reach. In the laboratory and in the field, when on a fence, where the lizard could not climb out of reach, it usually climbed as high as it could get and then dodged about there. Only very occasionally did one run down to the ground, though frequently one jumped to a nearby perch, if available, or ran out among the foliage and twigs at the ends of the branches. Mertens also notes that chlorocyanus usually climbs up its tree and conceals itself in the crown. The escape behavior of the three species closely parallels the feeding behavior. A. distichus conducts its evasive behavior on the tree trunk. A. cybotes willingly leaves its perch to escape on the ground, and A. chlorocyanus retreats whenever possible up into the top of the tree. Daily Activity. Since I did not spend long periods of time watching any one lizard or groups of lizards, I have no detailed information on the variations in activity during the day. All three species seem to be strictly diurnal. They were seen sitting in the sun more frequently in the early morning than at other times of the day, presumably to raise the body tempera- ture to the preferred level. A. distichus, at least, seems to feed more actively in the morning than at any other time. On several occasions I captured one with food in its mouth and three times I had one snap at the knot of the thread noose with which I was trying to snare it. Sleeping. I do not know where A. distichus spends the night but, since individuals sometimes use holes in trees as hiding places in the early mornings, they may possibly use these during the night. Two individuals of A. cybotes were found asleep at night, in the field. Both of them were males and were asleep on top of the foliage of the outermost twigs of small bushes. They were plainly visible from my vantage point, though perhaps not to a predator, such as a snake, climbing the bush. In the laboratory one evening during an examination of the room, I located seven cybotes asleep. Five of them were in the needles at the tips of the branches of the pine bush, one on a small vine where it stretched away from the tree trunk, and the last on top of the topmost leaf of a large-leaved potted plant. Shaking a branch on 12 BREVIORA No. 154 which a cybotes was sleeping woke the lizard but it did not move until the branch was shaken vigorously or the lizard touched. Then it jumped immediately to the floor and remained there un- moving. A. chlorocyanus was seen asleep only in the laboratory where during the period mentioned above I located six animals. Three of them were sleeping between large leaves of the rubber plant and the wall, one was between a board and the wall, one inside a cold radiator, and only one exposed, on top of its perch. When disturbed, the chlorocyanus immediately sought new hiding places. As Mertens notes, A. distichus and A. cybotes sleep with their hind legs partly flexed, while chlorocyanus usually sleeps with them extended along the tail. I do not know enough about the real and potential nocturnal predators of Anolis to speculate on the adaptive significance of these sleeping places. But it is striking that they are so different from the situations in which the animals spend the day. Male cybotes, which in the daytime live on substantial vertical surfaces, during the night sleep on the small flexible foliage and twigs of bushes and vines. A. chlorocyanus during the day is usually exposed to view and during the night sleeps under or behind some sort of cover. Reproduction. Since copulation was observed only in distichus and eggs found only of cybotes, it is not possible to compare these species with respect to reproductive behavior. I am including these data in the hope that eventually comparative data will be available. Three times pairs of distichus were seen in copulation in early to mid-afternoon on tree trunks from four to six feet above the eround. In no case was the preceding courtship observed. In one case the lizards were oriented diagonally up a large tree six feet from the ground. The larger male was on the higher side and on top of the smaller female. One of his front legs was across her shoulders and holding on to the tree in front of her front leg; one of his hind legs was across the base of her tail, the toes resting on her thigh and the trunk in front of it. His other legs were spread out holding the tree. His tail was bent under hers and his head was resting on but not biting her neck. Her position was that of any resting lizard except that her tail was strongly arched. The position observed in the other cases was virtually identical. 1962 THREE HISPANIOLAN ANOLES 13 SUMMARY AND CONCLUSIONS These observations on the behavior and ecology of these three species of lizards, Anolis distichus, A. cybotes and A. chloro- cyanus in the Dominican Republic are obviously incomplete. However, certain tentative conclusions can be advanced. These species are sympatric and occupy the same macrohabi- tats over much of their ranges. The species differ somewhat in their microhabitats but they are not clearly separated spatially in this way; members of each species occur on the same tree within feet and sometimes within inches of each other. They all have the same basic body form, though differing somewhat in size, in proportions and in morphological adaptations such as development of the enlarged subdigital pads. They are all in- sectivorous in diet. However, in the details of their behavior they are very different. These details can be fitted together to form a picture of three morphologically similar species of the same genus living together in the same habitat but living three very different lives. Reviewing these very briefly, Anolis distichus is strictly an animal of the lower tree trunks. Its territorial pattern results in spacing out the individuals living on the same tree with the largest at the bottom and the smallest higher up. This species feeds on small insects on the trunk, utilizing a posture and pat- ern of activity that enables it to forage effectively in this sort of place. When frightened it takes evasive action on the tree trunk, and copulation takes place there. A. cybotes, though partially an animal of the tree trunks, is also closely associated with the ground below. Its territorial behavior results in a spatial distribution with the large indi- viduals on the trees and the smaller ones on the ground. Even for the individuals on the trees the perches there seem to be pri- marily lookouts from which to survey the ground. The postures and patterns of activity seem fitted best for this and much less so for watching the tree on which the lizards sit. Certainly they do go to the ground to capture food spotted from these perches. They run to the ground when frightened and they bury their ego's in the ground. A. chlorocyanus in its behavior seems as closely related to the tree tops as cybotes is to the ground. It seems to be mainly the large males that descend the tree trunks. Some feeding un- doubtedly occurs high in the trees even in these large males and 14 BREVIORA No. 154 chlorocyanus retreats upwards when frightened. Its slow de- liberate movements seem adapted to the more precarious arboreal footing among the leaves and twigs. These are not just three similar animals doing the same things in the same way in slightly different places but three similar animals, each with a unique set of complex behavior patterns which interlock functionally so that each species has its own way of life within the same habitat. This situation can be an example of the operation of the Gause- Volterra hypothesis that closely related species can live together only if they differ in ecology. Since many of these differences can be correlated with feeding behavior they can be interpreted further as serving to reduce interspecific competition for food. However, the demonstration that these conclusions are valid must await the collection of further data, particularly data on just what environmental factors act to limit the population densi- ties of each species. LITERATURE CITED CocHRAN, D. M. 1941. The herpetology of Hispaniola. Bull. U. 8. Nat. Mus., 177: 1-398. COLLETTE, B. 1961. Correlations between ecology and morphology in anoline lizards from Havana, Cuba and southern Florida. Bull. Mus. Comp. Zool., 125: 137-162. GRANT, C. 1940. II. The Reptiles. Jn Lynn, W. G. and C. Grant, The Herpetology of Jamaica. Bull. Inst. Jamaica, Sci. Ser., 1: 61-148. , & 1959. Temperature responses and ecological relationships of two Bor- nean lizards. Ecology, 40 (1): 127-136. MERTENS, R. 1939. Herpetologishe Ergebnisse einer Reise nach der Insel Hispaniola, Westindien. Abh. Senckenb. naturf. Ges., 449: 1-84. OLIVER, J. A. 1948. The anoline lizards of Bimini. Amer. Mus. Novitates, no. 1383: 1-36. RvuIBAL, R. 1961. Thermal relations of five species of tropical lizards. Evolution, 15: 98-111. Scumip?T, K. P. 1928. Amphibians and land reptiles of Porto Rico, with a list of those reported from the Virgin Islands. Sci. Survey Porto Rico and the Virgin Islands, 10, part 1: 1-535. 1962 THREE HISPANIOLAN ANOLES 15) STEJNEGER, L. 1904. The herpetology of Porto Rico. 553-724. WILLIAMS, E. E. and A. S. RAanpD 1961. Notes on Hispaniolan herpetology. 2. A review of the Anolis semilineatus group with the description of Anolis cochranae, new species. Breviora, no, 135: 1-11. Rep. U. S. Nat. Mus., 1902: ae Coe hee Dalen Fea: - 3 aan t ae) aya i. one a cele ~~ hd i admis - Saea ne ears’ ef i iteg en areer yee pare a et va i ia (en) Silva a ne Poimaprs dle _ LS 7 di sang yi a? a) =e ened is iets 7 Ai > e rt WE ee he ; Ww oh - anal ; 7 #4 pha! oe miteid%: \ ‘i fe fer es q 12” bee and Fei Hh a ni’ 1s a Ap we 4 ee an an | yn WA! gay See ee ee va ci we ih 4 te di ay j : ie ee Tin LO 1" mY 1 ia ss : j l@ ¢ we ei) os Kh ! 7 as OF : 1) "4 a @ = 7 a ey : 7 : oe _ ty? re +e re t aha we a : i : ‘va : ~*~ - 7 ly F fy ; a af | Ve BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. APRIL 12, 1962 NUMBER 155 NOTES ON HISPANIOLAN HERPETOLOGY 6. THE GIANT ANOLES By Ernest E. WIuiamMs INTRODUCTION Mertens (1939) has called attention to the existence of geo- graphic variation in the giant anoles of Hispaniola and has distinguished a typical western race, Anolis ricordii ricordii Duméril and Bibron, and an eastern race, A. r. baleatus Cope. The distinction between these two forms is sharp and un- equivoeal; the situation is, however, more complicated than Mertens’ limited sample (16 specimens) led him to believe. Study of the unreported series of Hispaniolan giant anoles in the American Museum of Natural History (AMNH) plus the specimens in the Museum of Comparative Zoology (MCZ) and the United States National Museum (USNM) (91 specimens in all) makes it clear that at least three vicariant forms are recog- nizable. The form occurring from Port-au-Prince north to Cap Haitien and Port-de-Paix is the one to which Mertens has shown that the name ricordi Dumeril and Bibron must be attached. Another, occurring in the north and east of the Dominican Republic, may be called, following Mertens, by Cope’s name baleatus (Eupristis baleatus Cope 1864, Proce. Acad. Nat. Sci. Phila., p. 168, type locality ‘‘St. Domingo’’). A third unnamed population occurs on the Barahona peninsula. A fourth popu- lation inhabiting the southwest peninsula of Haiti may be dis- tinct. I list below the distinguishing characters of the three well-marked forms. bo ricordiu Very low nuchal and dorsal crests Nuchal crest scales as long as or longer than high, and not or but little higher than the very low dorsal crest Head scales small, numerous (7-9 across snout at level of second canthal 2) In ¢ 6 deep black spots above shoulder, sometimes also on occi- BREVIORA TABLE i Barahona population Low nuchal and dorsal crests Nuchal crest seales higher than long but small, not higher than the weakly developed dorsal erest Head seales larger, fewer (4-6 across snout at level of second eanthal ) Whole of body in both sexes with very irregu- lar small blotches and No. 155 baleatus 1 A prominent nuchal crest, a variable but lower dorsal crest Nuchal erest scales much higher than long, always much higher than seales of dorsal erest Head seales large, few (2-5 aeross snout at level of second canthal) Both sexes with no evident pattern or transverse banding or put; no other evident mottling reticulation pattern CHARACTER ANALYSIS The scales on the snout are swollen, bosslike in all Hispaniolan giant anoles. The differences are solely in the size of these boss- like scales. The contrast is extreme in this regard between the giant anoles of northern Haiti (ricordii) and those of the north- ern Dominican Republic (baleatus). The animals of the Bara- hona and southwestern peninsulas, however, are intermediate, overlapping in this regard the north Dominican (baleatus) and approaching the north Haitian populations. On this character alone it is not possible to separate every specimen of the north Dominican and the southern populations though there is a well- marked average difference. Number of scales across the snout is a measure of a more general feature — general scale size — which is somewhat greater in baleatus than in ricordu. Mertens has cited a number of key regions, i.e. loreal region, base of the tail, ete., which show this. 1A. Salle, who collected the type of Lupristis baleatus worked both in the northern Dominican Republic and at the base of the Barahona peninsula (map of travels compiled by W. J. Clench). It was therefore necessary to confirm the application of the name by checking the characters of the British Museum type. Miss A. G. C. Grandison has courteously confirmed that the nuchal crest scales of the type are indeed significantly higher than the dorsal crest scales. 21 count the canthals forward from the anterior border of the orbit (see Fig. 1). The reverse count—from the anteriormost canthal back —is sometimes used, e.g. Oliver (1948). This, however, has the disadvantage that the scales here are small and variable and do not provide a stable starting point. 1962 HISPANIOLAN GIANT ANOLES 3 The snout, however, provides the clearest expression of this general feature and one which can be reduced to a simple count with an adequate numerical range (2-9). Mertens has used a similar transverse count but makes it directly in front of the eyes. I have chosen a transverse count somewhat further for- ward because this transverse line, anterior to the supraorbital semicircles seems to me more suitable as a place for a standard count utilizable for all species of Anolis. BALEATUS RICORDI! Fig. 1. The two extremes of anterior head squamation in Anolis ricordii subspecies. The arrows indicate the place at which the count across the snout is to be made. The pattern of geographic variation of nuchal crest scales is not so simple. Again, the populations of Port-au-Prince and northern Haiti and northern Dominican Republic show the extremes, the crests of Port-au-Prince and of north Haiti popu- lations very low with a long anteroposterior base, while those of north Dominican Republic populations are tapering, high, short based anteroposteriorly. The southern populations could again be described as intermediate but on careful examination this is not quite accurate. Both the north Haitian and southern populations have a reduced nuchal crest as compared with the north Dominican population but the nature of the reduction is different. The nuchal crest scales of north Haitian animals are long-based, low, rounded scales, not at all tapering; they are 4 BREVIORA Now0'55 sometimes shorter but usually slightly higher than the very low, long dorsal crest scales. In the giant anoles of the Bara- hona peninsula the nuchal scales are, in contrast, still short anteroposteriorly, relatively high, tapered, but very small, not appreciably higher than the tallest dorsal crest seales and no- tably smaller in area than the largest dorsal crest scales. The few specimens from the southwestern peninsula do not permit adequate analysis of the populations of this important geographic area. The four adults examined all come from the vicinity of Fond des Negres halfway alone the peninsula. In squamation, these are very like the animals of Port-au-Prince and north Haiti. The one remaining specimen is a juvenile (78 mm snout-vent length) from the foothills of the Massif de la Hotte. Although it is less than half-grown, its dorsal and nuchal scales are appreciably higher and less broad based than those of the adults from Fond des Negres and indeed are very comparable to those of the Barahona population; in addition, in this individual these scales are double and even triple even on the nape. It thus seems very possible that the populations of the middle of the southwest peninsula and those of its extreme end differ significantly. (See further discussion below.) The nuchal crest scales seem in all populations to be some- what variable. However, within each adequately-sampled popu- lation the range of variation is very characteristic and does not blur the distinctions tabulated above. Despite statements by Boulenger (1885) and by Mertens (1939), I do not find a clear correlation with sex; females, for example, of baleatus do not consistently have lower nuchal crest scales than male baleatus of the same size. There is, however, clear ontogenetic change ; the smallest individual of baleatus at hand (AMNH 28651 from San Juan Bay, Samana, 41 mm snout-vent length) has the nuchal erest scales only incipiently enlarged and could not be recog- nized as a member of the baleatus population on that character. However, in baleatus the characteristic, tapering, spinelike nu- chal scales develop very early and even specimens little more than half grown (e.g. MCZ 57719, Santiago, ¢, 83 mm snout- vent length; MCZ 5445, Samana Peninsula, ¢, 88 mm snout- vent length) are very readily recognizable. Dorsal crest scales are more variable than nuchal crest scales, and variable in a peculiar fashion: there is sometimes a regular alternation of relatively high triangular single scales and pairs 1962 HISPANIOLAN GIANT ANOLES 5 of much lower, more quadrangular scales.1. However, such a regular alternation occurs —if present at all— on only a por- tion of the back and there is present between nuchal crest and dorsal crest, on the one hand, and dorsal and tail crests, on the other, various irregular conditions with double and single crests erratically intermingled and scale types somewhat inter- mediate and rather irregularly so. In ricordu itself with a very low dorsal crest the double condition with all low scales is most frequent; in baleatus the situation is individually very variable ; in the Barahona and the southwest peninsulas the double condi- tion of the dorsal crest predominates. The body color of live specimens and well-preserved alcoholics is probably useful. The usual specimen, however, requires much interpretation and its evidence must be received with some skepticism. In tabulating pattern, above, I have ignored all ill-defined discoloratious and have tried to assess pattern on the basis of real aggregations of pigment rather than fortuitous darkening of random areas by formalin. Even with this qual- ification the problem is not simple: these are Anolis and have the power of color change; an adequate discussion of their pat- tern would be possible only if the whole repertoire of color changes were known. I record my information below by popu- lation. Ricordii: Females from Port-au-Prince and north Haiti ap- pear to be plain green above with no markings of any sort. Males from Port-au-Prince are also mainly green (as preserved they may show an obscure, very fine brown reticulation) but are distinguished by a large, very characteristic patch of black above the shoulder but of ill-defined shape and varying extent. There may also be irregular black patches extending onto the back of the head. There may also be a white patch at the corner of the mouth. A Cap Haitien male (Senckenberg 10445) is quite similar to Port-au-Prince specimens. Of three males newly re- ceived from Ti Guinin a little to the east of Cap Haitien (MCZ 66147-9) two have almost no black at the shoulder (only small and inconspicuous spots) and none at all on the head. The other male is as devoid of black spotting as any female. All show fine brown reticulation quite like that of similarly preserved Port- au-Prince males. 1In the tabulation above I have compared the nuchal crest scales with the highest crest scales at midbody. 6 BREVIORA No. 155 Baleatus: Schmidt (1921, p. 10) records specimens from Sanchez and Villa Rivas as ‘‘usually green with dark-edged transverse bands of light greenish yellow.’’ Preserved specimens which have any definable pattern show this transverse banding which is present on dorsum, limbs and tail; it appears to be most persistent on the tail. It is very conspicuous in a specimen near hatchling size (AMNH 28651). There are frequently num- bers of small spots on the sides of the belly and on the venter. The MCZ Santiago series, though preserved in a light phase, show very irregular ight banding much invaded by dark retie- ulations, but the bands are very prominent on limbs and tail. Barahona Population: | have unfortunately been unable to find any description of Barahona giant anoles in Hassler’s notes. I must therefore rely entirely on the preserved specimens, which, however, are unusually consistent, though from several localities and with very different collection dates. Collected by Hassler, they differ strikingly from the baleatus specimens, most of which were also collected by Hassler. All Barahona specimens, and most clearly those from Valle de Polo, show irregular small blotches, very variable in tint and extent, scattered over the entire dorsum. The closest approach to this condition is seen in those baleatus which show reticulation. There are never in the Barahona specimens the large shoulder spots of ricordii, sensu stricto. One specimen (AMNH 51241 from Enriquillo) does show transverse banding, but more obscurely than in baleatus. The small blotches are also very obscure in this specimen, which therefore could not in this phase be recognized as a member of the Barahona populaticn on its color. The nuchal and dorsal crest scales are, however, typical barahonae, and the hypothesis which I have adopted is that the blotching so characteristic of most preserved specimens is characteristic only of one of the phases of the color repertoire possible to this population. Population of the Southwest Peninsula of Haitc: Again the material is inadequate for proper analysis, the more evidently so since the few available specimens suggest that this is not a unit sample. The juvenile from the Massif de la Hotte is, as preserved, reddish or purplish brown with indistinct broken dark longitudinal lines on head and nape and more evident narrow brown transverse bands on the back, six between shoulder and groin, in pairs with very slightly lighter reddish-brown be- tween each pair. The tail is obscurely annulate. 1962 HISPANIOLAN GIANT ANOLES 1 In contrast, the three adult females from the vicinity of Fond des Negres are essentially patternless (MCZ 66016 is dark and obscurely vermiculate, and USNM 72631, 72633 are plain light green except for a subocular half ring of scales that is conspicu- ously white). The single male is similarly nearly patternless but has also the subocular half ring of white and, in addition, a series of small black spots above the shoulder, very like a vestige of the large black shoulder spot of Port-au-Prince ani- mals and very lke those of the two Ti Guinin males. TAXONOMIC EVALUATION There appear to be at least three distinctive populations of giant anoles, all readily separable by nuchal crest development and body color, less sharply separable by the size of the scales on the snout. These populations are allopatric and thus may be species or subspecies. The differences between typical baleatus and typical ricordii are such that they could well imply specific distinction. How- ever, all the populations south of the Cul de Sac Plain, both the rather distinctive Barahona animals and the very poorly known populations of the southwest peninsula, are to some de- eree intermediate between the two northern extremes although presenting some features that are their own. In thus bridging the morphological gap between the extremes, they strongly sug- gest that the Hispaniolan giant anoles belong to a single species. The giant anoles are scarce and local. It is not to be expected that intergradation between the several populations will be easy to demonstrate; specimens from many of the critical intermedi- ate areas are conspicuously lacking. The Fond des Negres and Ti Guinin specimens, on the other hand, may be intergrade populations. The vestigial shoulder spots present in the two Ti Guinin males and the single Fond des Negres male suggest this conclusion, as does, in the latter case, the combination of nuchal and dorsal squamation most like typical ricordu with somewhat lower scale counts across the snout. No giant anoles are at hand from the northwest Domini- can Republic. This is the area in which intergradation between ricordu and baleatus is to be expected. It is thus not at all sur- prising to find the first suggestion of loss of characters of the typical race just to the east of Cap Haitien. 8 BREVIORA No. 155 On the southwest peninsula, the Fond des Negres area is one in which such intermediate populations are to be expected, as recent collections from the area show. Thus, Dromicus parvi- frons parvifrons and D. parvifrons protenus appear to meet in this area and several anole races (to be reported later in this series) show intergradation at just this point. I thus interpret the Fond des Negres giant anoles as inter- grades between typical ricordi and a population to the west occupying the tip of the southwest peninsula. This western popu- lation I infer to be represented at present by the single juvenile specimen collected by P. J. Darlington in the foothills of the Massif de la Hotte. The wide-banded specimen from Enriquillo in Barahona is reminiscent of the banded color phase in typical baleatus, but in squamation it is not intermediate and it is geographically quite unsuitable as a member of an intergradient population. Nevertheless, the presence of this color phase hints at a closer relation with baleatus than otherwise could be inferred. The absence of other evidence or hint of intergradation is easily accounted for by the gaps in the distributional record, and, granted the desirability of further evidence, recognition of subspecies status seems justified for typical ricordu, for baleatus, for the Barahona population and probably also for a population at the west end of the southwest peninsula of Haiti. The latter two populations are currently nameless. That from the southwest peninsula is at the moment very insufficiently known, and it would not now be appropriate to describe it. The Barahona population, on the other hand, is well recorded and may be formally named: ANOLIS RICORDII BARAHONAE hew subspecies Type: MCZ 43819, Polo, Valle de Polo, Barahona, Dominican Republic, an adult female collected by W. G. Hassler, Septem- ber 1932, J. C. Armstrong donor. Paratypes: Valle de Polo, MCZ 56141, AMNH 51036, 51235-6 ; Herman’s Finca near Paradis, AMNH 51231-3; Barahona, AMNH 50255-6, 50261; Halfway between Trujin and Enriquilo, AMNH 51230; Enriquillo, AMNH 51241. Diagnosis: A subspecies of ricordti distinguished from the typical form and from baleatus Cope by the nature of the nuchal crest (formed by small but slender tapering scales not or very 1962 HISPANIOLAN GIANT ANOLES 9 little higher than the highest scales of the dorsal crest), by the size of the scales of the snout (4-6 across snout between second canthals), and by a characteristic phase of coloration in both sexes in which small blotches are present, irregular in shape and of varying intensity. Inst of Specimens Examined: A. r. barahonae (17 specimens). DOMINICAN REPUBLIC. Polo, Valle de Polo: MCZ 43819; Valle de Polo: AMNH 51036, 51235-7, MCZ 56141; Barahona: AMNH 50255-6, 50261; Herman’s Finca near Paradis: AMNH 51231-3; Halfway between Trujin and Enriquillo: AMNH 512380; Enriqullo: AMNH 51241; locality uncertain: AMNH 51229, 51234, 51240. A.r. ricordu (12 specimens). HAITI. Port-au-Prince: AMNH 49501; Diquini: MCZ 8619, USNM 118902, 123988; Source Le- clerc, Morne Decayette, near Port-au-Prince: MCZ 65729-3831; Petionville: MCZ 60013-4; Marcaco: USNM 69437; Port-de- Paix: MCZ 63338; Ti Guinin near Cap Haitien: MCZ 66147-9. [Records by Mertens: Port-au-Prince, 4 specimens; Cap Haitien, 1 specimen. | A. r. baleatus (57 specimens). DOMINICAN REPUBLIC: Pena, near Santiago: MCZ 57713-9; Santiago: MCZ 7831; Las Bracitas: AMNH 41465-6; El Rio: AMNH 41294, USNM 62104- 5; Rio San Juan: USNM 74940-1; Samana Peninsula (various localities) : MCZ 5445, AMNH 28651, 39807-15, 39817-2383, 39825- 9, 39837, 40224-30, 40387-90, 42285, 42775, 44841-4; La Romana: MCZ 16321. [Records by Mertens: Santiago, 5 specimens; Moca, 4 specimens; Finca Arbol Gordo near Villa Altogracia, 1 speci- men. | A. r. subsp. nov. (1 specimen). HAITI: Foothills, Massif de La Hotte: MCZ 38277. A. r. subsp. noy. x r. ricordu. Pemel near Fond des Negres: MCZ 66015-6 ; Fond des Negres: USNM 72631, 72633 ; also 72632, skeletonized. BEHAVIOR The giant Anolis of Hispaniola is infrequently seen by collec- tors and consequently our information on its ecology and habits is limited. Such information as is available is summarized below by A. S. Rand. Quotation marks refer to Rand’s personal ob- servations. 10 BREVIORA No. 155 A. ricordu ricordu, Morne de Cayette, near Port-au-Prince, Haiti (A. S. Rand and J. Lazell). ‘‘ A large individual was seen about 15 feet up on a branch of a 30-foot high tree on the edge of a patch of brush on the hillside. It remained sitting quietly in sight for about half an hour until we climbed the tree to attempt to capture it. It then climbed out among the small branches where, despite an intensive search, we were un- able to locate it again.”’ Port-de-Paix, Haiti (A. S. Rand and J. Lazell). ‘‘A female was seen sitting head down on the trunk of a large tree about 4 feet from the ground. The tree was one of a series forming a fence row. The lizard allowed us to approach and noose it.’ A. ricordu baleatus, Pena near Santiago, Dominican Repub- lie (C. E. Ray and A. 8S. Rand). ‘‘ Local people who brought in a series at this locality reported that they hyed among the upper branches of the larger shade trees in the coffee plantations.’’ Finca Arbol Gordo, Dominican Republic. Mertens reports that the single specimen collected by him jumped to the ground from the trunk of a palm tree. Mertens kept this animal in cap- tivity for some time and wrote that it was active only in the late afternoon. When approached it squirreled around to the other side of its perch. It slept exposed on small branches with its hind legs flexed. A. ricordii subsp., Camp Perrin, Haiti (A. 8. Rand and J. Lazell). ‘‘A small boy guided Rand to a stand of coffee where a crowd of Haitians had treed a large individual. The lzard was about 20 feet up in a small tree. It was too high to noose and the tree too small to climb so we attempted to scare the lizard to a more favorable situation by poking it with a long pole. After considerable fuss the lizard ran out along a small branch to the next tree, down that to the ground, across the ground to hide among the roots of a large Cieba tree. After considerable prodding among the roots we dislodged the lizard which avoided the herd of assistants successfully and climbed the Cieba out of both sight and reach.”’ Above Maceline, near Camp Perrin (A. 8. Rand and J. Lazell). ‘“Tn a small stand of coffee we saw a single animal about 15 feet up in one of the large trees on a large branch. When Lazell climbed the tree it squirreled about the branch concealing all of itself except one eye. It then climbed up and out among the small branches and disappeared from view.”’ 1962 HISPANIOLAN GIANT ANOLES a ip | Mertens speculates that the giant anoles of Hispaniola were originally inhabitants of the lowland rain forest and have adapted to cultivated areas by living in the crown of the taller trees. The observations of Rand and Lazell tend to support this conelusion. INTRAISLAND ZOOGEOGRAPHY The portion of Hispaniola south of the Cul de Sae Plain, in- cluding the southwest and Barahona peninsulas and the La Selle and La Hotte ranges, has, as Mertens has emphasized, a number of endemic species and even genera. I have recently (Williams, 1961) stressed the importance of this area, which I have called the ‘‘southern island,’’ in the initial differentiation of forms how widespread in the portion of the island north of the Cul de Sac Plain. Barahonae as an isolate in ‘‘southern island’’ easily fits this pattern. Baleatus and typieal ricordiu do not. This is merely the first of many examples to be discussed in this series which will show that not all the patterns of differ- entiation in Hispaniola can readily be explained by the simple hypothesis of ‘‘southern’’ and ‘‘main island’’ isolates. As in the present case so in a considerable number of others, there appear to be additional loci of differentiation, e.g. northern Haiti, which are not so evidently marked off by present or recently past barriers to distribution. These cases of subsidiary loci of differentiation are not at present well analyzed. Hispaniola has not been really well col- lected, and it is now evident that the differentiation of local popu- lations that occurs within it is finer grained than the sporadic collecting of the past could reveal. For many of the instances of minor loci of differentiation, data are just now being gathered. ‘ RELATIONSHIPS OF RICORDIT AND THE OTHER WEST INDIAN GIANT ANOLES Anolis ricordi clearly belongs to the series of giant anoles that includes equestris of Cuba and cuviert and roosevelti of the Puerto Rican-Virgin Island complex. Anolis garmani of Jamaica seems, even on externals, to show no features, except size, which specially link it to this group, and Richard Etheridge in an unpublished thesis (University of Michigan) has shown that osteologically it belongs to quite a different section of the genus. 114 BREVIORA No. 155 No full study has ever been made of the Greater Antillean giant anoles as a group, though they clearly merit such a study. I do not intend to attempt this here, but in order to place the races of ricordu within a frame of reference which might add to our understanding of them and of their origin I have compared all of the species for a number of mostly qualitative characters which were simple to determine and to evaluate. Table 2 presents the results of this comparison. It is sufficiently clear even from the limited evidence presented by Table 2 that we are dealing with four good species. The Hispaniolan populations are a unit as compared with the forms of the neighboring islands and, as in the case of the other poly- typic giant anole, A. equestris, the intraisland differentiation has involved primarily scale size and color pattern. There is one unusual zoogeographical feature in the distribu- tion of the giant anoles — the occurrence of A. cuviert and A. rooseveltt on the same island bank, cuvicr? on the mainland of Puerto Rico, rooseveltt on Culebra in the Virgin Islands. These two are very distinct species. Their differences are at another level than those between ricordu and baleatus, and their oceur- rence on what must at various times during the Pleistocene have been a single land mass provides a special problem in accounting for their origin and dispersal. ACKNOWLEDGMENTS This paper was in part made possible by collections made under National Science Foundation Grants NSF G-5634 and G-16066 and grants from the American Philosophical Society and the Society of Sigma Ni. Assistance given by the govern- ments and people of the Haitian and Dominican Republics is also gratefully acknowledged. I am indebted also to Charles M. Bogert and Doris M. Cochran for loans from the collections under their care. Dr. Norman Hartwee permitted examination of the second known specimen of Anolis roosevelta Grant. Dr. W. J. Clench provided information on the areas visited by A. Salle in the Dominican Republic. 1 For further information on the non-Hispaniolan species see Stejneger (1904), Grant (1931), Schwartz (1958). 13 HISPANIOLAN GIANT ANOLES 1962 WUE Q9T &€ P2995] SMOL [BOTJIOA IB[NSIL UL posuvi4te oyearpenb ‘Tpeurg yyooug P ult yuesord 207 YJOOWS SaTRdS *pozVlyUIL1O ZIP Ips yoRoa WO [eIqe[qns JSOULLOTIO}UB ATWO yey ‘WJOOUIS SaTBaG nA (¢) sosptr Aq popunog ‘doo 8-9 peleey ‘WeT[OMG 19909800. WU GET && p9[99ST (FOGL ‘tesoulazg “gg pue 18 ‘ssy "JO) Jueur -O8UBIIB pu odLYS UL IV[NGILIT ‘OGAIVT LOYYEAY pelea ys P Ur yuosatg O}PVULIBOTI[NU WO}FO SelBos “poyeryUoa.lojfIp 20U MOL [BIGQRIGuYG sd[O19Q Ny aTdty[Nur [ITM SelBag Gg MOTTVYS 8-9 papeey “yep A[OATz Voy WIVAND wut JéT Té pees SMOL [BOTLIOA Iv[Nsal UL posuvs1e ayeipenb qpeug yyoowsg P UL JUOSOIg {joous SOlBOS “po PVIpUIIOJZIp TOM MOL [BIGVlqug UdT[OMS SOTBIG a7 MOTTVYS 6G PoPYULLA 10 pepeey ‘Wa[TOMG Mpsoors @ Wav, wut JCT yyooug SMOL [BOT}IOA Iv[NSII UL posuvIIV eyeipenb “peuug yooug P UL PUOSOAT yyoous SelVos ‘popVrIjuoloFIp [JOM MOL TBIGQRTqngG U9T[OMS SOTBIG SV ouo NT G-G esosnt ALVA WOTTOMG suugsanba P 4npe ‘qysuel JuUdA-JnoOuUg (eyeutxoidde ) 00} U}P Jopun ov] [owe'y S}ISTp pue SQ UIT WO SeTBIG [rez FO osvq JO opis UO SoPBog [leq FO oseq JapuN so[Rag STBUBYSO spTeIqelqug SO[OILOLULOS [VIIGr1ovrdns wor [BjoLied104UyT WOIssotdop [VPUO.LT JNOUS SSO.19V SaTBIG So[vos pvoT{ IopVIvYO 14 BREVIORA Noo 155 REFERENCES CITED BouLENGER, G. A. 1885 Catalogue of the lizards in the British Museum (Natural His- tory). London, vol. 2, i-xiii + 1-497. GRANT, C. 1931 A new species and two new subspecies of the genus Anolis. Jour. Dept. Agr. Puerto Rico, 15: 219-222. MERTENS, R. 1939 Herpetologische Ergebnisse einer Reise nach der Insel His- paniola, Westindien. Abhandl. Senckenb. Naturf, Ges., 449: 1-84. OLIVER, JAMES A. 1948 The anoline lizards of Bimini, Bahamas. Amer. Mus. Novitates, may, Isyj38 Ilsa. Scumipt, K. P. 1921 Notes on the herpetology of Santo Domingo. Bull. Amer. Mus. Nat. Hist., 44: 7-20. ScHWARTZ, A. 1958 A new subspecies of Anolis equestris from Eastern Cuba. Herpe- tologica, 14: 1-7. STEJNEGER, L. 1904 The herpetology of Puerto Rico. Rep. U.S. Nat. Mus., 1902: 549-724. WILLIAMS, HE. E. 1961 Notes on Hispaniolan herpetology. 38. The evolution and rela- tionships of the Anolis semilineatus group. Breviora, No. 136: 1-8. ANOLES GIANT HISPANIOLAN 1962 ‘sjoquiAs yuoovlpe Aq po}yeotpur suoTyeyndod UL ynouUs ssodoB SaTBos FO syuNOD oAv s1OquINN ‘SO[BOS [VSLOP IOMOT OY} FO OM} PUB SoTBOS [VYONU OMY FO oUTTZNO 9} “JUS 04 4JoT ‘poyuosordoa st [BAO Yovos UT] ‘SPIONII ,SUOJIOPT JV SNzNAIMG PUB WpLO9IU TOF SToqukS MOT[OFT ‘setoedsqns WpLooid SyoUpP FO WOTYNTAYST(T vo1njyndod saibay) SeD pvoy BB é uleyyod — vornjndod 3samy3n0¢ = spung asuaasvoiz Jo viajz0d — snyvajoq O @ sayo3o9 yows Jo vlazz0d — avvoyDI1Dg ® uv) Jods sapynoys — pio. O Hf £2 09 Sv OF SI oO SYaLIWOMY SO 31V90S _ a ae _ aed a ae — = a - = ——— - = 7 i 7 : —_ o ee Pa ad _ ae re = cal eo D gn | = é ae h — - in ry pt? sere 14 a 4 = i , up ry put gt aera) Baas fis e- is guawipe © — _ Bi -. . j i a ' b ! 7 a 4 nia . a a * te ie be ‘ as Ww) i af Preeie gE = ral ted - =. ° ta \ es - es = | vo ts © > ¢ ; tr) ia) ° e " % wh ea ae a7 & = - a) ny oe UV bh be oe! Foe Me - eo ae bie ro) 7 Wes he ch ” ; t + an ee 5 7 1s Ee “ft ae . an ’ Ve ae | _ ie fy oO 7 : - om gy 40) iv BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. APRIL 138, 1962 NuMBER 156 THE FOSSILIFEROUS TRIASSIC DEPOSITS OF ISCHIGUALASTO, ARGENTINA, AND PRELIMINARY DESCRIPTION OF ISCHIGUALASTIA, A NEW GENUS OF DICYNODONT By ALFRED SHERWOOD ROMER and C. Barry Cox THE FOSSILIFEROUS TRIASSIC DEPOSITS OF ISCHIGUALASTO, ARGENTINA By ALFRED SHERWOOD ROMER Early in 1958, a joint expedition of the Muséo Argentino de Ciencias Naturales of Buenos Aires and the Museum of Com- parative Zoology, Harvard University, was organized to explore continental deposits in the western Argentine in search of fossil vertebrates. The personnel included, for the Buenos Aires Mu- seum, Dr. Guillermo del Corro, geologist, and Mr. Orlando Gutiérrez, assistant ; from Harvard, Professor Bryan Patterson, preparators Arnold C. Lewis and James Jensen, Mrs. Romer and the writer. The earlier part of the trip was spent in the general region of Mendoza. In late April it was decided to move our base northward and explore the valley of Ischigualasto. This region hes some 300 miles to the northeast of Mendoza, in eastern San Juan Province, close to the border of La Rioja, in the department of Valle Fertil, at a latitude and longitude of approximately 68° W, 30°5'S. No adequate maps of this area exist; Ischigualasto is shown on the San Juan sheet (no. 37) of the 1/500,000 map of Argentina, but the details are highly inaccurate. Frenguelli (1948, fig. 1, pls. 1, 2) gives sketch maps showing the general topography of the valley and its relation to adjacent regions. The name derives from an Indian village once bo BREVIORA No. 156 present at the margin of the valley. This, however, disappeared long since and the valley is completely uninhabited. In 1874 Stelzner (1885, pp. xii, 74-75) rapidly traversed the valley on mule-back, presumably following a trail which runs from Jachal, in San Juan Provinee, through the valley and on eastward to Patquia in La Rioja. Bodenbender (1911, pp. 94-113) similarly crossed by this trail several decades later. A small coal seam had been discovered in the hills west of the valley at a locality named los Rastros because of dinosaur-like footprints discovered there; Huene (1931) was taken, in 1927, on a hasty trip to see these footprints; he traversed the valley but was unable to stop there. Until recent years nothing was known of the geology of the region except for such observations as Stelzner and Boden- bender could make as they crossed it, and nothing of fauna and flora except the footprints just mentioned and a few plants col- lected by Bodenbender. The reason for this former paucity of knowledge is obvious. The valley is extremely arid, and in the days of mule travel, a stay there was impossible because of the almost complete lack of water and fodder. With the coming of automobile transporta- tion and, particularly, of vehicles with four-wheel drive, the situation has changed radically, and in 1948 Frenguelli was able to publish a rather comprehensive account of the stratigraphy of Ischigualasto as the result of exploration of the area in 1943, primarily in the interests of paleobotany. During the course of his work there, three fragments of eyno- dont skulls and jaws were recovered, and were described by Cabrera (1943). These were of interest as indicating the pres- ence in Argentina of Triassic beds similar in fauna and presumed age to those of southern Brasil. But since so little reptile ma- terial was collected during Frenguelli’s survey of the region, it seemed unlikely that work there would be profitable for the ver- tebrate paleontologist. My attention was first called to Ischigualasto by Professor Huene before the results of Frenguelli’s work became known. As noted above, Huene had crossed the valley on mule-back during his inspection of the los Rastros footprint site and, al- though unable to stop and prospect, had been struck by the 1 Bodenbender (1911, p. 96) breaks off in the middle of his description of the region to insert a paragraph of emphatic and italicized warning: “NOTA —A los futuros exploradores hago presente que no se puede contar .. . con sufi- ciente pasto para los animales,” ete. 1962 TRIASSIC DEPOSITS OF ISCHIGUALASTO 3 seemingly favorable nature of the beds for vertebrate explora- tion. Like our Argentinian colleagues, I had failed to be im- pressed as to prospects by the few materials obtained by Frengueili. A different light, however, was shed upon the possi- bilities by a publication by Heim (1949). He had been commis- sioned in 1944 to report on the los Rastros coal mine, but in crossing the Ischigualasto valley had become so interested in the region that a considerable part of his report is devoted to the Ischigualasto beds and, incidentally, to the fossils found there. He photographed a cynodont skull in situ (pl. VIII, fig. 2) and says (p. 22): ‘‘El senor de la Vega me llam6 la atencién sobre los restos de vertebrados que se han hallado frecuentemente en el suelo de las arcillas, especialmente en la parte media de la formacién. Se presentan como acumulaciones de fragmentos de huesos, de color pardo oscuro. A veces aun se distingue la forma del créneo y de la dentadura con restos de dientes negros, pero sin que se pueda restaurar esqueletos.”’ This report indicated that investigation of the Ischigualasto region might prove profitable, and our expedition moved to that region. A road of passable quality runs to Valle Fertil, a village south of the valley, which serves as departmental capital. Be- yond this, settlement is sparse and presently ceases altogether, the roads deteriorate into a confusing series of trails, and it was necessary to hire a local guide to reach our destination.” At Cerro Morado, about 50 km. north of Valle Fertil, there is reached the southern edge of the valley which extends some 20 km. to the north-northwest. At its southern end its width is about 7 km.; it narrows to 214-3 km. at the north. Its western boundary is a range of rugged sandstone hills, the eastern boun- dary an unbroken cliff of red sandstone, ‘‘los Colorados,’’ strik- ing in appearance, with a height, for much of its length, of 150 to 200 meters. The valley itself, almost bare of vegetation except for scattered thorn bushes, exhibits an expanse of shales of varied pastel colors and occasional sandstones. It is drained by a series of broad sandy arroyos which converge at about the center of its western margin where, at Agua de la Pena, there is a small stream of water (non-potable, according to our guide). From this point the deep gorge of the Rio de la Pena runs west- ward through the hills to the Rio Bermejo. Northwestward from 2 Since our work there, a new highway has been constructed that links Patquia with Pagancilla and Chilecito. This passes a little to the northeast of the Ischi- gualasto basin, rendering it comparatively accessible. 4 BREVIORA No. 156 the main valley extends the narrower valley of the Rio de la Chilea, which turns westward to join the Bermejo. We made camp near Agua de la Pena, began exploration of the valley — and were immediately astounded by the abundance of vertebrate remains which it contained. Every vertebrate paleontologist dreams of finding, someday, a virgin territory strewn with fossil skulls and skeletons. Almost never does this dream come true. To our amazement and delight, it did come true for us at Ischigualasto. All about us, in the clays of the valley, were exposed specimen after specimen of fossil reptiles. In most instances the greater part of the time of a field party is taken up in prospecting for specimens. Here little of this sort of work needed to be done, and our energies could be de- voted to excavating the better specimens chosen from the wealth of materials readily available. Approximately six weeks were spent here. Our work was hampered by the shortness of the day- hight hours at this season of the year, and by the fact that a number of trips to Valle Fertil, each taking a full day, were necessary to earry out fossil blocks and obtain food supplies, gasoline and water. Nevertheless, well over 100 specimens, mainly plaster blocks containing skeletons, partial skeletons or skulls, were recovered. The geology of the region has been described by Frenguelli and by Heim, and discussed by Groeber and Stipanicie (1952, pp. 87-99). Apart from faults of a minor nature, the geologic structure is simple, the sediments uniformly dipping gently to the east-northeast with, in consequence, a regular succession of beds from west to east. Toward the Bermejo, in the western hills (an area not visited by us), are beds thought to be of Car- boniferous and Permian age — the Paganzo beds of Bodenbender —followed by the Estratos de Ischichuea.* Farther east, the hills bordering the Ischigualasto valley are formed by the Estra- tos de los Rastros, dominantly sandstones, which conformably overlie the Ischichuca and inelude the coal seam and footprint locality mentioned. The shales occupying the valley constitute the Estratos de Ischigualasto, with a thickness estimated at 400 to 500 meters; it is in this formation that nearly all the verte- brate fossils of the region have been found. The steep cliffs at the east side of the valley constitute the lower part of the Estratos de Gualo of Frenguelli, for which Groeber and Stipanicic prefer the designation of Estratos de los Colorados. The four upper 3 The beds beneath Cerro Morado may be the Ischichuca. 1962 TRIASSIC DEPOSITS OF ISCHIGUALASTO 5 formations, at least, of the series —the Estratos de Ischichuea, de los Rastros, de Ischigualasto, de los Colorados — are obviously parts of a single sedimentary cycle, without evidence of any dis- conformity. In the region of Agua de la Pena a local fault (Freneguelli, 1948, pl. II, profile A) obscures the transition be- tween los Rastros and Ischigualasto formations. But elsewhere (as Freneuelli’s profiles B-D, pl. IV) deposition can be seen to have been uninterrupted, and (apart from a conglomerate bed which is taken as the upper boundary of the los Rastros forma- tion) the transition is marked mainly by a diminution in im- portance of the sandstones which are so prominent in the los Rastros. The uppermost beds of the los Rastros yielded frag- mentary vertebrate remains which appear comparable to those of the Ischigualasto formation. The typical beds of the Estratos de los Colorados contrast strongly with those of the Estratos de Ischigualasto in color, predominance of sandstones, and re- sistance to erosion; but as Frenguelli notes, a zone of transition is apparent at the base of the cliffs. Stelzner ‘‘lumped’’ the entire series of beds found in this re- gion as ‘‘Rhaetic’’ (in the broad sense in which that term has frequently been used in Argentinian geology). Bodenbender believed the ‘‘Gualo’’ beds to be Cretaceous, the Ischigualasto Jurassic, the los Rastros ‘‘Rhaetic.’’ Frenguelli, more correctly, considers the series as a whole to be older, the ‘‘Gualo’’ to be Rhaetie or lower Liassic, the Ischigualasto to be upper Keuper, the los Rastros to be lower Keuper. Groeber and Stipanicie (table I) believe the spread in time of deposition of these beds to be rather narrower ; the los Colorados are assigned with doubt to the Rhaetie, and both Ischigualasto and los Rastros to the upper Norian —1.e., the uppermost Keuper. The vertebrate re- mains suggest a lower position (Romer 1960a, pp. 1291-1292; 1960b, pp. 86-87) although full discussion should be postponed until the fauna has been more thoroughly studied. The Norian stage is one in which is found the typical Upper Triassie dino- saurian fauna; the Ischigualasto fauna is, on the contrary, one in which there is little evidence of dinosaurs and in which eomphodont cynodonts and rhynehosaurs are dominant. The Ischigualasto formation is essentially comparable to the Santa Maria beds of Brasil and the Manda beds of Tanganyika. It is surely pre-Norian and not improbably pre-Carnian ; the gompho- dont-rhynechosaur faunas would appear to be essentially Middle Triassie in age. 6 BREVIORA No. 156 By agreement, the entire collection was shipped to Cambridge, where it is being prepared. All types and representative speci- mens of all forms found will be deposited in the National Museum in Buenos Aires. Complete preparation, however, will be a lengthy process, not only because of the considerable quantity of material but also because of the refractory nature of the mat- rix enclosing a large proportion of the specimens. In general, publication of the scientific results will not appear for some time, since we do not wish to publish before preparation has pro- ceeded to the point at which all material of a given form has become available for study. In the case of the dicynodonts, how- ever, the material included only a few specimens of a single large form; these have been prepared, and have been studied by Dr. Barry Cox, of King’s College, University of London. A prelimi- nary description is appended; it is hoped that a full account may be published within the year. It is gratifying to us to have been instrumental in opening up a new Argentinian area for exploration by vertebrate work- ers. Since our trip, several further expeditions to the region have been made by the University of Tucuman, under the direc- tion of Dr. Oswaldo Reig, with successful results; certain ma- terials collected on these later expeditions have already been described (Reig 1958, Casamiguela 1960). There are vast areas of late Paleozoic and early Mesozoic deposits in the western Ar- gentine which have never been visited by vertebrate paleontolo- gists. Although the chances of finding beds as unusually productive as those of Ischigualasto are not great, it is highly probable that other faunas which will aid in rounding out the early history of vertebrates in Argentina await discovery. I have elsewhere (Romer 1960a) expressed our thanks to a number of friends who aided us in the Mendoza region. We are further deeply grateful to various other persons who aided the general work of the expedition and our exploration of Ischigua- lasto. The cordial cooperation of Dr. Adolfo D. Holmberg, then Interventor, and members of the staff of the Buenos Aires Mu- seum, was much appreciated. Professor Rosendo Pascual of the La Plata Museum accompanied us during the early portion of the trip. Much valuable scientific information was given us by the geologists of the Yacimientos Petroliferos Fiscales, Comision Nacional de Energia Atémica and the Direccién Nacional de Mineria, including, among others, Drs. Pablo Groeber, Pedro N. Stipanicic, Martinez Cal, Héctor de la Mota, Luis A. Barrio- nuevo, and Vicente Ferreiro. The Comisién Nacional de Energia 1962 TRIASSIC DEPOSITS OF ISCHIGUALASTO th Atémica aided us in the difficult matter of water and gasoline supply at Ischigualasto. Dr. Mario E. Terrugi aided us greatly in many regards, and the Harvard members of the expedition appreciate very much the hospitality extended to them by Dr. and Senora Terrugi and by the late Dr. Bernhard Dawson and Sefiora Dawson of La Plata. Our expedition was made possible by grants from the National Science Foundation and [nfe magazine. REFERENCES CITED BODENBENDER, G. 1911. Constitucién geolégica de la parte meridional de La Rioja y re- giones limitrofes (Reptblica Argentina). Bol. Acad. Nac. Cience. Cérdoba, 19:1-220. CABRERA, A. 1943. El primer hallazgo de terfpsidos en la Argentina. Notas Museo La Plata, 8, Paleont. no. 55:317-331. CASAMIGUELA, R. M. 1960. Noticia preliminar sobre dos nuevos estagonolepoideos argen- tinos. Ameghiniana, 2:3-10. FRENGUELLI, J. 1948. Estratigrafia y edad del llamado Rético en la Argentina. Gaea, 8:159-309. GROEBER, P., AND P. N. STIPANICIC 1952. Tridsico, pp. 13-141 in Geografia de la Reptblica Argentina. Buenos Aires, Soc. Argentina Estudios geog. Gaea, 2 (1):1-541. Heim, A. 1949. Estudio geol6égico del Carbon ‘‘Retico’’ y del Valle de la Pena (Provincias de San Juan y la Rioja). Ministerio de la Industria y Comercio, Bol. Dir. General de Industria Minera, 69:1-31. HUuENE, F. 1931. Die fossilen Fahrten im Rhit von Ischigualasto in Nordwest- Argentinien. Palaeobiologica, 4:99-112. Reta, O. A. 1958. Primeros datos descriptivos sobre nuevos reptiles arcosaurios del Triassic de Ischigualasto (San Juan, Argentina). Rev. Assoc. Geol. Argentina, 13:257-270. Romer, A. S. 1960a. Vertebrate-bearing continental Triassic strata in Mendoza region, Argentina. Bull. Geol. Soe. America, 71:1279-1294. 1960b. Explosive evolution. Zool. Jahrb., Syst. Bd., 88:79-90. STELZNER, A. 1885. Beitriige zur Geologie und Palaeontologie der Argentinischen Republik. Geologischer Theil. Capel & Berlin, xxix + 329 pp. 8 BREVIORA No. 156 PRELIMINARY DIAGNOSIS OF ISCHIGUALASTIA, A NEW GENUS OF DICYNODONT FROM ARGENTINA By C. Barry Cox Department of Zoology, King’s College, University of London Above is given a brief account of a joint Buenos Aires-Har- vard expedition to the Valley of Ischigualasto, San Juan Province, Argentina. Among the remains collected from the Ischi- gualasto formation, presumably of Middle Triassic age, were several skulls and parts of posteranial skeletons belonging to a new genus of large dicynodont. A preliminary diagnosis of this new genus follows below; it is named [schigualastia jenseni after Mr. James Jensen, who was responsible for the extremely pains- taking collection and preparation of this material. ISCHIGUALASTIA JENSENI, gen. et sp. nov. Holotype of I. jensent: Number 18,055, Muséo Argentino de Ciencias Naturales, consisting of skull and partial skeleton. Geological Horizon and Locality: Ischigualasto formation (Triassic), approximately 100 m. above the base of the forma- tion ; about 2 km. north of Agua de la Pena, Ischigualasto Valley, Department of Valle Fertil, San Juan Province, Argentina. Genotype: Ischigualastia jenseni Cox. Generic and Specific Diagnosis: Large dicynodont (type skull 55 ems. long, 46 ems. broad). No teeth in upper or lower Jaws. Skull triangular in dorsal view, greatest width across occiput. Very wide interorbital region, very narrow intertemporal re- gion. Tapering snout, without nasal ridges or bosses. No pineal boss, but a slight mound in front of pineal foramen. No post- frontal bone. Preparietal bone probably present. Interparietal forms whole of posterior half of intertemporal bar, widely sepa- rating squamosals from postorbitals. No sharp median inter- temporal ridge. Zygomatie arches bowed outward. Sharp transition between dorsal and occipital surfaces. Occiput almost semicircular in outline. No tabular bone visible. Stapes lacks stapedial foramen. Short interpterygoid vacuity. No ectoptery- goid bone. Pterygoid broadly meets maxilla. Palatine and pre- maxilla meet, excluding maxilla from internal nares. Palatal surface of premaxilla bears pair of anterior ridges. Premaxilla 1962 TRIASSIC DEPOSITS OF ISCHIGUALASTO 9 extends some way anterior to maxilla. Ascending portion of epipterygoid slender, not expanded to form part of lateral wall of brainease. No lateral wing on dentary. Stout retro-articular process. Five sacral ribs. Acromion process of scapula absent or ves- tigial. Coracoid foramen between precoracoid and_ scapula. Sternum constricted halfway along its length; dorsal surface bears bosses for attachment of ribs. Ulna has large olecranon process, with cartilaginous epiphysial union with rest of bone. Femur with well-developed head set off from rest of bone. A more extensive and illustrated account of Ischigualastia will appear later. * a 4 va a : aa ee > = - ~~ 2 eal BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. May 28, 1962 NUMBER 157 A RHACHITOMOUS AMPHIBIAN, SPATHICEPHALUS, FROM THE MISSISSIPPIAN OF NOVA SCOTIA By Donaup Batrp Department of Geology, Princeton University, Princeton, New Jersey In the past few years field parties from the Museum of Com- parative Zoology at Harvard College, supported in part by funds from the National Science Foundation, have discovered several deposits of fossil amphibians which date from the earlier half of the Carboniferous period. Most of the material recovered comes from redbeds of the Upper Mississippian Mauch Chunk group in West Virginia (Romer, 1941) and Pennsylvania; more recently, amphibians have been found in beds of equivalent age at two sites in Nova Scotia. In continuation of the Harvard collecting program, field par- ties sponsored jointly by Princeton University and the Nova Scotia Museum of Science have collected additional bones and trackways of amphibians in Pennsylvanian as well as Mississip- pian beds. One of these finds is noteworthy because no member of the order Temnospondyli from the Mississippian system has hitherto been described. This specimen, the right half of a skull table preserved as a natural mold, comes from the same horizon and locality as the embolomerous lower jaw described as Pholiderpeton(?) breton- ense Romer (1958). The matrix is a flagey caleareous siltstone characterized by well-sorted quartz grains and silvery muscovite mica. bo BREVIORA No: asi Superorder LABYRINTHODONTIA Order TEMNOSPONDYLI Suborder RHACHITOMIE' Family LOXOMMIDAE Genus SPpATHICEPHALUS Watson 1929 [Spathiocephalus Romer 1945, errore | Emended Diagnosis. A loxommid extremely specialized in its expanded cheek and snout, constricted interorbital area and shortened skull table, and having numerous small, chisel-shaped marginal teeth. SPATHICEPHALUS PEREGER, Nl. Sp. Diagnosis. Differs from 8. mirus Watson, the only other known species, in its relatively narrower skull table. Type. PU 17182, Princeton University Geological Museum. Occurrence. Point Edward formation, Canso group, Upper Mississippian (probably early Namurian). Beach of cove be- tween Point Edward and Keating Cove, 4 miles northwest of Sydney, Cape Breton County, Nova Scotia. Collected by Donald Baird, William F. Take and Jane MeN. Take, 1960. DESCRIPTION The dorsal surface of the skull table bears a deep, coarsely reticular sculpture rather similar to that of Megalocephalus lineolatus (Cope) but with narrower ridges. Aside from the separation of prefrontal from postfrontal, the contacts between skull elements are normal for a loxommid. The narrow frontal bone is bounded medially by a deep, straight suture which is ridged rather like the edge of a file. Anteriorly the frontal is beveled to receive the end of the nasal; anterolaterally it bears a long, striated facet for the articulation of the prefrontal. As in other loxommids the prefrontal must have swelled laterally to form the antorbital process which dif- | Assignment of the Loxominidae to the Rhachitomi is now confirmed by the association of rhachitomous vertebrae with a skull of Megalocephalus lineolatus (Cope) from Linton, Ohio (cf. Baird, 1957). 1962 NEW SPATHICEPHALUS 3 ferentiates the orbit proper from its anterior extension, the lacri- mal fenestra, and gives the loxommid eye-socket its characteristic keyhole shape. Most of the lateral edge of the frontal forms the thick orbital rim. At its waist the frontal is 4.3 mm. wide, mak- ing the interorbital distance a mere 8.6 mm.— extraordinarily narrow for a skull with an estimated width of 185 mm. The short parietal extends laterally into a square lappet ; there is no indication that this lappet represents a former intertem- Be ee ee sao Ze | 7 pm | ye | Tg re / { = | Zp if \ | Ligeti ‘ / aes ae. \ | h ee i NG | / | | \ | i | | lief : | | Ly ‘ | ee ‘ | | \ : | | \ \ mor — = aa as ” fo) S \ SS 4+ Jo Ck 7. Vf, oe \ po\ie a ~~ Fig. 1. Skull of Spathicephalus pereger, n. sp., x %. Restoration in dashed lines based on S, mirus; dotted lines conjectural. Abbreviations: f, frontal; j, jugal; /, lacrimal; m, maxilla; 1, nasal; p, parietal; pf, post- frontal; pm, premaxilla; po, postorbital; pp, postparietal; prf, prefrontal; q, quadrate; qj, quadratojugal; sq, squamosal; st, supratemporal; f tabular. , 4 BREVIORA No: bi poral element. Centered in the anterior half of the inter-parietal suture hes the parietal foramen, 2.3 mm. in diameter, which housed the parapineal eye. The postparietal (dermal supraoc- cipital) is relatively large. A thin oval lamina slopes outward from the corner of the tabular. Ventromedial to this flanee lies the backward-jutting tubercle which is characteristic of Joxommid tabulars but which also occurs in such genera as Hdops (Romer and Witter, 1942). Whether this process is a true homoloe of the tabular horn in other labyrinthodonts and perhaps of the tabular lamina in Acanthostega remains to be determined. Unfortunately, the lat- eral margins of the tabular and supratemporal are not clearly defined in the specimen. The greatly broadened supratemporal, however, is evidently embayved by the apex of the otic notch —a condition also found in Baphetes and Megalocephalus brevicornis |Orthosaurus pachycephalus of Watson, 1926]. The postfrontal, which in most labyrinthodonts hes postero- medial to the orbit, here lies directly posterior to it. In conse- quence the postorbital has been thrust outward onto the cheek along with most of the supratemporal; its orbital margin is 19 mm. long and nearly straight. In its new orientation the post- orbital might be called exceptionally lone rather than excep- tionally broad. Watson’s (1929, fig. 22) illustration of the skull of Spathi- cephalus mirus, sealed to the interorbital width and post-orbital length of S. pereger, forms the basis for the reconstruction in Fieure 1. The restored skull of S. pereger measures 177 mm. in mid-sagittal leneth as compared to 198 mm. for the type skull of S. mirus. As nothing is known of the posteranial skeleton in this genus, the dimensions of the whole animal cannot be esti- mated. COMPARISON Inconsistencies between the Point Edward specimen and the published account of Spathicephalus mirus might be interpreted as generic distinctions. Many of the discrepancies, however, can be attributed to differences in the manner and quality of preser- vation. For example, the suture pattern illustrated by Watson is that of the ventral surface of the skull roof and can be ex- pected to differ somewhat from that of the dorsal surface. 1962 NEW SPATHICEPHALUS 5 The single element labeled ‘‘postorbital’’ in Watson’s figure of S. mirus occupies the position of the postfrontal and half of the postorbital in S. pereger. Since the suture between jugal and squamosal meets the outer corner of this element —as it does the corner of the postorbital in all known loxommids — Wat- son’s identification must be correct. If (as he suggests) the postfrontal forms part of the undifferentiated interorbital bar in S. mirus, then the two species differ markedly in this respect. Watson illustrates a suture which bisects the antorbital process at right angeles to the lateral margin of the nasal bone, and in- terprets this suture as separating the lacrimal from a wing of the interorbital bar. New evidence now permits a more conven- tional reconstruction of this region. No parietal foramen was observed in the Scottish material, but a foramen of normal size and position is evident in the Nova Scotian specimen. One clear distinction hes in the relatively narrower skull table of the new species: in S. mirus the ratio of interorbital width to table width is 1:7 while in NS. pereger it is 1:6. Another difference is that even when allowance is made for post-mortem distortion of the cheek region, the orbits seem to have been more obliquely placed in S. pereger than in S. mirus. DERIVATION OF SPATHICEPHALUS Grotesquely modified though it is, the skull structure of Spathicephalus (as elucidated by the new specimen) can be read- ily derived from that of more typical loxommids. As it happens, one species which can be said to represent a pre-Spathicephalus morphological stage is also the only older loxommid known: Loxomma allmanni Huxley from the Gilmerton ironstone of Scotland. The Lower Limestone group in which it was found directly underlies the Limestone Coal group in which Spathi- cephalus mirus occurs. In the transition between the morphological stages represented by Loxomma and Spathicephalus, posteromedial migration of the orbits has displaced the supraorbital elements so that the frontal bone forms much of the orbital rim—a condition repeated in various genera of labyrinthodonts but more commonly associated with enlarged orbits. Concomitantly the cheek and snout have broadened and flattened into a condition which recalls the early Permian zatracheids Acanthostoma and Zatrachys. The con- striction of the frontals into a narrow bar, together with the 6 BREVIORA No. 157 shortening of the skull table, loss of the intertemporal bone and lateral displacement of the postorbital and supratemporal, are unexpected extremes of specialization in so early a genus. Such a condition did not reeur, so far as we know, until the de- velopment of the plagiosaurs in late Triassic time (cf. Panchen, 1959; fig 16). The specializations of Spathicephalus evidently adapted it to life as a bottom-dwelling fish-eater. Among labyrinthodonts of this ecotype — e.g. Erpetosaurus, Zatrachys, Capitosaurus, Eu- pelor |Buettneria|, Gerrothoraz —the lower jaw articulation tends to be aligned with the occipital condyle, so that the mouth could be opened by raising the snout while the thorax remained prone on the bottom (ef. Watson, 1951, pp. 67-70). A similar tendency is evident in Spathicephalus, for its occipital and quad- rate condyles are more nearly aligned than those of any other loxommid except the late Westphalian species of Megalocephalus. Surely the extent of modification which Spathicephalus had achieved by late Mississippian time presupposes a long previous history — yet to be revealed — tor the Loxommidae. ASSOCIATED FAUNA A Gyracanthus spine, a supposed Sagenodus quadrate and fragmentary fish remains were found in the flagstone layer with the mandible of Pholiderpeton(?) bretonense in 1956. From this same layer the 1960 field party collected the type of Spathicepha- lus pereger and the proximal third of another Gyracanthus spine (GEUS ae eyey ie About 12 feet lower in the section, bones coated with red iron oxide occur in an 18-inch zone of highly caleareous mudstone, rich in ostracodes and Spirorbis, which is intercalated with layers of limey shale. A Gyracanthus spine (PU 17186), a skull bone of the lungfish Sagenodus (the left ‘‘E’’ plate, PU 17187), and scales and scrap of the crossopterygians Megalichthys and Strep- sodus (PU 17188, 17189) represent a typical assemblage of Carboniferous fresh-water fishes. Amphibian remains from this horizon include a left clavicle, a phalanx and a ventral scale (PU 17190), all apparently embolomerous. STRATIGRAPHY The Point Edward formation is assigned to the lower part of the Canso group. While the Canso cannot yet be correlated pre- cisely with European and other American sequences, it appears 4 1962 NEW SPATHICEPHALUS on the basis of fossil floras (Bell, 1944, pp. 23-24) and inver- tebrates (Copeland, 1957, pp. 6-8) to be more or less equivalent stratigraphically to the Lower Namurian of Europe. Since in current practice the lower boundary of the Upper Carboniferous system coincides with that of the Namurian, the Point Edward formation is classified in European terms as basal Upper Car- boniferous. In North America, however, the Carboniferous is subdivided differently: its upper half constitutes the Pennsyl- vanian system which includes equivalents of the Upper but not the Lower Namurian. Thus the Point Edward formation is clas- sified in American terms as Upper Mississippian (Weller et al., 1948). The Scottish specimens of Spathicephalus were found in the Loanhead No. 2 ironstone of the Midlothian coalfield, a bed which hes in the upper half of the Limestone Coal group (Tulloch and Walton, 1958). This group is believed to be approximately equivalent to Goniatite Zone EK, (characterized by Eumorpho- ceras) and the lower part of Brachiopod-Coral Zone Ds (charac- terized by Dibunophyllum) ; these zones correspond to the Lower Namurian of continental Europe (Trueman, 1954). At the risk of confusion it should be added that although the Eumorphoceras Zone forms the base of the Upper Carboniferous, the flora and the fish fauna of this zone show Lower Carboniferous affinities and are separated from their Upper Carboniferous counterparts by a distinct biotic break. The sequence of amphibian-bearing beds in the Scottish Carboniferous has been reviewed by Westoll (1951) and most recently by Panchen and Walker (1961). In summary, although the correlations involved are still rather tentative on both sides of the Atlantic, current stratigraphic practice assigns an early Namurian age to both the Scottish and Nova Scotian species of Spathicephalus. The epoch in which they lived was evidently one of biotic transition between the earlier and later phases of the Carboniferous. In view of the stratigraphers’ uncertainty, the possible utility of Spathicephalus as an index fossil is worth investigating. For the most part the loxommid amphibians are too imperfectly known for their distribution to be stratigraphically significant. The best-known genus, Megalocephalus |Orthosaurus|, ranges through early and middle Pennsylvanian time, i.e. from West- phalian substage A to late Westphalian D (Panchen and Walker, 1961; Baird, 1957). But since Spathicephalus is the most ex- tremely specialized member of the family it may have existed over a comparatively short span of geologic time. 8 BREVIORA No. 157 SUMMARY A partial skull roof of the loxommid labyrinthodont Spathz- cephalus, previously known only from the Limestone Coal group of Scotland, has been found in the Point Edward formation of the Canso group near Sydney, Nova Scotia. The suture pattern (hitherto incompletely known) of this extremely specialized genus is described and derived from that of more typical loxom- mids. Spathicephalus-bearing beds of Scotland and Nova Scotia are considered to be early Namurian in age, corresponding to the base of the Upper Carboniferous in Europe and the top of the Mississippian system in North America. S. pereger n. sp. is the first-deseribed temnospondylous amphibian of Mississip- pian age from the Western Hemisphere. ACKNOWLEDGEMENTS This study owes much to the hospitality and cooperation of Director Donald K. Crowdis and staff members William F. and Jane MeNeill Take of the Nova Scotia Museum of Science in Halifax. F. B. Van Houten analyzed the source rock; A. L. Panchen provided most helpful advice on matters of British stratigraphy ; Edward S. Belt has generously shared the results of his current research on Canso stratigraphy and sedimenta- tion. Financial support from the Wiliam Berryman Scott Re- search Fund of Princeton University (administered by Glenn L. Jepsen) and the National Science Foundation made possible the field work. REFERENCES BairRD, DONALD 1957. Rhachitomous vertebrae in the loxommid amphibian Megalo- cephalus [abstract]. Geol. Soc. America Bull., vol. 68, p. 1698. BELL, W. A. 1944. Carboniferous rocks and fossil floras of northern Nova Scotia. Canada Geol. Survey Mem. no. 238, 277 pp., 79 pls. COPELAND, M. J. 1957. The arthropod fauna of the Upper Carboniferous rocks of the Maritime Provinces. Canada Geol. Survey Mem. no, 286, 110 pp., 21 pls. PANCHEN, A. L. 1959. A new armoured amphibian from the Upper Permian of East Africa. Roy. Soe. London Phil. Trans. (B), vol. 242, pp. 207- 281, pl. 8. 1962 NEW SPATHICEPHALUS 9 ——--———. AND A. D. WALKER 1961. British Coal Measure labyrinthodont localities. Ann. Mag. Nat. Hist seralloyvOla ds ppaosl-ooe- Romer, A. S. 1941. Earliest land vertebrates of this continent. Science, vol. 94, p. 279. 1945. Vertebrate paleontology. Uniy. Chicago Press, ix + 687 pp. 1947. Review of the Labyrinthodontia. Mus. Comparative Zool. Bull., vol. 99, pp. 1-368. 1958. An embolomere jaw from the Mid-Carboniferous of Nova Seo- tia. Mus. Comparative Zool. Breviora, No. 87, 8 pp. ————— AND R. V. WITTER 1942. Edops, a primitive rhachitomous amphibian from the Texas Red Beds. Jour. Geol., vol. 50, pp. 925-960. TRUEMAN, ARTHUR (ED.) 1954. The coalfields of Great Britain. London: Edward Arnold, xi + 396 pp., 7 pls. TULLOCH, W., AND H. S. WALTON 1958. The geology of the Midlothian Coalfield. Geol. Survey Scotland Mem., vii + 157 pp., 5 pls. Watson, D. M. S. 1926. The evolution and origin of the Amphibia. Roy. Soc. London Phil. Trans. (B), vol. 214, pp. 189-257. 1929 [**1923’’]. The Carboniferous Amphibia of Scotland. Palaeon- tologia Hungarica, vol. 1, pp. 219-252, pls. 1-3. 1951. Paleontology and modern biology. New Haven: Yale Univ. Press, xii + 216 pp. WELLER, J. M., ET AL. 1948. Correlation of the Mississippian formations of North America. Geol. Soc. America Bull., vol. 59, pp. 91-196. WESTOLL, T. S. 1951. The vertebrate-bearing strata of Scotland. Internat. Geol. Cong., Rept. 18th Session (11), pp. 5-21. ‘OSMOAIL UL PoJULId PUB poPBULUM][L Wood SBy TorM ydeasojyoyd B WOLF patoqsot JLB Jfory “YSvo xopVp poaopootq B woz “ds cu “wahadad snjoydaoiyjody Jo opqey [ays BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. May 29, 1962 NUMBER 158 A FOSSIL GERRHOSAUR FROM THE MIOCENE OF KENYA (REPTILIA: CORDYLIDAE) By RicHarp Estes! Many thousands of fossils, both plant and animal, have been collected since 1947 in the Kavirondo Gulf area, Lake Victoria, Kenya. Most of these fossils come from sediments on Rusinga and Mfanganu Islands, within the Gulf. The vertebrate fauna is diverse, though principally mammalian; thus the discovery of the fossil lizard described here is of special interest. Much of the fossil mammalian fauna has been described in a British Museum of Natural History series (see e.g. LeGros Clark and Leakey, 1951; Whitworth, 1958). Part of the extensive seed-nut flora has recently been described by Chesters (1957). Leakey (1952) has noted and figured some of the remarkably well-pre- served invertebrates. The age of the fossils in this sequence has been assigned en- tirely on faunal grounds, and is usually considered to be early Miocene (Burdigalian). The reasons for this age determination are given in the papers mentioned above. The presence of both archaic and modern elements in the mammalian fauna as well as our lack of knowledge of the Cenozoic sequence in tropical areas make it difficult to assign a firm date to these deposits. Suggestions ranging from Oligocene to ‘‘considerably younger than the lower Miocene’’ (LeGros Clark and Leakey, 1951) have prompted Whitworth (1958, p. 45) to state that ‘‘it is clear that the question of geological age may have to be revised when the description of the East African fossils is completed.’’ 1 Department of Biology, Boston University, and Research Associate, Museum of Comparative Zoology, Harvard University. 2 BREVIORA No. 158 Class REpTILIA Order SAURTA Family CORDYLIDAE Subfamily GERRHOSAURINAE GERRHOSAURUS ef. G. MAJOR Dumeril Referred specimen. — Coryndon Museum, Nairobi, M. F. W. 1955/1. Locality. — Mfanganu Island, Lake Victoria, Kenya, Africa. Area B, of Red Earth Series as mapped by Whitworth (1953). An extensive flora and invertebrate fauna have also been col- lected from these deposits, but have not yet been described. Age. — Karly Miocene (Burdigalian?). Preservation and major features of the specimen. — As pre- served, the total length of the specimen is 55 millimeters. It consists of most of the head and the greater part of the neck, though it is probable that much more if not all of the entire body was originally present. The external features, many of them from the soft anatomy, are in part completely replaced with calcite. Some bone is preserved internally, but selective replacement has occurred. The external caleite covering is about 3 or 4+ millimeters thick, and internal to this the neck and pos- terior skull region are filled with a soft waxy black sediment which contains small calcite erystals. No trace of the posterior bones of the skull is visible. The head is shehtly twisted to the left, and the half-open mouth is filled with caleite. A striking feature of this specimen is the replacement of the tongue by ealeite, though no important details of structure are visible. Dorsally, especially in the neck and posterior skull region, the specimen is heavily crushed; otherwise, there is little distortion. The shape of the depression in the top of the skull, the half-open mouth, and lack of crushing in the facial segment suggest that the animal may have been stepped on, perhaps by the sharp hoof of a grazing animal. This might have caused the animal’s death, or could also have happened shortly after it was buried by sediment. The palatal, facial, and marginal bones of the snout are broken or missing. Most of the teeth are eroded, though their outlines are preserved as imprints in the ealeite filling of the mouth, but in some places a few tooth crowns still remain, as well as frag- ments of tooth shafts. 1962 MIOCENE GERRHOSAUR FROM KENYA 3 The eye sockets are completely filled with calcite, though the right one is badly eroded. On the left side, the calcite has taken an impression of the underside of the supraorbital plates, but their number cannot be determined. One of the most unusual features of this specimen is the preservation of the external shape of the left eve, in calcite. The lids are sharply delineated, and between them, a low, domed area reflects the outline of the cornea or perhaps the underlying lens. Fig. 1. Gerrhosaurus ef. G. major, Lower Miocene, Kenya, right lateral view, x 1.5. The outline of the outer ear and tvmpanum is perfectly pre- served on the right side, though on the left much of it is broken away. The anteroposterior diameter of the ear opening is less on the left as a result of twisting of the head. Many of the granular scales which surround the ear region and extend into the lateral fold are clearly visible. The tip of the extracolumella is stronely imprinted on the tympanum. The lateral fold is prominent on both sides, but is especially clear on the right, where it has been stretched open by twisting of the body on the median axis be- fore deposition. Very little remains of the posterior skull bones. A thin film of prefrontal and palatine surrounds the left eye, and directly below, small portions of the vomers protrude. Maxillae and dentaries are badly broken, and little remains other than thin edges of their internal processes. Under both eves parts of the internal faces of the Jugals are visible as imprints. Dorsally, there are imprints of seven or eight postcranial scale rows. They end anteriorly near a smooth flat surface which 4 BREVIORA No. 158 represents the skull roof, but whatever bone or scale imprints may have been preserved are no longer present. Description. — The large imbrieate osteoscutes of the throat region are compound; each is formed of multiple polygonal or trapezoidal osteoderms. The anterior ones are small, smooth, and subequal, the posterior ones elongated and either smooth or faintly wrinkled. The posterior borders of the osteoscutes are rounded or slightly squared. The throat osteoseutes are in seven longitudinal rows between the lateral folds, and they alternate rather than being aligned in straight transverse rows. Pos- teriorly, their arrangement is somewhat distorted as a result of post-mortem dislocation of some of the seales. Two pairs of large non-imbricate chin shields seem to have been present. formed of small subequal polygonal osteoderms. The lateral folds appear to have contained small or granular scales, but this is uncertain. Both folds extend as far forward as the ear. The subtriangular outer ear opening is covered ventrally with small squarish or lenticular scales which grade into those of the lateral fold. On the posterior border of the ear opening, several marginal rows of small seales are followed by a row of slightly larger ones. The tympanie shield is strap-shaped, narrow, and not at all posteriorly expanded. The dorsal scales of the neck, of which imprints of seven or eight straight transverse rows are preserved behind the occipital marein, are large and subrectanegular. Abrasion has removed almost all imprints of scales from the cheek, but a few large ones are visible anteriorly. The pleurodont teeth are robust, tall, and columnar. Imprints of their closely-spaced shafts indicate that replacement teeth were formed in subcircular basal excavations. Preserved tooth crowns are faintly tricuspid. Discussion. — Union of the gerrhosaurs and cordylines as sub- families of the Cordylidae, as recently suggested by McDowell and Bogert (1954), is undoubtedly correct. The presence of compound osteoderms in cordylines (sensu stricto), a character not mentioned by these authors, further emphasizes the separa- tion of this group from the anguimorphs and allies them with the scincomorphs. Until now, the cordylds have had no clear fossil reeord. Pseudolacerta mucronata (Filhol) has been tentatively placed in this family by Romer (1956, p. 552). De Stefano (1903, p. 413) 1962 MIOCENE GERRHOSAUR FROM KENYA 5 and Filhol (1877, p. 489) do not mention any characters of taxonomic value, and Filhol’s illustration (ibid., fig. 423) is vague and diagrammatic. Hoffstetter (1944, p. 553) considers it possibly a skink, but notes that vertebrae similar to those of Cordylus (sensu latw) occur in the same deposit. Later, he indi- cates (1955, p. 621) that these fossils ‘‘rappellent les piéces homologiques de l’actuel Cordylus,’’ but the assignment is still tentative. The fossil described here is referable to the subfamily Gerrho- saurinae on the basis of the large, broadly imbricate, rounded or slightly squared throat scales, contrasting with the much smaller, non-imbricate, diamond-shaped throat scales of the cor- dylines. Chamaesaura has larger throat scales than other cordy- lines, but they are anteroposteriorly elongate and mucronate, rather than smooth and transversely widened as in gerrhosau- rines. Fig. 2. Gerrhosaurus ef. G. major, Lower Miocene, Kenya, left lateral view, x 1.5. Though the published generic characters of Gerrhosaurus are in soft anatomy and scale details not preserved here (Loveridge, 1942, p. 488; FitzSimons, 1943, p. 268), similarity between this fossil and Recent members of the genus seems to indicate that it belongs here. The throat scales, what remains of the dorsal scales, shape of the ear and lateral fold all agree closely with these characters of the Recent genus. Moreover, a number of resemblances discussed below strongly suggest reference to the Recent species G. major. 6 BREVIORA No. 158 1. The fossil has the size and general proportions of a large adult individual of G. major. The latter is the largest of the species of Gerrhosaurus; most other species are considerably smaller. 2. The shape of the opening of the external ear in G. major may be rounded or slightly angular dorsally. In the available specimens, G. m. major most frequently shows the rounded con- dition, but G. m. grandis usually has a more angular dorsal edge, as in the fossil. G. m. bottegoi resembles G. m. major in this character. 3. In gerrhosaurs, the shape of the tympanic shield is con- sidered taxonomically significant. This scale hes on the anterior border of the outer ear, and in all species of Gerrhosaurus except G. major, is thin, flattened, and often expanded to cover and protect the cavity of the outer ear (see e.g. FitzSimons, 1943, figs. 150, 156). Loveridge (1942, pp. 515, 518) states that tym- pame shields of G. favigularis are also narrow and band-like as in G. major. This is grossly true, but in detail the two can be distinguished easily. G. flavigularis has a narrowly ecrescent- shaped tympanic shield (see e.g. FitzSimons, ibid., fig. 154; ef. fig. 164 of G. m. grandis) which is thin and flattened, while that of G. major is strap-shaped, and thickened. 4. Another similarity to G. m. grandis is the presence, on the posterior border of the outer ear opening, of a small anterior row of scales, flanked by a larger posterior row. In G. m. major and G. m. bottegoi these scales tend to be subequal. This charac- ter varies somewhat, and in any ease the time separation as well as lack of further preserved characters precludes reference of this fossil to one of the vine subspecies. However, this charac- ter and that given as number 2 above seem to suggest a closer relationship to G. m. grandis and G. m. bottegoi than to G. m. major. The other subspecies, G. m. zechi, is known from only a few specimens, none of which were available to me. It is very closely related to G. m. bottegot and its status is not clear at this time. Distribution of Recent Gerrhosaurus major G. major occurs today in principally arid savanna along the eastern coast of Africa, north to Eritrea and south to Zululand. G.m. grandis, the most southern subspecies, is found from Zulu- land north to Morogoro, Tanganyika. G. m. major is a coastal 1962 MIOCENE GERRHOSAUR FROM KENYA " subspecies, principally in Tanganyika, but reaching as far north as Kenya. G. m. bottegoi ranges from central Tanganyika north through central Kenya and reaches north to coastal Eritrea, far- ther north than any other gerrhosaur, and is the only subspecies of G. major found today in the Kavirondo Gulf region of Lake Victoria, the same region as the occurrence of the fossil. The problematical G. m. zechi has, so far as known, a disjunct distri- bution limited to the northern Belgian Congo and Togo. CONCLUSIONS The fossil described here is closely related to, and perhaps conspecific with, the Recent species Gerrhosaurus major. Thus it is extremely probable that the habitat of the lizards repre- sented by the fossil was semi-arid or arid savanna, like that of the modern species. The presence of a mammalan fauna of Ay Fig. 3. Gerrhosaurus ef. G. major, Lower Miocene, Kenya, ventral view, dominantly savanna aspect in these deposits supports this con- clusion. Whitworth (1953, p. 82) states that ephemeral lakes were probably present in this region in the late Miocene, allow- ing the savanna mammals access during the periods of desicca- tion indicated by the sediments. He also points out that similar ioe) BREVIORA No. 158 situations occur today in areas of the northwestern Sudan. Ches- ters (1957), on the basis of the flora, has concluded that a ‘‘egal- lery-type forest in which trees festooned with climbers overhung the watercourses’’ lived near to the site of deposition, and that many of the fossil nuts and seeds represent living tropical Af- rican genera. Close relationship of the fossil with the Recent species indi- cates that at least part of the pattern of speciation seen within the genus today is of considerable antiquity. Moreover, similari- ties of the fossil to some of the Recent subspecies of G. major perhaps indicate that some of the geographic variants seen today were beginning to appear, as far back as the Miocene. The two Recent subspecies which the fossil most closely resembles, G. m. bottegot and G. m. grandis, are northern and southern popula- tions which intergrade in the area immediately south and east of Lake Victoria. It is interesting, but highly speculative, to suggest that the occurrence of the fossil near the present area of intergradation of these two Recent subspecies might either indi- cate a stage in the development of the Recent subspecifie pat- terns, or an intergrade between the two which could possibly be duplicated today if sufficient specimens were available. Moreau (1951, esp. pp. 877, 881) has gathered evidence which suggests that at least from the mid-Cenozoic to the present, cen- tral and eastern Africa had a climate and broadly defined vege- tational types which differed relatively little from those occurring there today. If this is so, the above alternatives are quite pos- sible, yet additional fossil evidence, both biotic and climatic, is necessary to accept or reject either of them. SUMMARY The well-preserved head of a fossil lizard from the Lower Miocene (Burdigalian?) of Mfanganu Island, Lake Victoria, Kenya, is tentatively referred to the Recent species Gerrhosaurus major (Reptilia: Cordylidae). Many external features of soft anatomy are preserved as casts in calcite, including the eye, tongue, and tympanic membrane. Close relationship to the Re- cent semi-arid or arid savanna species indicates a similar habitat for the fossil, a conclusion corroborated by the savanna aspect of the fossil mammals from contemporaneous deposits on nearby Rusinga Island. This specimen is one of the oldest vertebrate fossils even tentatively referred to a Recent species and must in- dicate that at least part of the pattern of speciation seen in Recent gerrhosaurs is of relatively ancient origin. 1962 MIOCENE GERRHOSAUR FROM KENYA 9 LITERATURE CITED CHESTERS, K. I. M. 1957. The Miocene flora of Rusinga Island, Lake Vietoria, Kenya. Palaeontogr., Abt. B, 101:30-71, 4 figs., 21 pls. Dr STEFANO, G. 1903. I sauri del Quercy appartenenti alla collezione Rossignol. Atti Soe. Ital. Sci. Nat., 42:382-418, 2 pls. FILHOL, H. 1877. Recherches sur les phosphorites du Quercy. Etude des fossiles qu’on y rencontre et spécialement des mammiféres. Ann. Sei Géol., 8:1-561, 55 pls. FirzSimons, V. F. 1943. The lizards of South Africa. Transvaal Mus., Mem. no. 1, xv + 528 pp., 384 figs., 24 pls. HorrstTETTER, R. 1944. Sur les Seincidae fossiles. I. Formes européennes et nord- americaines. Bull. Mus. Nat. Hist. Nat. Paris, 16:547-553, 2 figs. = ce) il | Squamates du type moderne. Jn: Piveteau, Traité de Paléon- tologie, 5:606-662, 26 figs. Leakey, L. S. B. 1952. Lower Miocene invertebrates from Kenya. Nature, 169:624- 625, 2 figs. LeGROS CLARK, W. E., AND LEAKEY, L. S. B. 1951. The Miocene Hominoidea of East Africa. Brit. Mus. Nat. Hist., Fossil Mammals of Africa, no. 1:1-117, 28 figs., 9 pls. LOVERIDGE, A. 1942. Revision of the African lizards of the family Gerrhosauridae. Bull. Mus. Comp. Zool., Harvard Univ., 89:484-543. McDoweELL, S. B., JR., AND BoGert, C. M. 1954. The systematic position of Lanthanotus and the affinities of the anguinomorphan lizards. Bull. Amer. Mus. Nat. Hist., 105:1- 142, 43 figs., 16 pls. Moreau, R. E. 1951. Africa since the Mesozoic, with particular reference to certain biological problems. Proc. Zool. Soc. London: 121:869-9135, 4 tables. Romer, A. 8. 1956. Osteology of the Reptiles. Univ. Chicago Press, xxi + 772 pp., 248 figs. WHITWorRTH, T. 1953. A contribution to the geology of Rusinga Island, Kenya. Quart. Jour. Geol. Soc. London, 109:75-96, 2 pls. 1958. Miocene ruminants of East Africa. Brit. Mus. Nat. Hist., Fossil Mammals of Africa, no. 15:1-50, 18 figs., 12 tables. 158 No. BREVIORA 10 JO oUTTINO ATLA “APISIOATUL) UOJSOG ‘pavuAVI_Y YOUopoty AG YdBisoj,oyd “poyonojat ATYSI[TS “e[Bos 07 JON “souTpJNoO o[vos podtosoid-[fam pure fonsuo0, surpnajor1d ATJYSIS *VoW10) pue sptjaco \dSo OJON “AT[VSAOP poysnayd [NYG “MOTTA [RAayVT 4Jop JO YdRasojyoyq “wolMw “4H “Jo snanpvsoy.sa+y L 81d O BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. May 31, 1962 NuMBER 159 AGE IN A SMALL SAMPLE OF BLUEFISH (POMATOMUS SALTATRIX (LINNAEUS))! By RicHarp H. Backus Woods Hole Oceanographic Institution and Museum of Comparative Zoology Little is known about the life-history of the bluefish (Poma- tomus saltatrix) even though this long-known species is widely distributed in warm and temperate seas and is valuable for food and sport. Not only have the eges and larval youne of this interesting fish not been positively identified, but nothing exact is known of its rate of growth although the ‘‘annuli’’ on its scales prove to be quite easily read. Perhaps it is the great population fluctuations of this fish and its erratic appearance in many waters which have discouraged naturalists from study- ing it. Bigelow and Schroeder (1953) give a summary of what is known of bluefish life history. Little has been added since their account was written. To learn something of the growth of the species, advantage was taken of the abundance of bluefish around Woods Hole dur- ing the autumn of 1961 and of the angling expertness of Asa Wing, Henry Cain, and Carl Grant, Jr. These men saved heads and scale samples together with records of fork length from the 34 fish that they caught between October 9 and 19 in Great Harbor at Woods Hole, in Woods Hole passage, and along the east shore of Buzzards Bay north of Woods Hole. The saccular otolith of the bluefish has been figured by Le Gall (1934) and by Sanz Echeverria (1950). Le Gall (op. cit.) also figured the scales, and in another paper (1935) he said that his examination of the scales in North African specimens suggests that ‘‘young individuals attain their adult size and their first sexual maturity at the age of 4 or 5 years.’’ On the other hand, 1 Woods Hole Oceanographic Institution Contribution No. 1234 bo BREVIORA No. 159 Borcea (1936) says of Black Sea bluefish: ‘‘During the second year they reach dimensions of 14-20 em and can attain the first sexual maturity.”’ TABLE | Fork length and age in a small sample of bluefish. Tork Length Age Fork Length Age (inches) (annuli) (inches) (annuli) 18) ISP 15.2 EF 14.0 ila= 19.2 lie 14.0 lee 15.5 tse 14.0 Ise 15.5 [Ee 14.0 4 (ke oie UEP) bea) [Se 14.1 2 UES oe WES) 15.5 T+ 14.1 IAF 15.8 ict 14.2 I+ 16.0 le 14.2 ae 16.2 Mee 14.4 ict 16.5 lier 14.5 ict IAD) SF 14.5 I+ 15 iss 14.8 I+ 18.0 NE 15.0 fet 19.0 ee 15.0 lO 19:0 nee 15.0 Y (Clse Ge IUI-=)) 20.1 IDIGF hel Se 24.0) UOIe= An otolith extracted from a specimen in our sample 14.4 inches in fork length measured about 0.45 by 0.15 inches. The otolith is much sculptured and does not seem suitable for use in age de- termination by simple visual inspection though it might be so suited if x-ray methods were used. Since the scales of this fish are large, and those examined by us seem to show the annul rather clearly (Figure 1), we have relied on the scales alone for the ages reported here. Fork leneth and age for the specimens in our sample are given in Table 1. Fork lengths were taken to the nearest eighth of an inch, were converted to decimals and rounded to the nearest tenth. Ages (expressed in annuli) were determined by examin- ing at least 10 legible scales from each specimen. In three cases consistent results could not be obtained. Thus, age has been deter- mined with some confidence in 31 instances. Of these, 21 speci- mens show an age of I+, nine an age of II+, and one an age of 1962 AGE IN BLUEFISH 3 III+. Specimens of age I+ range from 13.9 to 16.5 inches in fork length and have a mean length of 14.85 inches. Specimens of age II+ range from 15.0 to 20.1 inches and have a mean leneth of 17.48 inches. The sole II[I+ specimen measures 24.0 inches in fork length. Because of the small size of our sample these data ean do little more than show that snappers (as small bluefish are called ) of four to nine inches, seen in the autumn (Bigelow and Schroe- der, 1953), are indeed young of the year and that such fish ap- proximately double their length in the succeeding 12 months. Bluefish left the Woods Hole area during the course of a five- day northeast storm which commenced on the evening of October 19 and lasted until the early hours of October 25. The last fish in our sample was caught on October 19. None were caught after the passage of the storm although the fishing effort continued and produced good catches of striped bass (Roccus sazxatilis). Surface temperature records for Great Harbor, Woods Hole, show fluctuations between 64.9 and 64.5°F for the period October 6 to 12, a decline from 64.5 to 60.8°F from October 12 to October 19, the day the storm began, a decline from 60.8 to 55.0°F during the storm, and small fluctuations about 55°F for the remainder of the month. One may suppose that at about 60°F the bluefish were near the minimum temperature that they can tolerate at this stage of the life cycle and that the catastrophe of the storm, with the accompanying further drop in tempera- ture, was enough to start them on the journey to their winter haunts. LITERATURE CITED BicrLow, Henry B. and WILLIAM C, SCHROEDER 1953. Fishes of the Gulf of Maine. U.S. Fish and Wildlife Service Fishery Bulletin, 74 (Volume 53): 1-577. Borceta, I. 1936. Notes sur la Biologie du Pomatome (Lufar) de la Mer Noire. C. R. Acad. Sci. Roumanie, 1: 222-223. Lr GALL, JEAN 1934. Le Tassergall ou Bluefish (Pomatomus saltatriz Lacépéde = Temnodon saltator Linné). Rey. Tray. Office Péches Maritimes, 2 PAT (=hsyay- 1935. Le Tassergall ou Blue Fish. Bull. Soc. Sci. Nat. Maroc, 15: 239-933 Sanz ECHEVERRIA, JOSEFA 1950. Notas sobre otolitos de peces procedentes de las costas del Sahara. Bol. Inst. Espafiol Ocean., 27: 1-14. 4 BREVIORA No. 159 hee ANNULUS I Fig. 1. Scale from a bluefish (Pomatomus saltatriz) 20.1 inches in fork length taken in Woods Hole passage on October 17, 1961, showing two annuli. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. JUNE 12, 1962 NuMBER 160 TWO NEW ARTHROPOD CARAPACES FROM THE BURGESS SHALE (MIDDLE CAMBRIAN) OF CANADA By W..D. [An Roure Three carapaces of an undescribed arthropod were found by the writer when curating the large Museum of Comparative Zoology collection of Burgess Shale arthropods, collected by P. EK. Raymond, H. C. Stetson, W. E. Schevill and C. H. Burgess in August, 1930. Subsequent search through the material in the U. S. National Museum, Washington, D. C., yielded eleven further specimens and two specimens of another new form, which had been set aside for description by C. E. Resser. The writer is indebted to Dr. G. A. Cooper for permission to borrow and describe the U.S.N.M. material and to Dr. H. B. Whitting- ton for the photographs and for criticism of the manuscript. The USNM specimens came from Walecott’s quarry at locality 35k near Field, British Columbia (Walcott, 1911, pp. 51-52; Resser, 1929, p. 2; = locality S11f of Rasetti, 1951, pp. -37-38, 103, 129). The MCZ specimens were also probably collected from this quarry, although it is impossible to be certain of this since the 1980 expedition also collected from ‘‘a second layer . very fossiliferous . . . some seventy feet further up the mountainside’’ (Raymond, 1930, p. 32; 1935, p. 205). This second locality possibly corresponds with Rasetti’s Sllg (1951, pp. 38, 104, 130) and details of the stratigraphy of these two horizons are given in that work. Letters a and b following a specimen number indicate that part and counterpart are present. Carapace shape alone is insufficient to determine the affinities of any arthropod, as Roger has pointed out (1946, p. 59), and hence it seems better to group such isolated carapaces together as follows. 2 BREVIORA No. 160 TRILOBITOIDEA or CRUSTACEA incertae sedis PROBOSCICARIS gen. nov. Type species. Proboscicaris agnosta sp. nov. Diagnosis. Carapace valves only known, large, with antero- dorsal region produced into a spatulate beak. PROBOSCICARIS AGNOSTA Sp. Noy. Plate 1, figures 1, 2; Text-figure 1 Diagnosis. Anterior beak prominent; length of posterior mar- gin 0.43 to 0.82 of greatest depth of carapace; posterior margin indented at or near midpoint. Description. Valves elongate, ranging from shallow to deep. Since the orientation of the valves is unknown, the straight to she¢htly concave margin will be treated as dorsal and the pro- duced region as anterior. Postero-dorsal angle rounded and obtuse; posterior margin slightly indented at or near its mid- point. Ventral margin moderately convex in posterior half to three-quarters of carapace length; strongly concave in anterior region and separating off a tongue-shaped anterior beak. This beak has been lost from USNM 139866 (see Text-fig. 1) and it seems likely that the distinct outlines shown by USNM 139869 and 139873 (Text-fig. 1) are only due to the loss of this region. The absence of a recognizable rim or doublure pre- vents certainty on this point. The valves were doubtless jomed by a membranous hinge along the dorsal border as in Canadaspis. Only one specimen shows evidence of two valves preserved, and this is shown on Text-figure 1, USNM 1389872. Only the anterior beak of the right valve is preserved and this is displaced anteriorly rela- tive to the almost complete left valve. A reticulate pattern may be seen in patchy areas on USNM 139867 and 139873. The wrinkles subparallel to the ventral margin of USNM 139866 (Text-fig. 1) are clearly due to flat- tening of the originally convex test. Specimens USNM 139867, 139870, 139871, 13898738, MCZ 5979/2 and MCZ 5979/3 are blotched by the alga Morania parasitica Walcott, previously recorded on Canadaspis and figured by Waleott (1919, p. 232, pl. 50, fig. 1) on a carapace of Hurdia victoria Walcott. ARTHROPODS NEW BURGESS SHALE 1962 ‘TL86EL WNSD OdAJOLOFT “popPVorpul saoquinu Sopeye YIM ‘sou “ds Jo “Was Y]soUsD si.ipaiwsogoig JO Stowtoods UMOUY WodZINOF OY} FO Woop JO SSurmevarp ourpyNO “TL “Py USD Ul 3/D2 OSC SZ E6EI We /6LES OL86LI $Le6Ll ZOW, : XK e986LI 4 BREVIORA No. 160 Remarks. The form of the carapace is so distinct that little confusion is possible with previously described species. Such a small beaked specimen as USNM 139874 approaches Canadaspis perfecta (Walcott) in outline, but the posterior and antero- ventral embayments readily distinguish the new form. Some specimens of Hurdia victoria in the Museum of Comparative Zoology show an indentation of the ?posterior margin similar to that in Proboscicaris, and in addition show an identical retic- ulation of the carapace surface, so that fragments of the pos- terior ends of the two forms might prove difficult to distinguish. Large-mesh reticulation also occurs in Tuzoia and Carnarvonia (Walcott, 1912, pp. 157-158, 165, 187, 189) so that this char- acter is of little value for suggesting relationships. Small-mesh reticulation is visible in species of Caryocaris, Dictyocaris and Concavicaris (as well as in the olenellid trilobites: Raw, 1936; Moore, 1958, fig. 5.22), and in Ceratiocaris and Montecaris such reticulation can be shown to arise from differential corrosion of the cuticular prisms (Rolfe, 1962a, pp. 45-47). The cuticle of the Burgess Shale specimens is too poorly preserved to ascertain whether this reticulation is sculptural or structural. Holotype. USNM 139871. Plate 1, figure 2 and Text-figure 1. Other material. The twelve specimens shown on Text-figure 1: USNM 139866a/b, 139867a/b, 139868-139870, 139872-139875. MCZ 5979/1, 5979/2a/b, 5979/3a/b. Another specimen, USNM 139876, is a fragment of the anterior end only. Dimensions of holotype. Maximum length parallel to hinge line: 98 mm. Maximum depth perpendicular to hinge line: o2 mm. PROBOSCICARIS INGENS sp. nov. Plate 1, figure 3; Text-figure 2 Diagnosis. Carapace valves only known; anterior beak rela- tively small; length of posterior margin ca. 0.28 of greatest depth of carapace; posterior margin sigmoidal. Description. The ventral margin is more of a simple skewed curve than the convex and coneave outline of P. agnosta. The posterior margin is shorter and hence is situated more dorsally than in P. agnosta and in addition this margin is sigmoidal rather than indented. The carapace margin is curled under, except alone the dorsal border, suggesting the marginal rim common in the later phyllocarids. 1962 NEW BURGESS SHALE ARTHROPODS 5 The surface of the carapace is smooth but little is preserved of the original test save blotches of filmy black material. Occa- sional circular areas of silver sheen on the holotype may repre- sent Morania parasitica. Fig. 2. Outline drawing of the holotype of Proboscicaris ingens sp. nov. — USNM 1389865b. Postulated anterior to left. Remarks. Again the carapace shape is distinetive, though the asymmetrical outline recalls that of Jsoxrys, which, however, is smaller and has the anterior and posterior dorsal extremities acuminate rather than truneate. This species is the largest of the known Burgess Shale arthro- pods in terms of surface area of the carapace and was doubtless the one which Walcott had in mind when he wrote, ‘‘there are also fragments of the carapace of a very large form that possibly may be related to Hurdia victoria’’ (1912, p. 183). Individuals of H. victoria may be longer but they are also slenderer. Such large carapaces are of particular interest in this fauna since it is among them that a suitable adult for the hypothetically larval Waptia fieldensis might be sought (Fedotov, 1925, pp. 386, 389; Heldt, 1954, p. 180; Tiegs and Manton, 1958, pp. 292, 314; cf. Henriksen, 1928, p. 14; Stormer, 1944, p. 100). In this connection is seems worth recalling the striking resemblance of Marria, from this same deposit, to a erustacean nauplius (Ruedemann, 1931, p. 8) or metanauplius. A comparable Upper Ordovician form, Paramarria, occurs in association with an arch- aeostracan carapace, Galenocaris (Wells, 1944). If Paramarria is a nauplhiar stage, and the larval stage of Galenocaris, and its 6 BREVIORA No. 160 aspect 1s not merely due to convergence for a planktonic exist- ence (as that of Mimetaster and Bostrichopus seems to be), it would contrast with Recent Leptostraca. In the latter group, development is direct, the young hatching at a late stage. Simi- larly, Naraoia might be regarded as a larval merostomoid. It is possible that P. ingens is simply an older instar of P. agnosta. However, such radical changes in shape are not ecom- mon except in early ontogeny, and it seems better to distinguish this form as a separate species. Holotype. USNM 139865a/b — part and counterpart. Dimensions of holotype. Maximum leneth parallel to hinge line: 156 mm. Maximum depth perpendicular to hinge line: 75 mm. Other material: USNM 139890 —a fragment of the posterior of a carapace valve, which must have exceeded 150 mm. long by 95 mm. deep when complete. DISCUSSION The lack of Limbs or body segments precludes any discussion of the affinities of this new genus and the problem of classifica- tion of these early crustacean-trilobitoid forms has been sum- marized elsewhere (Rolfe, 1962b). It seems worthless to classify such isolated carapaces above the generic level in view of the limited number of characters available. Many of the genera attributed to phyllocarid families, or made the types of new famihes such as the Isoxyidae (junior synonym of Tuzoiudae Raymond, 1935) and Pseudoarctolepididae of Brooks and Caster (1956, p. 18), will need to be brought together under the incertae sedis category shown above. Some idea of the relative abundance of Proboscicaris agnosta in the Burgess Shale fauna may be gained from the following list of numbers of individuals of non-trilobite arthropods col- lected by the 1930 MCZ expedition, and recently curated by the writer: TRILOBITOIDEA Burgessia bella Walcott 54 Emeraldella or ?Molaria spp. indet. 3 Leanchoilia superlata Wale. 12 Marrella splendens Wale. 202 Naraova compacta Wale. 3 1962 NEW BURGESS SHALE ARTHROPODS if Opabinia regalis Wale. 1 Sidneyia inexpectans Wale. 9 ?Yohoia plena Wale. 1 TRILOBITOIDEA or CRUSTACEA incertae sedis Anomalocaris canadensis Whiteaves ca. 22 Canadaspis obliqua (Wale. ) 7 C. ovalis (Wale. ) 1 C. perfecta ( Wale.) 76 C. sp. indet. 12 Fieldia lanceolata Wale. ] Hurdia triangulata Wale. ] H. victoria Wale. 16 Tsoxys acutangulus Wale. 10 Proboscicaris agnosta sp. nov. 3 Protocaris ef. pretiosa Resser 1 Tuzova retifera Wale. i T. sp. indet. 2 REFERENCES Brooks, H. K. and K. E. CASTER 1956. Pseudoarctolepis sharpi, n. gen., n. sp. (Phyllocarida) from the Wheeler Shale (Middle Cambrian) of Utah. Jour. Paleont., 30 :9-14. Frpotoy, D. 1925. On the relations between the Crustacea, Trilobita, Merostomata and Arachnida. Akad. Nauk S.S.S.R., Leningrad, Bull., VI Série, 18:383-408. HeEwpt, J. H. 1954. Waptia fieldensis Walcott et les stades larvaires des pénéides. Bull. Soe. Sei. nat. Tunis., 6: 177-180. HENRIKSEN, K. L. 1928. Critical notes upon some Cambrian arthropods described by Charles D. Walcott. Vidensk. Medd. fra Dansk naturhist. Fore- ning i Kebenhayn, 86:1-20. Moors, R. C. 1958. Introduction to Historical Geology. 2nd ed. McGraw-Hill, New York, Toronto, London, 7 + 656 pp. RASETTI, FRANGO 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Misc. Coll., 116, No. 5: 5 + 277 pp. 8 BREVIORA No. 160 Raw, FRANK 1936. Mesonacidae of Comley in Shropshire, with a discussion of classification within the family. Quart. Jour. geol. Soc. London, 92 :236-293. RAYMOND, P. E. 1930. Report on invertebrate paleontology. Ann. Rep. Mus. Comp. Zool. for 1929-1930:31-83. 1935. Leanchoilia and other Mid-Cambrian Arthropoda. Bull. Mus. Comp. Zool., 76:205-230. ResseEpr, C. E. 1929. New Lower and Middle Cambrian Crustacea. Proce. U.S. Nat. Mus., 76, Art. 9, 18 pp. ROGER, JEAN 1946. Les invertébrés des couches & poissons du Crétacé Supérieur du Liban. Mém. Soe. géol. France, Nouvelle Série, 23, No. 51, 92 pp. Roure, W. D. I. 1962a. The cuticle of some Middle Silurian ceratiocaridid Crustacea from Lanarkshire, Scotland. Palaeontology, 5:30-51. 1962b. Grosser morphology of the Scottish Silurian phylloearid crus- tacean, Ceratiocaris papilio Salter in Murchison. Jour. Paleont. (in press ). RUEDEMANN, RUDOLF 1931. Some new Middle Cambrian fossils from British Columbia. Proe. U.S. Nat. Mus., 79, Art. 27, 18 pp. ST6RMER, LEIF 1944. On the relationships and phylogeny of fossil and recent Arachno- morpha, Skr. Norske VidenskapsAkad. Oslo, I, Math.-Kl., No. 5, 158 pp. Trees, O. W. and 8. M. MAnTON 1958. The evolution of the Arthropoda. Biol. Rey., 33:255-337. Watcort, C. D. 1911. Cambrian geology and paleontology, II. No. 38 — Middle Cam- brian holeothurians and medusae. Smithsonian Mise. Coll., 57:41- 68, 1912. Cambrian geology and paleontology, II. No. 6— Middle Cam- brian Branchiopoda, Malacostraca, Trilobita, and Merostomata. Smithsonian Mise. Coll., 57:145-228. 1919. Cambrian geology and paleontology, IV. No. 5 brian Algae, Smithsonian Mise. Coll., 67:217-260. WELLS, J. W. 1944. Two new planktonic crustaceans from the Maquoketa Shale (Ordovician of Illinois). Amer. Jour. Sci., 242 :436-441, Middle Cam- 1962 NEW BURGESS SHALE ARTHROPODS Plate 1 The scale beneath the Figures represents five centimeters. Figs. 1-2. Proboscicaris agnosta gen. et sp. nov. 1. Left valve and dis- placed anterior beak of right valve of carapace. USNM 139872. 2. Holo- type — USNM 139871 with anterior of valve at right. The silver blotches are the enerusting alga Morania parasitica Walcott. 3. Proboscicaris ingens sp. nov., holotype — USNM 139865b. Postulated anterior to left. From an ad BREVIORA Museum of Comparative Zoology * CAMBRIDGE, Mass. JuLy 16, 1962 NuMBER 161 A COMPARATIVE STUDY OF THE RESPIRATORY MUSCLES IN CHELONIA By R. V. SHAH Division of Comparative Anatomy and Embryology, Department of Zoology, M.S. University of Baroda Baroda, India INTRODUCTION The unique skeletal modifications in Chelonia have greatly influenced the morphological features of the soft parts in these animals. Of these modifications the rigid shell formation is the most prominent characteristic feature of the Chelonia; this has made their body wall immovable and hence the normal respira- tory movements characteristic of amniotes are lost. This loss of active body wall movements has given much reason for dis- cussion about the respiratory mechanism adapted by the chel- onians, and from the time of Malpighi and Cuvier many attempts have been made to explain the phenomenon. Some have sug- gested that the throat movements similar to those seen in frogs are responsible for bringing about the expirations and inspira- tions in Chelonia; others have supposed that the movements of the limbs and the neck indirectly effect the expiration and inspiration. Recently McCutcheon (1943) has summarized the evidence that, on the contrary. the movements of certain abdominal muscles, the diaphragmaticus,! the transversus ab- dominis, the serratus magnus and the obliquus abdominis, bring 1A matter of terminology must be mentioned here. McCutcheon (1948) de- seribed the flank cavity muscles in Walaclemys centrata where the diaphragmati- cus muscle as described by him seems to be the same as the muscularis striatum pulmonale and quite different from the diaphragmaticus as described in this paper. Owen (1866), deseribing the musculature of Emys europea, describes the diaphragmaticus as formed of three parts originating from the carapace; two parts insert on the wall of the lung while the third one inserts on the plastron. This description compares well with the account given here, except that the two parts of the muscle described by Owen as inserting on the lung are here regarded as the muscularis striatum pulmonale, and only the third part which inserts on the plastron is the true diaphragmaticus. 2 BREVIORA No. 161 about expiration and inspiration. He interprets the throat move- ments in Chelonia as functioning in olfaction and not in respira- tion. George and Shah (1954) have studied the respiratory mechanism in Lissemys and have confirmed MeCutcheon’s view that the abdominal muscles are effective for respiratory move- ments and that the throat movements are only for olfaction. In addition, they have also described the presence of an extra pair of muscles which cover the lungs completely in Lissenys. These muscles are composed of striated muscle fibres. On con- traction of these muscles the pulmonary air is pushed out of the lungs, and on their relaxation the atmospheric air is taken in. Thus these lung muscles, the muscularis striatum pulmonale, aid the action of the abdominal muscles in bringing about the expiration and inspiration. Fig. 1. Diagrammatic sketch of the disposition of the respiratory muscles in Cyclanorbinae where the lungs are completely covered by the muscularis striatum pulmonale. George and Shah (1955, 1958 and 1959) made a compara- tive study of the abdominal muscles and of the lung muscles in some additional chelonians: Lissemys punctata (all three 1962 RESPIRATORY MUSCLES IN CHELONIA 3 subspecies), Geoemyda trijuga, Trionyx gangeticus, Testudo ele- gans, Malacochersus tornerti and Eretmochelys imbricata. Ac- cording to their observations the lung muscle, the muscularis striatum pulmonale, covers the lungs completely in Lissemys punctata, partially in Geoemyda, while the muscle is totally absent in the rest of the forms they studied. Of the flank cavity muscles the diaphragmaticus and the transversus abdominis are Fig. 2. Diagrammatic sketch of the disposition of the respiratory muscles in Trionychinae. well developed in Lissemys and Trionyx where they join with each other to form a continuous muscle sheath covering the visceral organs including lungs. The diaphragmaticus in Geo- emyda trijuga and Eretmochelys imbricata does not join with the transversus abdominis to form a continuous muscle sheath, but there is a bridge of connective tissue between them. In Testudo elegans and Malacochersus tornert the diaphragmaticus muscle is totally absent leaving only a thin membranous sheath of connective tissue in its place. The other flank cavity muscles, viz. the serratus magnus and the obliquus abdominis are present with sight variation in all the animals they studied. In the light of these observations on the respiratory muscles in a very few chelonians, it was thought desirable to examine 4 BREVIORA No. 161 Fig. 3. Diagrammatic sketch of the disposition of the respiratory muscles in Malaclemys terrapin terrapin, Ag Vs LLL. Fig. 4. Diagrammatic sketch of the disposition of the respiratory muscles in Pseudemys floridana and Pseudemys texana. 1962 RESPIRATORY MUSCLES IN CHELONIA 5) more forms representing, as far as possible, almost all the major groups of the order Chelonia, and make a comprehensive com- parative study of these muscles to get an overall idea of the morphological features of the respiratory mechanism adapted by the animals of this order. For this study some fifty different cryptodiran and nine pleurodiran forms were selected. A list of the animals chosen is given below. This work was carried out at the Museum of Comparative Zoology at Harvard University, Cambridge, Massachusetts, U.S.A. I am thankful to Dr. A. 8S. Romer, then Director of the Museum, and Dr. E. E. Williams, Curator of Herpetology, for all the facilities given and for their constant help and en- couragement during the course of the study. M.D- Fig. 5. Diagrammatic sketch of the disposition of the respiratory muscles in Emydinae forms in which the diaphragmaticus muscle is absent. MATERIAL STUDIED All the animals selected for the study were alcohol preserved and were found in excellent state of preservation. Careful dis- sections of the flank cavity muscles, the diaphragmaticus, the transversus abdominis, the serratus magnus and the obliquus 6 No. 161 abdominis and the lung muscle, the muscularis striatum pulmon- ale, were done on these animals and following is the report of the comparative study. List of chelonians selected for the present study: CRYPTODIRA TESTUDINOIDEA TESTUDINIDAE Emydinae Chinemys reevesii Chrysemys picta dorsalis Chrysemys picta marginata Chrysemys picta picta Clemmys caspica caspiea Clemmys caspica leprosa Clemmys guttata Clemmys mutica Cuora amboinensis Deirochelys reticularia Emydoidea blandingii Emys orbicularis Geoemyda manni Geoemyda punctularia funerea Geoemyda spinosa Geoemyda trijuga Graptemys kohni Graptemys pseudogeographica Kachuga tectum tectum Malaclemys terrapin terrapin Malayemys subtrijuga Ocadia sinensis Pseudemys floridana Pseudemys texana Terrapene yucatana Testudininae Geochelone pardalis Pyxis arachnoides Testudo graeca Testudo hermanni Testudo horsfieldii Testudo kleinmanni CHELYDRIDAE Kinosterninae Sternotherus carinatus minor Sternotherus odoratus Chelydrinae Chelydra serpentina TRION YCHOIDEA TRION YCHIDAE Cyclanor binae Cyelanorbis sp. Cycloderma frenatum Lissemys punctata (all three subspecies) Trionychinae Dogania subplana Trionyx gangeticus Trionyx sinensis Trionyx triunguis CHELONOIDEA CHELONIDAE Caretta caretta Chelonia mydas Kretmochelys imbricata Lepidochelys olivacea DERMOCHELYOIDEA DERMOCHELYIDAE Dermochelys coriacea PLEURODIRA PELOMEDUSIDAE Pelomedusa subrufa subrufa Pelusios subniger Podocnemis expansa Podocnemis lewyana Podocnemis unifilis CHELIDAE Chelodina longicollis Emydura krefti Hydromedusa maximiliani Phrynops geoffroana Platemys platycephala 1962 RESPIRATORY MUSCLES IN CHELONIA ff DESCRIPTION OF THE MUSCLES Of all the respiratory muscles mentioned above, the lung muscle, the muscularis striatum pulmonale, shows the most ex- treme variation in Chelonia. It is so well developed in the forms belonging to the subfamily Cyclanorbinae that it covers the lung completely while in the other subfamily, Trionychinae, the muscle is totally absent. In Cyclanorbis sp. and Cycloderma frenatum (Fig. 1) the muscle arises from the carapace in the vicinity of the second and third thoracic vertebrae and also from the lateral side of these vertebrae. The fibres arising from the carapace run over the entire dorsal surface of the lung and when they Fig. 6. Diagrammatic sketch of the disposition of the respiratory muscles in Kachuga tectum. reach the outer, anterior, and the posterior limits of the lung they turn onto the ventral side and continue to run towards the entrance of the bronchus. The fibres arising from the lateral side of the thoracic vertebrae run over the medial side of the lung and then come onto the ventral side and reach the entrance of the bronchus. All the fibres of the muscle closely adhere to the wall of the lung. From the place of the origin of the fibres, the muscle could be arbitrarily divided into two parts, a lateral 8 BREVIORA No. 161 part which arises from the carapace and a medial part which arises from the lateral side of the thoracic vertebrae. The mas- cularis striatum pulmonale in Lissemys punctata (Fig. 1) (George and Shah, 1954) differs slightly from the one in Cycla- norbis and Cycloderma; the muscle is otherwise very similar in its course and insertion in all the three genera of Cyclanorbinae. In Lissemys the muscle arises entirely from the carapace and does not have its lateral part arising from the side of the verte- brae. The muscle in all the forms of the group Cyclanorbinae is Gales: Fig. 7. Diagrammatic sketch of the disposition of the respiratory muscles in marine turtles. innervated by the branches of the intercostal nerves. As said before, the muscle on its contraction pushes out the pulmonary air of the lungs, and on its relaxation the atmospheric air rushes in. In Trionyx gangeticus, Trionyx sinensis, Trionyx triunguis and Dogania subplana (Fig. 2) belonging to Trionychinae, the muscularis striatum pulmonale muscles is totally absent. All the forms belonging to the Emydinae have a muscularis striatum pulmonale which partly covers the lung. In this group the muscle shows great variation in different species. 1962 RESPIRATORY MUSCLES IN CHELONIA 9 Fig. 8. Diagrammatic sketch of the disposition of the respiratory muscles in Testudininae, except Pyxis arachnoides. Fig. 9. Diagrammatic sketch of the disposition of the respiratory muscles in Pyzxis arachnoides. 10 BREVIORA No. 161 In Malaclemys terrapin terrapin, Clemmys caspica caspica, Clemmys guttata, Graptemys pseudogeographica, Graptemys kohmi, Cuora amboinensis, Pseudemys floridana and Pseudemys texana, the muscularis striatum pulmonale, though only partly covering the lung, is well developed compared to other Emydinae. In these emydines the medial part of the muscle arising from the side of the thoracic vertebrae is well developed and quite extensive, while the lateral part of the muscle with its origin from the carapace, though well developed, is comparatively small Fig. 10. Diagrammatic sketch of the disposition of the respiratory muscles in Pleurodira forms in which the muscularis striatum pulmonale is partial and the diaphragmaticus muscle is absent. in extent. However, in Malaclemys terrapin terrapin, Graptemys pseudogcographica and Graptemys kohni the lateral part of the muscle is comparatively more developed than in the other emy- dines listed above. The place of origin of the lateral part of the muscle in these three forms is parallel to that of the medial part of the muscle (Fig. 3). In Pseudemys floridana and Pseu- demys texana the place of origin of the lateral part of the muscle is perpendicular to that of the medial part of the muscle (Fig. 4). 1962 RESPIRATORY MUSCLES IN CHELONIA AT In Emys orbicularis, Kachuga tectum tectum, Ocadia sinensis, Chrysemys picta picta, Chrysemys picta dorsalis, Chrysemys picta marginata, Deirochelys reticularia, Emydoidea blandingii, Clemmys mutica, Chinemys reevesti, Malayemys subtrijuga, Geo- emyda punctularia funerea (Figs. 5 and 6), Geoemyda manni, Geoemyda spinosa, and Geoemyda trijuga (Fig. 12) the medial part of the muscularis striatum pulmonale is very poorly devel- oped and only covers a very small portion of the anterior medial side of the lung. The lateral part of the muscle is also less developed compared to that of Malaclemys and others and shows variations in its extent, never covering more than a small portion of the anterior and anterolateral side of the lung. ean Br. ge A,\ 4/?. Ext. Fig. 11. Diagrammatic sketch of the disposition of the respiratory muscles in Podocnemis. Six species belonging to the Testudininae, Testudo hermanni, Testudo graeca, Geochelone pardalis, Testudo horsfieldui, Testudo klemmmanni (Fig. 8) and Pyxis arachnoides (Fig. 9) have been examined. In all these except Pyxis arachnoides the muscularis striatum pulmonale is totally absent and a thin sheet of con- nective tissue is present in its place. In Pyxis arachnoides (Fig. 9) there is a poorly developed lateral part of the muscularis striatum pulmonale present, covering only a very small part of the anterior region of the lung. The presence of part of the 11) BREVIORA No. 161 muscle in Pyzxis appears to be a case of an intermediate stage between the typical condition of the Testudininae, on one hand, where the muscle is absent, and that of Emydinae, on the other, in which it is better developed. In Sternotherus odoratus and Sternotherus carinatus minor, belonging to Kinosterninae, the muscularis striatum pulmonale is similar in its origin, course and insertion to that described for the Malaclemys terrapin terrapin (Fig. 3). In Chelydra ser- pentina of the Chelydrinae the muscle is completely absent and instead a thin layer of connective tissue is present in its place. In Chelonia mydas, Caretta caretta, Lepidochelys olivacea and Eretmochelys imbricata (Fig. 7), which all belong to the family Chelonidae, the muscularis striatum pulmonale is totally absent. Even in Dermochelys coriacea (Fig. 7), of the Dermochelyoidea, Fig. 12. Diagrammatic sketch of the disposition of the respiratory muscles in such Emydinae as Geoemyda spinosa, ete. the muscle is absent. Thus, it appears that none of the marine forms possess any lung muscle, and that unlike the land forms or the fresh water ones there is no variation in this regard. All the members belonging to the suborder Pleurodira possess a partial muscularis striatum pulmonale rather similar to that seen in the Emydinae (Fig. 5). The members of the genus 1962 RESPIRATORY MUSCLES IN CHELONIA 13 Podocnemis show an unusual extension of the muscularis stria- tum pulmonale. In Podocnemis unifilis and the other two species of Podocnemis, the medial part of the muscle, after its usual origin, runs forward adhering to the medial wall of the lung and (unlke the normal condition where the fibres terminate on the lung near the entrance of the bronchus) after reaching the anterior limit of the lung continues forward and _ closely adheres to the dorsolateral side of the pericardial membrane. Finally, these fibres insert on the membrane at the level of the anterior side of the auricles (Shah, in press). Such a pericardial extension of the muscularis striatum pulmonale is not found in any of the other pleurodirans that were studied. No trace of such an extension is present in any of the Crypto- dira, nor does there appear to be any previous record of the presence of a striated muscle layer on the pericardium in any vertebrate. FLANK CAVITY MUSCLES The diaphragmaticus and the transversus abdominis form the expiratory set of the flank cavity muscles in Chelonia, while the serratus magnus and the obliquus abdominis form the in- spiratory set (McCutcheon, 1943; George and Shah, 1954, 1958 and 1959). In all Chelonia the transversus abdominis muscle is well de- veloped. In the Cryptodira studied, the transversus abdominis muscle is the most highly developed in the Trionychoidea, and in this group it joins with the anteriorly placed diaphragmaticus muscle of the same side to form a continuous muscular sheath which envelops the visceral organs including the lungs. The muscle arises in all the chelonians from the posterior half of the carapace, but the place of origin is not constant in all forms since great variations in its extent are seen in different individ- uals. In Pseudemys floridana and Pseudemys texana the trans- versus abdominis muscle extends almost up to the level of the apex of the heart on the ventral side (Fig. 4). In no Emydinae, whether the diaphragmaticus muscle is present or not, does the transversus abdominis muscle have the extensive spread seen in all the Trionychoidea. In those Emydinae where the dia- phragmaticus muscle is present, there is a bridge of connective tissue between it and the transversus abdominis of the same side. Such a bridge of connective tissue between the diaphrag- maticus and the transversus abdominis muscle is present in all the species of the genus Geoemyda and all the marine chelonians. 14 BREVIORA No. 161 The diaphragmaticus muscle, in all the chelonians in which it is present, arises from the undersurface of the second or third costal plates of the carapace. Its place of origin is oriented transversely with respect to the vertebral column. In all the forms of Trionyehoidea (Figs. 1 and 2) the muscle is very highly developed and, as mentioned above, it joins the trans- versus abdonunis muscle of its side to form a continuous mus- cular sheath to envelop the viscera. In some Emydinae the muscle is present (Fig. 12); in others it is absent (Figs. 3, 4, 5, and 6). No members of the Testudinae have the diaphrag- maticus (Fig. 8); there is a thin layer of connective tissue in its place. In all the chelonian studies the inspiratory muscles, the ser- ratus magnus and the obliquus abdominis (Figs. 1 to 11), are present with such shght variation that these are not worth detailed discussion. On contraction of these muscles the volume of the body cavity is increased and thus a negative pressure is created in this cavity and so the lungs expand. On expan- sion of the lunes the atmospheric air rushes in and in this way inspiration is brought about. DISCUSSION From the present study of the respiratory muscles in Chelonia it is evident that there is a great deal of noticeable variation in two muscles, the diaphragmaticus and the muscularis striatum pulmonale. The variations in these two muscles range from a highly developed condition to a total absence, with all inter- mediate stages. The other respiratory muscles are always pres- ent, and although slight variations in different forms are seen, these are very minor ones. The presence of the muscularis striatum pulmonale in all the Cyeclanorbinae, where the muscle covers the lung completely, is regarded as a primitive character which is retained in these forms. The early ancestral chelonians presumably developed these muscles as a substitute for the intercostal muscles lost when their body wall was covered by the rigid shell and could not have the normal movements which are the main component of the respiratory mechanism in all other amniotes. The muscu- laris striatum pulmonale must thus have been of survival value to the early ancestral chelonians, and it is retained fully in all the Cyclanorbinae but shows a gradual trend toward total dis- appearance in other chelonians. Some chelonians, the Triony- chinae, Testudinae and all the marine forms, have totally lost 1962 RESPIRATORY MUSCLES IN CHELONIA A5 this muscle. It is quite obvious that the presence of the muscle covering the lung is a hindrance to full expansion of the lungs; it must therefore have developed as a stop gap arrangement to tide over the loss of the body wall movements until some better physiological adaptation for respiration was achieved. Unpub- lished work by the author on the blood of some chelonians shows some interesting results. The oxiphoric capacity of the blood of Inssemys punctata where the muscularis striatum pulmonale covers the lungs completely is much less than that of the blood of Trionyx or Testudo elegans where the muscle is totally absent. The oxiphorie capacity of the blood of Geoemyda trijuga, where the muscularis striatum pulmonale muscle is incompletely de- veloped (Fig. 12), shows intermediate values. Thus from the study of blood some light is thrown on the new physiological adaptations which have taken place, substituting for some of the morphological adaptations of the primitive forms. More work on the physiology of respiration in different chelonians will be necessary for a better understanding of the problem of respiratory mechanism in this order. Some aspects of this are being worked on at present in my laboratory at the University of Baroda. ABBREVIATIONS USED IN FIGURES Br. Bronchus Cu Connective tissue C.T.B. Bridge of connective tissues between diaphragmaticus and the transversus abdominis muscles. D. Diaphragmaticus muscle ist Heart Ln. Lung M.D. Connective tissue in place of the diaphragmaticus muscle. N.S.P. Muscularis striatum pulmonale muscle covering the left lung M.S.P.1 Muscularis striatum pulmonale muscle cut horizontally and the right lung removed so as to show the place of origin of the muscle M.S.P.L. Lateral part of the muscularis striatum pulmonale M.S.P.M. Medial part of the muscularis striatum pulmonale O.A. Obliquus abdominis muscle P.M. Pericardial membrane 12, Wat Pericardial extension of muscularis striatum pulmonale. S.M. Serratus magnus muscle eA: Transversus abdominis muscle 16 BREVIORA No. 161 REFERENCES GEORGE, J. C. and R. V. SHAH 1954. The oceurrence of a striated outer muscular sheath in the lungs of Lissemys punctata granosa Schoepff. J. Anim. Morph. Phys- iol., 1: 13-16. 1955. Respiratory mechanism in Chelonia. J. Anim. Morph. Physiol., 1: 30-32. 1958. The structural basis of the evolution of the respiratory mech- anism in Chelonia. Proe. XVth Int. Congress of Zool. London. Papers read by title: 24: 1-2. 1959. The structural basis of the evolution of the respiratory mech- anism in Chelonia. J. Anim. Morph. Physiol., 1: 1-9. McCutrcHeon, F. H. 1943. The respiratory mechanism in turtles. Physiol. Zool., 16: 255- 269. OwEN, R. 1866. Anatomy of vertebrates. Vol. I. London. xxxvii + 650 pp. WILLIAMS, E. E. and S. B. McDOWELL 1952. The plastron of the soft shelled turtles (Testudinata, Triony- chidae). A new interpretation. J. Morph., 90: 263-275. BREVIORA Museum of Comparative Zoology a CAMBRIDGE, Mass. JuLy 25, 1962 NuMBER 162 AUSTRALIAN CARABID BEETLES X. BEMBIDION By P. J. DARLINGTON, JR. Museum of Comparative Zoology, Cambridge, Mass. This is the tenth in a series of papers on Australian Carabidae. Some earlier parts, including a list of localities at which I col- lected in 1956-1958 and a discussion of transition of wet forest carabid faunas from New Guinea to Tasmania, are given under References. The present paper deals with the Australian species of Bem- bidion (sensu lato). This is a zoogeographically important genus, which tends to be bi-zonal in distribution, occurring mainly in the north and south temperate zones of the world (Darlington 1959, pp. 332-333). The distribution, ecology, relationships, and possible history of the Australian forms are therefore note- worthy and will be summarized after discussion of the separate species. I am indebted to Prof. Carl H. Lindroth for dissecting males of all the Australian species and telling me how he thinks they are related to Kuropean and North American forms. I could have made the dissections myself, but I am not familiar with the genitalic characters of northern Bembidion and could not have interpreted the characters of the Australian species. However, neither Prof. Lindroth nor, I have investigated most of the Asiatic species or those of New Zealand or southern South America. This paper is therefore only a limited contribution to the zoogeography of Bembidion. At the time of Sloane’s last study of Australian Bembidiini (1921), he knew five Australian species of Bembidion and two of Cillenus. Two supposed ‘‘Bembidion’’ of Blackburn’s that Sloane did not know (hobarti and wattsense) are in fact not Bembidion but Tachys. I plan to treat them in my next paper. I have series of all five real Bembidion known to Sloane and 2, BREVIORA No. 162 have seen no other native species, and it may be that these five species of the genus (exeluding Cuillenus) are all that are native in Australia, although it is too soon to be sure about this. References and synonymy of the species are given by Sloane (op. cit.) and will usually not be repeated here. The species should be identifiable by the following key, which is based partly on Sloane’s key (1921, p. 193). All the species are winged and presumably able to fly, except that the wings are dimorphic in proprium (q.v.). Key to Australian Species of BEMBIDION 1. Large (ec. 5.2-6.5 mm.); dull bronze, elytra with 2 incomplete transverse pale fasciae; clypeus with several fine converging grooves on each side Gntroducedstrom South»America)y 4.940 ar AON ats raters brullei — Smaller; not marked as above; (mative) ........... Sele arate 2 2. Frontal sulei long, impressed and converging on clypeus; upper surface of insect dull or shining; elytron with 6 or 7 dorsal striae .......... 3 — Frontal sulei shallow and rather short, not crossing clypeus; upper Sumas, Chodlls lhyanon yak %/ Clonmel pee - 25. .c6e5escce0000cg000- 5 3. Dull brown with vague paler elytral markings; male upper surface microreticulate; elytron with 6 dorsal striae (stria 7 absent or faint) ; lenge thye 6422-48. ef septate) be. cep A age Go aa vou’ ii ; 1 ’ i i _ 0 ra i a oF : 4 i 3 a - ; a 7 an BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. August 22, 1962 NuMBER 164 NOTES ON THE HERPETOLOGY OF HISPANIOLA. 7. NEW MATERIAL OF TWO POORLY KNOWN ANOLES: ANOLIS MONTICOLA SHREVE AND ANOLIS CHRISTOPHE! WILLIAMS. By ERNEST E. WILLIAMS Recent expeditions in Haiti have obtained material of two anoles, one of which (A. monticola Shreve) had been previously recorded on the basis of the unique type specimen, and the other (A. christophec Williams) was known only from type and para- type. Examination of the new material suggests that the two species, though in a number of respects strikingly different, may yet be related. It is, therefore, appropriate to discuss these two species jointly. Comparison is made also with Anolis darlingtoni Coch- ran = A. etheridget new name,! regarded by Cochran (1989, 1941) as alhed to monticola. ANOLIS MONTICOLA A. monticola Shreve 1936 was described from a single male (lacking most of the tail and darkened by formaldehyde) col- lected by P. J. Darlineton in ‘‘the northern and eastern foothills, Massif de La Hotte, 10060-4000 feet, Haiti.”’ No other specimens have been reported since the original description. However, specimens had been collected for the American Museum of Natural History by W. G. Hassler in 1935 in the vicinity of Aux Cayes and Camp Perrin. These, hike the type, are in a dark phase and show only the faintest trace of pattern. In Hassler’s notebook. on the other hand, there 1 Etheridge (unpublished thesis, University of Michigan) has shown that the genus Xiphocercus cannot be retained. Thus, Xiphocercus darlingtoni Cochran 1935 joins the genus Anolis, and Anolis darlingtoni Cochran 1939 must in conse- quence be renamed. I propose. with Miss Cochran’s kind consent, that the later named species be called Anolis etheridgei. 2 BREVIORA No. 164 is an excellent description of colors in life. The American Mu- seum specimens (as well as Hassler’s notes) were examined by Max Hecht some years ago and the material tentatively referred to monticola Shreve. In 1960, A. S. Rand and J. Lazell (collecting in Haiti with the aid of a grant from the American Philosophical Society) re- turned with the first well-preserved specimens of this very beauti- ful small lizard. A. 8S. Rand reports the circumstances under which this mate- rial was collected as follows: ‘The monticola were found along the trail one-half day’s ride above Camp Perrin. The trail there ran along a hillside mostly cut over and grown up to dense bushes and grass. Along this trail were many hendersoni subsp. in the bushes, particularly where they were shaded by occasional trees. I remember also seeing cybotes and one ricordu. ‘“ At one point we came to a small patch of rocks . . . boulders of various sizes, some very large, heaped one on top of another a bit like but less extreme than the boulder heaps in the Panduras Mts. in Puerto Rico. This area had not been cleared and growing up from among these boulders were sizeable trees and much smaller woody vegetation. The result was a dense, heavily shaded patch of bush isolated in an open sunny area. It was in this spot on the trees, bushes, fallen branches among these large boulders that we found the monticola.”’ The available sample of Anolis monticola now consists of the following specimens :! Haiti. Department du Sud. Northern and eastern foothills, Massif de La Hotte, 1000-4000 ft.: MCZ 38296 (type). Mountains 25 miles north of Aux Cayes on Jerenne Road: AMNH 49818, 49845, 50108-9, MCZ 56139 (formerly AMNH 50110). In mouwn- tains on road to Jerenue about 8 miles from Camp Perrin, 2000- 3000 ft.: MCZ 56140 (formerly AMNH 50097). Tombeau Cheval between Camp Perrin and Beaumont : MCZ 62998-63003. Grande Cayemite: MCZ 58026. The last specimen requires special discussion. It is the Kyerdam specimen (formerly MCZ 25483B) discussed by Miss Cochran (1941, p. 179) as a juvenile coelestinus with keeled ventral scales. Though like many Eyerdam specimens poorly 1 Museums from which specimens are cited in this paper are abbreviated as follows: AMNH, American Museum of Natural History; MCZ, Museum of Com- parative Zoology ; UMMZ, University of Michigan Museum of Zoology. Oo 1962 TWO POORLY KNOWN ANOLES preserved and dried, it seems a typical monticola with the char- acteristic swollen middorsals as well as keeled ventrals. It is important in extending the range of monticola to this small island off the north coast of Haiti’s southwestern peninsula. Hassler’s and the more recent collection were all made in a relatively small area in the southeastern foothills of the Massif de La Hotte. However, the type, like the Grande Cayemite speci- men, came from north of the Massif de La Hotte. (Recent evi- dence, still unpublished, indicates that there may be striking differences between the forms to the north and to the south of this ridge.) The new specimens show that the species reaches at least 46 mm snout-vent length. Color notes and sketches are available for A. monticola both from W. G. Hassler and from the Rand-Lazell expedition. Hassler’s color notes are for specific specimens: 1. Male (Field no. 74, now AMNH 49845). General dorsal color Hooker’s Green. Saddles brown green, three in number, narrowest middorsally, one across shoulder, two between fore and hind legs. A light crescent in the temporal region. Throat and belly dark olive green. Legs barred. Tail barred. Eyes Antwerp Blue, sometimes changing to greenish. Edge of orbit yellowish brown. Skin of fan (which is relatively small) blue, seales light and dark green. Occurring also in a dark phase almost without pattern. 2. Female (Field no. 70, AMNH 50097 = MCZ 56140). Back brownish, bounded laterally by a wavy reddish line edged with a dark line. Sides below the dark line olive shading to very light yellowish green. Sides of neck yellowish green. Tail nearly plain brown. Legs reddish brown posteriorly. Belly nearly white. Upper lips bluish. Throat yellowish with some green. I have examined also by courtesy of Mr. Hassler (now of the Fort Worth Children’s Museum, Fort Worth 7, Texas) a color sketch, made by Melville P. Cummin, of the head and neck of a live male from ‘‘mountains on Jeremie road about 30 miles from Cayes, 2000-3000 ft. alt., August 28-29, 1935.’’ This shows very well the blue of the chin and the yellow green ground color of occiput and nape as well as the black white-centered ocelli which occur laterally on the occiput and nape of some male specimens. Hassler’s photographs of a live male and a female contrast the ocelli and the strong banding of the body in the male with the almost patternless female. 4 BREVIORA No. 164 The observations of Rand and Lazell are in excellent agree- ment with those of Hassler. A pencil and crayon sketch from life is included in their field book and Lazell’s description of the same specimen, now MCZ 63004, may be paraphrased as follows: Ground color yellow green, top of head darker. Body banded with velvety black. Upper and lower jaws blue, this color con- tinued as a stripe fading to whitish above shoulder. An area behind the eye and a V-shaped band across the neck ochre yellow. In the black mark on the neck behind the yellow is a long, thin sky-blue spot. Another such blue spot in the black area on the occiput and anterior to it, still within the black area, a larger white spot with a pale pink center. Legs banded, with a brown- ish wash. Venter and fan bright yellowish green, but base of fan with a bluish wash on both scales and skin. Lazell comments (in agreement with Hassler) that the males ean turn very dark after capture, obscuring the markings to an extreme degree, such that only the markings on head and neck are still visible, but the jaws always retain their bluish color and the fan its bright yellow green with a blue wash. (The type was apparently preserved in this phase; no markings at all are visible, but the blue of the chin remains evident. ) All the males in the Rand and Lazell collection have two prominent pairs of ocelli, one on the occiput, one on the nape. In the figured specimen the light spots in the interior ocelli are divided. In the preserved specimens the hght zone between the two pairs of ocelli may be hehter than the ground color or not. (This is one of the areas described by Lazell as ochre yellow in life.) Banding on body and limbs is always conspicuous; on the flanks each dark band is divided by a lighter, narrow, vertical streak. The two females obtained by Rand and Lazell show the lighter mid-dorsal zone bounded by a wavy dark line mentioned by Hassler. Rand, who has described these specimens, cites the color of the middorsal zone as greenish brown. The inward curves of the dark line are gentle, the convex portions of the line are produced into points by larger or smaller spots of color even lighter than the broad middorsal zone. There is blue on the sides of the chin, and the throat is green and the belly whitish in life. ANOLIS CHRISTOPHEI The species A. christophei Williams 1960 was based on two female specimens obtained in the vicinity of La Citadelle of 1962 TWO POORLY KNOWN ANOLES 5) King Christophe, one collected by W. J. Eyerdam for the Mu- seum of Comparative Zoology in 1927, and one collected by W. G. Hassler for the American Museum of Natural History in 1935. A series of this species, including the previously unknown males, has now been secured again in the vicinity of the Cita- delle by Lue and George Whiteman collecting for the Museum of Comparative Zoology as part of a collecting program in Haiti partly supported by N.S.F. grant G 16066. Unfortunately these are without notes on color in life or any record of dewlap color. They are recorded as occurring on walls and on the ground. The type (MCZ 25485) and paratype (AMNH 49736) are now supplemented by the following specimens (all from La Cita- delle) : MCZ 66900-19, UMMZ 122818(5). The color as ascertained from preserved specimens is not very different from that described by Cochran (1941) for the type, as quoted in the original description. Male. Snout mottled. Two dark supraorbita! stripes, neither sharply defined, the posterior stripe coalescent with an are of dark pigment surrounding a light area which encloses the inter- parietal. A scalloped, crescent-shaped transverse dark mark on the occiput. A butterfly-shaped mark middorsally in front of shoulder, light-edged behind. Three similar marks between the fore and hind limbs and above the sacrum, but in each of these the anterior margins are ill defined. Tail indistinctly banded. On each side a hght lne from the ear arching downward to join a dark-edged ght line just above root of arm, continuing halfway along the body. Flanks above and below light line stronely mottled with darker. Limbs mottled hght and dark. Labials above and below darkly pigmented. Venter washed and powdered, with throat and undersides of limbs and tail darker, with some obscure light spotting, least evident on the throat. Dewlap scales white, skin grey. Female. Pattern as far as discernible the same but much more obseure. Throat and limbs more distinctly spotted with white. Even in the male the pattern, while complex and distinct, is not prominent. Near hatchlings show the same pattern especially well-defined, but even in these the contrast of dark and light areas is not really bold or striking. In all cases specimens must be fully immersed in liquid before the pattern becomes at all evident. 6 BREVIORA No. 164 COMPARISON OF A. CHRISTOPHE, A. MONTICOLA AND A. ETHERIDGEI A. monticola, A. christophei and A. etheridgei are all small anoles (44-48 mm snout-vent length) of, as far as known, quite local distribution. (See map.) Only A. etheridget is known to be montane, reaching at least 6000 feet elevation. A. monticola, despite its name, is not known above 3000 feet, and A. christophei is known only from the Cita- delle which is not more than 2000 feet high. All three belong to the set of Hispaniolan anoles that have the ventral scales arranged in transverse rows. They differ from all other Hispaniolan anoles (except the very different giant anole A. ricordw) in having the subocular scales separated from the supralabials by a row of intervening scales. In the other Hispaniolan anoles the suboculars and supralabials are in contact. The three species share other minor details of squamation: 10-14 scales across the snout; 6-7 loreal rows; 6-7 supralabials to center of eye; temporals and supratemporals finely granular ; interparietal smaller than ear: only one sublabial in contact with the infralabials; lamellae under phalanges 11 and i of fourth toe 14-19. They are somewhat similar also in basic color pattern. All have the dorsum transversely marked. In monticola the dorsal Gone bob ace a | DN ireeanethligil Fe | S “& Dia * aes * \ | ) a > = ee ——— : | 3 | o # Anolis monticola AdDnolis christophei $e Anolis etheridgei Figure 1. Map of the distribution of Anolis monticola, A. christophei and A. etheridgei on Hispaniola. ca | 1962 TWO POORLY KNOWN ANOLES marks extend on to the flanks as transverse bands; in christophet and etheridgei they are restricted to the dorsum as saddle-like markings and in etheridgeit rather broken up into mottling.! Monticola is strongly sexually dimorphic in pattern; etheridger weakly so; christophei hardly so at all. In spite of all resemblances, however, we deal with three strongly marked species, almost as distinct from each other as from other Hispaniolan anoles. Table I compares the differences in squamation between the three taxa. As Cochran suggested in the description of darlingtoni (= etheridget), this species seems the closest relative of monticola. Christophei in its lack of the specializations of the other two forms — in particular, in its possession of a well-developed dew- lap — seems the most primitive of the trio. The bright bold patern of male A. monticola would seem prima facie an obvious compensation for the extreme reduction of the dewlap, which is in most species a striking species-recognition mark. We do not have a description of colors in life of A. etheridgei; it would be reasonable to expect a more vivid pattern than appears on the available specimens. We also lack ade- quate information on the color in life of christophei. In regard to the latter species I can only state that the material recently re- ceived and in a state of preservation that normally retains well any vivid pattern seemed nondescript and obscurely patterned until very closely examined. Whether there are any more members of this small sub-group of Hispaniolan anoles will have to be determined by more thor- ough search of the island. A. etheridgei is still known only from four localities in the Cordillera Central of the Dominican Repub- lice (Loma Vieja, Loma Rucilla, Valle Nuevo, Constanza), A. monticola from four localities in the foothills of the Massif de La Hotte and on the island of Grand Cayemite, and A. christopher only from the single locality, the vicinity of La Citadelle. These are widely separated areas. If related species are equally local in distribution, they may well have been missed. Of course, we lack at the moment any information which would permit sig- nificant hypotheses on the history of these forms. There is indeed evidence that suggests that these three forms are one branch of an extremely interesting intra-Hispaniolan radiation; this con- ception, however, will be documented in another paper. 11 rely on Miss Cochran’s description of A. darlingtoni = etheridgei for evi- dence of transverse, dorsal, saddle-like markings in this form. At the time of writing twenty years later they are not at all evident on the specimens. No. 164 BREVIORA IvpMooevidns 10T19}UL [[jvus AIOA = ‘vd [eyUoIyZ 10T10}80d 1ot1943s80d uvyy 19] -[eus ATJAnNIQe 2OU S[VYyUBd IOLIOJUL azIS ut [enbe ‘sepeos ¢ Aq peyeivdos sopditotuies [e}iqiovidns pajaay “Qnuohpxayqns ‘sno -19UUNU UOTSsatdep [BJUOLZ UL SoTBos qNOUS SSO10¥ SoTBos FI-eT vB prsayyza IvpnooVidns 10110} -ue JO azIs &{ ‘po [eyuody 10119}80d 1o1iajsod ueyy Jayjews fAypdnuiqo speyjued 10TI0}Ue Se[bos g-T Aq poyeivdes sopoitotuas pejiqioeidns YZoous ‘puno.s 10 youohpxay, ‘sno -19WUNU WOTsseidap [eJUOIF UT SeTBos qNOUS SSO1DB SoTvoS ZL-OL 2aydo)sr.yo uorewuenbs jo uostiedwo0g J avy, Ie[noo0ev1dns 10119} “ub oaZzIS %&% ‘po [eyUuoIy «A0t19}8s0d 1ot19}sod UvY} Io[[eVUs ATJAnNAG’ JOU SapBVos [VYJUBO IOTIO}UB Ao[[etus Wetpeut ey} ‘sateos Eg Aq poyeiedas Ssopoitorwmes [eyrqioevidns pa)jaay ‘azvbwoja ‘sno -19tUNU WOLSsoidep [e},UOIZ UT SoTvos JNOUS SSOLOV SoCs ZT-OT pjO01jUOW TWO POORLY KNOWN ANOLES 9 1962 yyoous {ews sopvos ‘ploy [vurpnytsuoyT Be Aq pozBotpur “[eistsea de[Map ayo -TIQWIIGNS ‘SMO OSLOASUVIY UT SYST -Ivnbs 10 papunot “[[eus s[TRIyWeA posiv[uo SMo.a [esopprut Ou S[BIGVIGNS Wod9MJOG SoTBoS E PTA qoVjUOD UL SUOT UY} JOpTA sTezyUUT ayVotIquit YVpropaséd ‘STV.14 -UdA UVY} AISUO] BIB Salvos 910M ospoa ye ydooxo) ulys poyeu jo svoie Aq poyeredoes s[BiqUeA Uvtyy Id[][VUIS SofBos YJOowWS F-E JO SMO YQtIm juoutrmoid ‘aatsuezxe depMop posod -BYxNE ‘SMOL OSIOABIBAY UL ‘TYStavNbs 10 papunor Ypeuts AdodA s[RipWOA (ATWO sayeut UT) posivlud ATJYSTS SMOL [VSLOpprul F (sdr} olf} WaeMyoq po}tosut F-Z) Apsotseysod speVrqey -qus UWoeMjJoq Sa[VoS g YIM 4ov4 -U0) UL OPIM UY} JasUOL s[eyUuoUT (‘qu0/))) T ATaV], pojeey “[peus ‘prog [eurpnzytsuol, = 4yaT78 ve sq poyeotpur “erdstysea depaMop So[vos ayeuoroNnU ‘oyeoTIquit ‘SMOI ASIOASUBI} UL ‘papoody “[[RUIS S[RIQUAA polaoy ‘WTTOMS SMOI [VSLOPplul F-F B[UIGBTqns WweeMJoq A[LOLIOJSOd soleos fF YYTA 4oBJUOD UL SWOT UBYY JOPIM s[eyuTT 10 BREVIORA No. 164 LITERATURE CITED CocHRAN, D. M. 1935. New reptiles and amphibians collected in Haiti by P. J. Darling- ton. Proce. Boston Soc. Nat. Hist., 40: 367-376. 1939. Diagnoses of three new lizards and a frog from the Dominican Republic. Proc. New England Zool. Club, 18: 1-3. 1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177: 1-398. SHREVE, B. 1936. A new Anolis and new Amphibia from Haiti. Proce. New Eng- land Zool. Club, 15: 93-99. WILLIAMS, E. E. 1960. Notes on Hispaniolan herpetology. 1. Anolis christophei, new species, from the Citadel of King Christophe, Haiti. Breviora, INO, dls I=". et KNOWN ANOLES TWO POORLY 1962 “F009 Z KASAOTUHIULS 'N JIN "OA x YG Dp) 0. DUPWOUW SL)OUP (@ \ LOB CT BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. Aucust 22, 1962 NuMBER 165 AN EXTINCT SOLENODONTID INSECTIVORE FROM HISPANIOLA By Bryan Patterson During the summer of 1958, Drs. Clayton E. Ray and A. Stanley Rand earried on field work! for this museum in Puerto Rico and the Dominican Republic. Particular attention was paid to cave deposits, and a number of previously unexplored caves were examined. The cave containing the material re- ported on here is in the Sierra de Neiba near Rancho La Guardia in the Province of San Rafael, Dominican Republic; it has no local name. Situated in a limestone cliff and accessible only by a nearly vertical climb of some 30 feet, the cave con- tains a number of chambers connected by narrow passages. Bones were encountered in the deposits on the floor of the antechamber and of the first chamber beyond it. In the ante- chamber two layers are present, an upper, dark grey to black one some six inches thick, and a lower, reddish-brown one eight to ten inches in depth. The upper layer is either lacking or negligible in depth in the first chamber, the lower there attaining a thickness of six to twelve inches. Secanty remains from the upper dark layer include Rattus, which clearly indicates post- Columbian age. No introduced forms were encountered in the reddish-brown layer, which contains indigenous rodents and the three species of Nesophontes? known to have inhabited the island, together with scanty remains of bats, birds, lizards, and solenodontids. In addition to the bones of the extinct form de- scribed below, one specimen of Solenodon paradoxus was ob- tained. This is the left horizontal ramus of a young individual 1 Partly supported by a Sigma Xi-RESA grant-in-aid. 2N. paramicrus, hypomicrus and zamicrus of Miller (1929). N. “naramicrus,” the largest of the three, agrees in size with the Cuban N. micrus, from which Miller separated it on molar characters. I have examined several hundred speci- mens of the Hispaniolan form and compared them with Cuban material. The supposed differences are not constant and I have so far been unable to find others that might validate Miller’s species. 2 BREVIORA No. 165 with I, in process of eruption and alveoli of the other teeth (M.C.Z. no. 7260) ; as far as I am aware this find constitutes the first fossil (or subfossil) record of the species. Judging from the good preservation and completeness of most of the bones from the reddish-brown layer, Dr. Ray (MS field notes) is inelined to doubt that their occurrence is a result of owl-roost accumulation. Most of the material, including all the solenodontid remains, was found in the first chamber of the cave. As regards the age of the layer, all that can presently be said is that it is almost surely pre-Columbian. During their stay in the Dominican Republic, Drs. Ray and Rand were accompanied by Professor Eugenio de Jesus Mareano F., Universidad de Santo Domingo, whose aid was invaluable in all phases of the work. It is a pleasure to name the extinct solenodontid in his honor. Dr. Ray will discuss the rodents from this and other localities in the course of his comprehensive studies on the Antillean Rodentia. I am indebted to Miss Linda Loring for the cleaning, sorting and cataloguing of the other groups represented in the collection. The drawings are by Mrs. Dorothy Marsh. I have been fortunate indeed as regards com- parative material, thanks chiefly to the excellent series of Solenodon in the Mammal Department of the museum. Thirty- six mandibles, fifteen humeri and six ulnae of S. paradoxus and three mandibles of S. cubanus have been available. I have not seen limb bones of 8S. cubanus, but published figures (e.g. Peters 1863) indicate that these do not differ appreciably, either in structure or in proportions, from those of the larger species. INSECTIVORA SOLENODONTIDAE ANTILLOGALE* gen. nov. Type: — A. marcanoi sp. nov. Distribution: — Quarternary, Hispaniola. Diagnosis: — Differing from Solenodon as follows: Py and lower M smaller relative to size of jaw, lower M with lingual cleft between paraconids and metaconids deeper, paraconid wings directed more anteriorly, paraconids consequently farther from metaconids, low ridge between bases of paraconids and metaconids isolating slight valleys at bases of trigonid basins, anterior 1The Antilles, plus ya\7 , weasel. 1962 EXTINCT SOLENODONTID INSECTIVORE 3 cuspules below bases of paraconids very weakly developed, heels less broadly shelf-like labially. Post-dental portion of ramus larger relative to anterior portion. Humerus and ulna shorter, much wider relative to length, ulna with pit proximo-medial to sigmoid notch. ANTILLOGALE MARCANOI sp. nov. Type: — M.C.Z. no. 7261, incomplete right ramus of mandible with Ps3-Ms and alveoli of other teeth. Hypodigm: — Type and the following specimens: M.C.Z. nos. 7262, incomplete left ramus with Py-Ms and alveoli of Ps, Mz: 7266, posterior portion of left ramus with alveoli of Ms, juvenile; 7263, right humerus lacking ectepicondyle; 7264; left humerus lacking proximal epiphysis; 7265, right ulna. Horizon and locality: — Late Pleistocene or Recent ; unnamed cave 2 kilometers SE of Rancho La Guardia, Municipio de Hondo Valle, Provincia de San Rafael, Reptiblica Dominicana. Diagnosis: — As for the genus. Description: — Knowledge of the lower incisors, canine and anterior premolar derives solely from alveoli. On this evidence, the relative sizes of the incisors and the degree of enlargement of Ip were essentially as in Solenodon, as was the relative size of the single-rooted canine. The anterior premolar would appear to have been larger than Ps, to about the degree seen in S. para- doxus. Ps is small relative to Py, nearly as much so as in SV. paradoxus, and much smaller, both actually and relatively, than in S. cubanus. In structure it is very similar to the corresponding tooth of S. paradorus, and rather different from the larger, more globular one of S. cubanus. The paraconid of Py is a larger cusp than in the majority of specimens of S. paradoxrus and much larger than in S. cubanus. The essential characters of the molars have been given in the diagnosis and ean be seen in the figures: I may add that, as far as these teeth are concerned, Antillogale eould only with difficulty be distinguished generically from Apternodus. The horizontal ramus is shorter and more slightly built than in Solenodon and, instead of the two, or even three, mental foramina almost invariably present in that form, there is only a single large one, situated beneath Ps. The postdental portion of the mandible is relatively large and robust. Whereas the horizontal ramus is rather shorter than in S. cubanus the pos- terior part is rather larger and longer, being intermediate in 4 BREVIORA No. 165 a Figure 1. a) Antillogale marcanoi, type, M.C.Z. no. 7261, incomplete right ramus, dorsal view. b) Solenodon paradoawus, M.C.Z. no. 12384, crown view of right P3-Mo. X 3. these respects between cubanus and paradoxus. The coronoid process, so far as can be judged from the juvenile M.C.Z. 7266 and the incomplete M.C.Z. no. 7262, appears to have been less — tapering than in the living species. The base of the anterior border of the process is convex, as in S. cubanus, and not ex- eavated as it is in NS. paradoxus. The masseteric fossa is small, oO 1962 EXTINCT SOLENODONTID INSECTIVORE relatively and actually, oval in outline, and not very sharply defined. The marginal process, for the insertion of M. digastricus, is comparable to those of the living forms. The angle is not complete in any of the specimens but enough is preserved in M.C.Z. no. 7262 to demonstrate that, in contrast to S. paradoxus and in agreement with S. cubanus, the ventral border is not b Figure 2. Antillogale marcanoi. a) M.C.Z. no. 7261, type, incomplete right ramus, lateral view. b) M.C.Z. no 7266, incomplete left ramus, lateral view, X 2. downcurved and that, consequently, there is no Innate notch between angle and marginal process. The limb bones differ markedly in proportions from those of Solenodon. The humerus of Antillogale may be summed up as a bone having the width but not the length of that of S. para- doxus, and hence massive. The relative leneths of humerus and ulna appear to be essentially the same in both. The ulna, like 6 BREVIORA No. 165 the humerus, is shorter than but equally as wide as that of the living species; other differences of note between the two are the presence in the fossil of a very distinet pit proximo-medial to the sigmoid notch and of a rugose interosseous border. Si iN Figure 3. Lateral views of left rami of a) Solenodon paradoxrus, M.C.Z. no. 34828, b) Antillogale marcanoi, M.C.Z. no. 7262, and c¢) Solenodon cubanus, Y.P.M. no. 1203. In b) areas in solid outline are restored from M.C.Z. nos. 7261 and 7266. 3/2. 1962 EXTINCT SOLENODONTID INSECTIVORE —] Figure 4. Anterior views of humeri (a-c) and ulnae (d, e). Antillogale marcanoi, a) M.C.Z. no. 7264, b) M.C.Z. no. 7263, d) M.C.Z. no. 7265. Solenodon paradoxus, ©) and e) M.C.Z. no. 12416. X 4/3. 8 BREVIORA No. 165 Measurements in mm. M.C.Z. nos. 7261 7262 P3, length 2.7 — width 2.0 — P4, length Soll ay width 2.3 2.5 My, length 3.0 3.4 width 2.9 — Mo, length 3.3 3.4 width 3.0 Be) Depth of ramus beneath My, 6.5 — 7263 7264 Humerus, total length 40.0 _— a.-p. diameter of proximal end O39 tr. diameter of proximal end 10.9 a.-p. diameter at center of shaft oll 6 tr. diameter at center of shaft U3 6.5 a.-p. diameter of distal end 5.1 4.9 tr. diameter of distal end — WH tr. diameter of trochlea -- 9.2 7265 Ulna, total length 45.6 a.-p. diameter of olecranon 4.8 tr. diameter of olecranon 7.4 tr. diameter of proximal articular surface 6.4 a.-p. diameter at center of shaft 4.3 tr. diameter at center of shaft 23 a.-p. diameter at distal end 3.9 tr. diameter at distal end 2.8 Discussion: — The Solenodontidae have thus far been known from one genus with two species. Splitting of these taxa has of course been attempted. Solenodon cubanus has been made the type of Atopogale by Cabrera (1925), and Barbour (1944) has described a second Cuban species, S. poeyanus, based on pelage characters. Barbour explicitly stated that he could detect no cranial or dental distinctions. Aguayo (1950) and Koopman and Ruibal (1955) have regarded this form as a subspecies of cubanus. The latter authors noted that some fragmentary remains (M.C.Z. no. 7054) from a cave in Camaguey, which they referred to the living species, were somewhat larger than Recent specimens of cubanus examined by them. In recent years, members of the Sociedad Espeleol6gica de Cuba have obtained further material, including some skulls, from the Province of Havana. Thanks 1962 EXTINCT SOLENODONTID INSECTIVORE 9 to Sr. Oscar Arredondo, I have seen most of these specimens ; they also represent individuals rather larger than, although not other- wise different from, any in the small sample (3) of cubanus available to me. It is of some interest that the type of poeyanus (M.C.Z. no. 6957) is also rather larger than this sample, a point not mentioned by Barbour; in fact there is agreement in this respect between the type and these fossils (or subfossils). To conclude from this that poeyanus is distinguishable from cubanus on the basis of size and that the fossils so far found are referable to the former would, I think, be premature. The available series are too small to rule out accidents of sampling. The size differ- ence is slight and if the size range of cubanus is comparable to that shown for paradoxus by the adequate series at hand we could be dealing simply with segments of a normal distribution. Reinforcing caution is the fact that one of the two mandibles from Maisi, Oriente (M.C.Z. no. 10065), mentioned by Allen (1918) is smaller than the other fossils. Whatever the solution of this minor problem may prove to be, I agree entirely with Aguayo and with Koopman and Ruibal that subspecies, at most, are involved. There is at present no good evidence of more than one species of Solenodon in Cuba. Atopogale is recognized, either as a genus or as a subgenus, by some authors. There can be no question that paradoxus and cubanus are clear-cut taxa but they seem to me, as to others, to merit no more than specific rank. Antillogale marcanoz helps to clarify matters here; it is sharply distinct from either of the living species and the latter share most of the characters that differentiate them from it. Antillogale is quite evidently a member of the Solenodontidae. The structure, although not the proportions, of the known limb bones; the general structure of the mandible, and especially the presence of a marginal process; the small I, and the degree of enlargement and mode of implantation of I, (as revealed by the alveoli) —all these characters combine to place this con- clusion beyond reasonable doubt. Only in lower molar structure is there a closer resemblance to members of another family, the Apternodontidae. The difference here I believe to be the result of specialization in the Solenodon phylum. The molars of the two living species give the impression of having undergone anteroposterior compression of the trigonids. A majority of the characters in which they differ structurally from those of Antillogale could be directly correlated with such a change. Aside from molar structure, the differences between the extinct and the living species are chiefly of a proportional nature, as 10 BREVIORA No. 165 pointed out above. Antillogale evidently had a somewhat shorter facial region and much shorter, more heavily built fore limbs than Solenodon. It may well have been more fossorial in habits. Solenodon with its two species, one on each of the larger West Indian islands, has always been one of the most isolated of mam- mals, zoogeographically — and even taxonomically — speaking. As knowledge of the past history of mammals has improved, this isolation has become, if anything, even more apparent. In earlier days it was at least possible to assume a common ancestry with if not membership in the Tenrecidae. There is now no real evidence for such an assumption — tenrecids, as far as the record goes, appear always to have been African and Madagascan in distribution — and we must look elsewhere for close relatives and possible ancestors. The only group that seems to me to come close to fulfilling the requirements is the Apternodontidae, a family which at least had the merit of inhabiting North America, the only probable source area.! The described forms, the Oligocene Apternodus and Oligoryctes, were certainly not ancestral to the solenodontids, but apternodontids are now known in North Amer- ica at least as far back as the Bridgerian Eocene.? The possibility, I would go so far as to say probability, exists that solenodontids were derived from relatively unspecialized apternodontids that inhabited the Central American peninsula during the earlier Tertiary. Rafting of the ancestral stock to the Antilles, for rafting was certainly involved, may have taken place in the later part of the Eocene, at roughly the same time as the rafting of the ancestors of ecaviomorph rodents and platyrrhine primates to South America. This event, if it occurred at the time suggested, would have insured for the solenodontids a very long residence in the West Indies. There has, I feel, been some reluctance to accept such a possibility, a reluctance based, consciously or un- consciously, on the fact that Solenodon alone has hitherto repre- sented the family and on the assumption that we now have an adequate idea of the Pleistocene, or at least pre-human, faunas of the archipelago. I strongly doubt if we do have a good 1 Allen’s belief (1918) that solenodontids were derived from nesophontids. a view recently supported by McDowell (1958). is not at all convincing. McDowell's attempt to read Apternodus out of the Insectivora is utterly unrealistic, being in large part based on misinterpretation of the fossils he examined. 21I am indebted to Dr. Craig C. Black for the opportunity of examining a speci- men from Tabernacle Butte, Wyoming. This is the earliest known eutherian with zalambdodont molars. Contrary to earlier statements (including one of mine made on the basis of the literature), the molars of the Paleocene Palaco- ryetes are not zalambdodont in structure, nor are they even pre-zalambdodont in the way those of Potamogale are. 1962 EXTINCT SOLENODONTID INSECTIVORE 11 knowledge of the faunas (cf. Koopman and Williams 1951) ; as regards birds at any rate Sr. Arredondo and I will demonstrate the contrary in a forthcoming paper. And now, in Antillogale, we have a hint that there may have been a radiation of solenodon- tids on the Antilles in some degree comparable to that of the tenrecids on Madagascar. REFERENCES Aquayo, C. G. 1950. Observaciones sobre algunos mamiferos cubanos extinguidos. Bol. Hist. Nat. Soe. Felipe Poey, 1: 121-134. ALLEN, G. M. 1918. Fossil mammals from Cuba. Bull. Mus. Comp. Zool., 62: 131- 148. Barsour, T. 1944. The solenodons of Cuba. Proce. New England Zool. Club, 23: 1-8. CABRERA, A. 1925. Genera Mammalium. Insectivora, Galeopithecia. Madrid: 1-282. Koopman, K. F. anp R. Rupa 1955. Cave-fossil vertebrates from Camaguey, Cuba. Breviora, Mus. Comp. Zool., no. 46: 1-8. Koopman, K. F. anp E. E. WILLIAMS 1951. Fossil Chiroptera collected by H. E. Anthony in Jamaica, 1919- 1920. Amer. Mus. Novitates, no. 1519: 1-29. McDoweE tt, S. B. 1958. The greater Antillean insectivores. Bull. Amer. Mus. Nat. Hist., 115: 113-214. MILLER, G. S. 1929. A ' L y'? ’ iis ii {¢ or eae? ea puaninds PT ey oe °@/) aha ' sab te m8 pais i 0 phe 7 D . BREVIORA Museum of Comparative Zoology palin Sn Mie eT) A Bs te Re Pe PES a Td RN a I EIT oe a NO er CAMBRIDGE, Mass. SEPTEMBER 5, 1962 NuMBER 166 LECTOTYPES OF SPECIES OF OGCOCEPHALIDAE SELECTED FROM SYNTYPES IN THE MUSEUM OF COMPARATIVE ZOOLOGY By Marcaret G. Brappury ! Hopkins Marine Station Pacific Grove, Calif. During the 1891 expedition of the U. S. steamer ‘‘ Albatross”’ to the eastern tropical Pacific, about 80 specimens of fishes of the family Ogecocephalidae were taken. Garman, who studied the collection, concluded that it represented twelve species, ten of which were described by him as new (Garman, 1899). The remarkable thing is that subsequent collecting in the same area has produced additional examples of only about half of these species; the rest remain known only from specimens taken on the original expedition. It became necessary for me to examine this valuable ‘‘ Albatross’’ collection in the course of work on the family Ogeocephalidae. Then it was discovered that most of the specimens upon which Garman based his original descriptions had been designated as syntypes, a common practice at the time. In the interest of stabilizing the nomenclature of this group, I have selected lectotypes for the species heretofore represented by syntypes. The list of species below includes only those for which Garman wrote original descriptions. They are arranged in the order in which they appear in his paper. OGCOCEPHALUS PORRECTUS The original lot contained four specimens from ‘‘ Albatross’’ Sta. 3368A, and of these Hubbs (1958) selected the largest speci- men as lectotype; it bears the number MCZ 28733. The other three specimens are now removed to MCZ 41594. ‘This investigation was supported by Publie Health Service Fellowship No. 8543 from the Division of General Medical Sciences, Public Health Service. bo BREVIORA No. 166 HALIEUTOPSIS TUMIFRONS The only two specimens known of this species are those originally taken by the ‘‘Albatross.’’ It was not possible for me to determine which of the two had been figured by Garman, so the choice of a lectotype was arbitrarily made, the two ex- amples being closely similar. The lectotype, MCZ 28729, is from ‘* Albatross’’ Sta. 3400A. The second specimen, from ‘‘ Albatross”’ Sta. 3413A, bears the number MCZ 28730. DIBRANCHUS HYSTRIX Garman assigned three examples to this species, and from these I have chosen the specimen that appears to be the one illustrated in the original description; therefore the lectotype is MCZ 28726, ‘‘ Albatross’’ Sta. 3375A. The other two specimens are: MCZ 28727, ‘‘Albatross’’? Sta. 3392A, and MCZ 28728, ‘* Albatross’’ Sta. 3362A. DIBRANCHUS SCABER Two specimens represented this species originally, the larger of which is apparently that figured in the original description. It has the number MCZ 28724, ‘‘ Albatross’’ Sta. 3431A, and is hereby designated the lectotype. The second specimen, MCZ 28725, ‘‘Albatross’’ Sta. 3364A, is a specimen of Dibranchus hystrix, not Dibranchus scaber. The lectotype of Dibranchus scaber is therefore the only known specimen of the species. DIBRANCHUS ASPER This species was described from a single specimen, MCZ 28723, ‘‘Albatross’’ Sta. 3418A; accordingly this specimen is the holotype. My own studies show Dibranchus asper to be the same as Dibranchus hystrix; of the two nominal species, I hereby place Dibranchus asper in the synonymy of Dibranchus hystrix for the reason that no figure accompanies the description of Dibranchus asper. DIBRANCHICHTHYS NUDIVOMER Altogether, two lots totalling seven specimens represent this species. A specimen in relatively good condition from MCZ 28719, ‘‘Albatross’’ Sta. 3353A, was selected as the lectotype; there was one other specimen in the lot with the lectotype, and 1962 LECTOTYPES OF OGCOCEPHALIDAE 33 it is now removed to MCZ 41595. The second lot of five speci- mens bears the number MCZ 28718, ‘‘Albatross’’ Sta. 3395A. These, along with an eighth example from ‘‘Albatross’’ Sta. 3395A which had been sent to the U. S. National Museum (USNM 57867), are the only known specimens of this species. MALTHOPSIS SPARSA Garman originally assigned three lots totalling 19 specimens to this species. Of these it is impossible to tell exactly which example was used for the illustration in the original descrip- tion, and the lectotype was chosen more or less arbitrarily from a series considered to be in good condition and which most resembled the original illustration. The lectotype, MCZ 28717, bears a tag indicating that it came from ‘‘ Albatross’’ Sta. 3386A, but the situation is not so clear with respect to most of the specimens that were originally in the same lot with the lectotype. Of these there are eight specimens as follows: two specimens with the same station number as the lectotype, 1.e., ‘‘ Albatross’’ Sta. 3386A ; one specimen tagged ‘‘ Albatross’’ Sta. 3354A ; five specimens with no tags and no indication in the original de- scription as to whether they came from station 3386A or 3354A. These untagged specimens together with those from station 3386A are now removed to MCZ 41596. The specimen from ‘Albatross’? Sta. 3354A is certainly an example of Malthop- sis spinosa, not Malthopsis sparsa, and is now removed to MCZ 41597. The remaining two lots of Malthopsis sparsa stand as fol- lows: MCZ 28715, ‘‘ Albatross’’ Sta. 3396A, six specimens; MCZ 28716, ‘‘ Albatross’’ Sta. 3385A, four specimens. MALTHOPSIS ERINACEA From the original series of eight, the one specimen that ap- pears to have been used for the illustration accompanying the original description is hereby designated the lectotype; it bears the number MCZ 28712, ‘‘ Albatross’’ Sta. 3402A. A second specimen originally in the same lot with the lectotype is now removed to MCZ 41598. The other six specimens bear museum numbers as follows: MCZ 28711, ‘‘ Albatross’’ Sta. 3358A, one specimen; MCZ 28713, ‘‘Albatross’’ Sta. 3425A, one specimen (this is probably not a specimen of Malthopsis erinacea, but it 4 BREVIORA No. 166 is small and difficult to place at this time without more com- parative material) ; MCZ 28714, ‘‘ Albatross”’ Sta. 3418A, four specimens (these four are actually examples of Malthopsis spinosa, not Malthopsis erinacea). MALTHOPSIS SPINOSA Two rather similar specimens were assigned to this species by Garman, but I am unable to tell which was used for the illus- tration accompanying the original description. I have selected MCZ 28710, ‘‘Albatross’’ Sta. 3392A, as the lectotype. The re- maining specimen from ‘‘ Albatross’’ Sta. 3393A, originally with the same museum number as the lectotype, is now removed to MCZ 41599. MALTHOPSIS SPINULOSA From a series of 21 specimens from ‘‘ Albatross’’ Sta. 3394A, I have selected a specimen on the basis of its similarity to the illustration used with the original description and the fact of its relatively good condition. This specimen bears the number MCZ 28709, and the remaining 20 specimens of the lot are removed to MCZ 41600. From my own examination of this material in conjunction with newer collections, I must conclude that Malthopsis spinulosa is the same as Malthopsis spinosa. Inasmuch as the original descriptions of both nominal species are accompanied by illustrations of equal quality, I give prefer- ence to Malthopsis spinosa as the name to represent this taxon by virtue of its page priority. Thus, Malthopsis spinulosa falls into the synonymy of Malthopsis spinosa. I am extremely grateful to Mrs. Myvanwy Dick and to Dr. Giles W. Mead, who both took great trouble to assist me in the loan of material. LITERATURE CITED GARMAN, S. 1899. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer ‘¢ Albatross’? during 1891, Lieut.-Commander Z. L. Tanner, U.S.N., commanding. XXVI. The Fishes. Mem. Mus. Comp. Zool. Harvard Coll., 24: 1-481, pls. 1-88, A-N. Husss, Cari L. 1958. Ogcocephalus darwini, a new batfish endemic at the Galapagos Islands. Copeia, 1945 (3): 161-170, pls. 1-5. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. SEPTEMBER 5, 1962 NuMBER 167 Bathyclupea schrocderi, a New Bathyclupeid Fish from the Western Tropical Atlantic By Myvanwy M. Dick Museum of Comparative Zoology, Harvard University The species described below is based on specimens collected by the Atlantis during the Harvard-Havana Expedition to Cuba, and by the Oregon (exploratory vessel of the U.S. Bureau of Com- mercial Fisheries) during her work in the Gulf of Mexico. To Mr. Harvey R. Bullis, Jr., director of the Oregon’s program, go my thanks for representatives of both western Atlantic species of Bathyclupea. I am also indebted to Mr. Loren P. Woods of the Chicago Natural History Museum, and to Dr. Leonard P. Schultz, U.S. National Museum, for the loan of comparative specimens of related species and for additional representation of the new form. BATHYCLUPEA SCHROEDERI, new species Holotype: A specimen 140 mm. in standard length, taken at Atlantis Station 2987, 23°23’N., 79°39’W., 280 fathoms. MCZ 41498. Paratypes: MCZ 39416, 4 specimens, Atlantis Station 2987, 23°23’/N., 79°39’W., 280 fms.; MCZ 39380, 1 specimen, Atlantis St. 2987a, 23°22’N.; 79°39’ W., 260 fms. ; MCZ 40600, 1 specimen, Oregon St. 2635, 17°37’N., 63°28’W., 220-235 fms.; CNHM 65145, 1 specimen, Oregon St. 1872, 16°41’N., 82°20’W., 300 fms.; CNHM 65147, 1 specimen, Oregon St. 1888, 16°41’N., 81°02’W., 250 fms.; CNHM 65146, 1 specimen, Oregon St. 1886, 1G 2a5 Ne oil 127W., Zia ims: Diagnosis: The following species of Bathyclupea have been described : Bathyclupea hoskynii Alcock, 1891, from the Andaman Sea (the type species) ; Bathyclupea argentea Goode and Bean, 9 BREVIORA No. 167 1895, from Nevis, West Indies; Bathyclupea maylayana Weber, 1934, from the Flores Sea, Indonesia; Bathyclupea megaceps Fowler, 1937, from off Mindanao, Philippines; and Bathyclupea gracilis Fowler, 1937, from off Makyan Island, Moluceas. Bathyclupea schroederi differs from all of these by having a spine and 37 to 39 rays in the anal fin (ef. 33 or fewer). It can also be distinguished from B. hoskynw by its less deep body (3.9 to 4.4 in standard length; ef. 3.3), and from B. argentea and B. maylayana by the placement of the anal fin, which originates in the anterior half rather than in the posterior half. B. schroederi can be distinguished from B. megaceps by the relative length of the base of its anal fin, which is longer than the head in the new form but shorter in B. megaceps. The length of the head in B. schroedert, 3.3 to 3.4 in standard length, is shorter than that of B. gracilis (3.0 in length). Description: The description which follows is based on the holotype and paratypes, 120 to 144 mm. in standard length, listed above. Counts and proportional dimensions are provided in Table 1. Body laterally compressed, its greatest depth at origin of anal fin, 3.9 to 4.4 in standard leneth. The dorsal profile is nearly straight. There is a well defined, rather straight lateral line bear- ing 38 pored scales which are somewhat more adherent than the seales elsewhere on the body. Scales ecycloid, deciduous, thin and relatively large. Head naked with large mucous cavities. Verte- brae 10-20-1, total 31. Head 3.3 to 3.4 in standard leneth, interorbital flat, 16.3 to 17.1 in standard length. Eye mainly in the posterior half of the head. Nostrils small, contiguous, almost superior. The mouth is nearly vertical. Minute conical teeth on the lower jaw, a single row of minute teeth on the vomer and premaxillary. The anterior end of the vomer protrudes into the mouth, frequently extend- ing forward to beneath the tip of its snout. There are four gill arches. Moderately long gill rakers, with slight protuberences, on the first and second arches; rakers much reduced and stublike on the third and fourth arches. Gill rakers on the first arch 3+ 16. The gill opening is wide, the branchi- ostegal membranes free from one another and from the isthmus. Pseudobranchiae present. The pectoral fin is large, pointed, the upper rays the longest, about 3.5 in standard length. The ventral fins are anterior to the pectorals, short, very slightly separated. The rays are rather 1962 NEW BATHYCLUPEID FISH 3 like the ribs of a fan, fitting into a shallow groove when con- tracted. The dorsal fin originates in the postmedian half of the body, the anal fin in the anterior half. Color in alcohol: Opercle and abdominal wall dusky. In life the general color was silvery. It is a pleasure to name this species for William C. Schroeder of the Department of Fishes, Museum of Comparative Zoology, who collected the holotype and several of the paratypes. His many publications, the collecting he has done, and his work in the department have greatly enriched the field of ichthyology. TABLE 1 Counts, and proportional dimensions expressed in percent of standard lenath, of the holotype and paratypes of Bathyclupea schroederi. re) We) we} Ne} we) ° as) ° ~ t - ” ” ay ay On + Sn Standard length (mm.) 144 144 140 139 137 1S 37/ 130 120 116 Greatest depth of body (percent) 27.8 26.4 PO hab eS FAz/a(t) 24.6) 25:20) 42353 Greatest depth of body at midpoint 26:46 2356 2570) 25.10) pean 24a 2301 (25.00 23.5 Least depth of caudal peduncle 10.4 CLT 9.3 9.5 9.5 EY) 9.2 G2. 8.6 Greatest width of body 10.4 9.7 7.9 9.5 9.5 8.7 7.7 9.2 7.8 Greatest width of head by alae} 252) L252 eA Lor: WG) ai) abl Snout to origin of dorsal fin Saree 5536 Siapli) Beh) Wee l7 0 = bie pe 5476 50:8 (5157 Snout to origin of anal fin 43.8 43.1 42.8 42.8 45.2 43.8 43.1 47.5 44.8 Snout to insertion of ventral fin 27.0 s2t.o 28.6 28.6 31.4 26.9 29.2 29.3 Snout to insertion of pectoral fin, ventral angle 2825) 2728 2728) 12923) 2929) (2959 26.9 28.0 29.3 Length of base of dorsal fin 10.4 10.4 10.7 10.6 10.9 10.9 10.8 10.0 10.3 Length of base of anal fin 51.4 49.4 475.) 4953) 482) 51-1 48.4 48.3 50.9 Distance between insertion of ventral fin and origin of anal fin 20.7 20-2 W728) LiekS V20:4. idee Load. alten Length of head Pashia) 743\05) 30.0 29.3 29.2 30.6 Zit) 30-0) 3220 Length of snout 8.3 8.3 8.6 8.6 8.7 8.7 8.5 Ore 7.8 Greatest diameter of eye LTSt ELL 4 0054 Bey Vy Os Ale hie Width of interorbital space 5.6 5.6 5.0 5.0 a9 Bre 5.4 5.8 6.4 Length of upper jaw nie}72) ali) WA SCR alec alee | 1253) liebe sD Counts: Dorsal fin 1-9 1-9 1-8 1-9 1-8 1-9 1-9 1-8 1-9 Anal fin 1-37 1-38 1-39 1-37 1-38 1-39 1-38 1-39 1-37 Pectoral fin 1-28 1-28 1-29 1-28 1-28 1-28 1-29 1-28 1-29 No. 167 BREVIORA “Bqny WOLF “QGFTF ZOW ‘ed«jzo[ox] Sere Byf GCERCECECEECEEEEE “aou “ds “ttopaoiyos vadnphyjvog "T AXODLAT BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. SEPTEMBER 7, 1962 NUMBER 168 TWO NEW SPECIES OF FOSSIL TALPID INSECTIVORES 3y KATHERINE M. REED During work on the talpid subfamily Proscalopinae, two new scalopine moles came to my attention and are deseribed in this note. I am grateful to Mr. Richard Tedford, University of Cali- fornia Museum of Paleontology, and Dr. C. W. Hibbard, Uni- versity of Michigan Museum of Paleontology, for the loan of the material. The work was carried out at the Museum of Compara- tive Zoology. I am obliged to Professor Bryan Patterson for critical comment and to Miss Barbara Lawrence for access to the collections of Recent insectivores. I also appreciate the com- ments and assistance of Dr. J. R. Macdonald and Dr. Mary Dawson. The illustrations were made by Mr. Richard Stafford. The following abbreviations are used: UCMP—University of California Museum of Paleontology UMMP—University of Michigan Museum ot Paleontology l., w., trig., tal—leneth, width, trigonid, talonid TALPIDAE Subfamily ScALOPINAE DOMNINOIDES Green 1956 DOMNINGIDES VALENTINENSIS hn. sp. Type: UCMP 33152, right ramus with P,, P3-4, Mo-s. Hypodigm: Type and UCMP nos. 36150-36157, ineluding iso- lated teeth, partial rami and some limb bones. UCMP 29215 and 29215-A refer to limb material marked ‘“‘float.’’ Horizon and locality: Late Miocene, Valentine formation, from the quarries near the middle of the exposed Valentine formation 2 BREVIORA No. 168 at the Fort Niobrara locality on the quarter section line between the NW and SW quarters of Sec. 24, T 34 N, R. 26 W, Cherry County, Nebraska, UCMP loeality V3218 (See Macdonald 1947). The specimens marked ‘‘float’’ are, according to Mr. Tedford, ‘undoubtedly Barstovian, but come from an undetermined horizon within the Valentine.’ Diagnosis: No diastema between Ps and P,; slightly larger and more robust than Domninoides riparensis; metastylid defi- nite on M..* Description: The only upper tooth in the material, and the first known to belong to this genus, is a broken, isolated P*. This tooth has a blade-shaped paracone. A very narrow anterior cingulum widens to a shelf-like lingual cusp and extends up the posterior side of the tooth to join the paracone erest posteriorly. The lingual cvsp is widest opposite the stoutest portion of the paracone. The tooth has at least two, possibly three roots, two labial, one lingual. In the lower dentition, P; is small, conieal and single rooted. Ps is not represented in the material but is double rooted. Ps is considerably larger than P,, double rooted as in D. riparensis, and with a slight heel. It is situated very close to P,, with no diastema between these teeth. Py is double rooted with a very small anterior cuspule. It has a larger heel than Ps, the heel sloping downwards labially to a small cingulum which connects the heel to the anterior cuspule. The main cusp is conieal. M, is not represented in the material studied; thus no com- parison with D. riparensis is possible in this respect. Relative root sizes are in accord with this species. In Ms the paraconid is smaller than the metaconid, which is the highest lingual cusp. The entoconid is the stoutest lingual cusp. A crest, the erista obliqua, runs from the hypoconid to the metastvlid, as in D. riparensis. The protoconid is higher than the hypoconid and is slightly labial to the metaconid. There is a large anterior ein- gulum, wide labially and with an irregular border, that extends around the paraconid to the opening of the trigonid valley. The posterior cingulum is small and does not reach to the lingual’ face of the tooth. The talonid has a narrow opening. In M. the size relations of the cusps are as in Ms. All examples of this tooth Ipr. J. R. Maedonald has kindly sent me a copy of a forthcoming manuscript in which he describes a new species of Demninoid«s. D. valentinensis is distin- guished from this species by lacking any trace of cingula on the lingual face of the molars and by the crista obliqua running to the metastylid rather than to the metaconid. 1962 NEW FOSSILS TALPIDS 3 are worn, but there are strong suggestions of a metastylid and of a relatively wide anterior cingulum. No posterior cingulum is present. The talonid is relatively longer than in Ms. There are usually two mental foramina which vary in_posi- tion. In UCMP 36151, they are below the anterior roots of P, and between Py, and M,; in UCMP 33152, between Ps-. and the roots of Py. In UCMP 36152, there is only one foramen, below the anterior root of Py. In D. riparensis, the foramina are below P. and between the roots of Py. The ramus is much like that of D. riparensis and is in general similar to that of Macdonald’s species, although shghtly stouter than in either. Some comparison of the limb material of the fossil has been made with modern species. It must be noted that the majority of the limb material is ‘‘float’’ and association with the teeth could perhaps be doubted; the extremely talpid-like nature of both does, however, strongly suggest association. The humeri of the fossil are very similar to but slightly smaller than those of Talpa europaea. They are larger than in Parascalops breweri and wider than in Condylura. Discussion: Macdonald has suggested (pers. comm.) that the jaws of his new species may represent the lower dentition of Proscalops sccundus. This now seems extremely unlikely, first because of the discovery of P+ of Domninoides valentinensis. which is very unlike that of the Proscalopinae, and second, the greater development of the anterior cingula on lower molars than I would expect on the basis of the known trends in the Proscalopinae (See Reed 1961). Wilson (1960) has recently de- scribed lower molars which he assigns to Proscalops sp. ef. P. secundus; these are much nearer to the proscalopine type of molar than to that of Domninoides. It is evident that Domninoides is a talpid, not a soricid as Green (1956) originally deseribed it. Measurements UCMP 33152 Ms UCMP 36152 Ms UCMP 33152 Ms l. 3.1mm IL 3.2 mm 1. (approx.) 2.4 mm Weibrie. 225 we trigy 22 w. trig. 1.9 w. tal. 2.7 w. tal. 2.7 w. tal. 1.6 depth of jaw below M,, UCMP 33152: 3.7 mm. 4 BREVIORA No. 168 HIESPEROSCALOPS Hibbard 1941 HESPEROSCALOPS SEWARDENSIS Nn. Sp. Hesperoscalops recroadi Hibbard 19538, p. 23, fig. 1D. Type: UMMP 27276, partial right ramus with Mo-3, fragment of a left ramus, anterior part of a left ramus with worn M,, ulna, part of a scapula, and a femur. Hypodigm: Type only. Horizon and locality: Late Pliocene or early Pleistocene, from area of Saw Rock local fauna, NE 1/4 See. 35, T 34 8, R 31 W, Seward County, Kansas. Diagnosis: Distinctly larger than H. rerroadi and with greater development of basal accessory cuspules. Description and discussion: The difference in size between the new species and specimens of H. rexroadi in which the teeth are in nearly the same state of wear indicates that these cannot be wear differences in the same species and that the two are distinct. Although the cusp pattern is very similar, as Hibbard states (1953, p. 23), the anterior cingular cuspule on Ms is larger and better developed than in H. rexroadi, as is the anterior basal ac- cessory cuspule on My. If Hesperoscalops rexroadi is ancestral to Scalopus, this new, but closely related species must represent an extinct side line, for /7. sewardensis is considerably larger than Scalopus aquaticus and has much better developed basal cuspules and cingula. Measurements H. sewardensis UMMP 27276 Mo. UMMP 27276 M2, Il, 3.0 mm ll 2.8 mm Wa tric Zea w. trig. 2.4 we tale 226 War bales wale depth of jaw below Ms on lnegual face: 4.5 mm. length of Ms-3 : 5.8 mm. H. rexroadi UMMP 27278 Mo. UMMP 27278 Mz lle 2.38 mm ll; 2.2 mm w. trig. 1.9 w. trig. 1.6 w. tal. 2.1 Wobales ens depth of jaw below Ms on lingual face: 3.2 mm. leneth of Mo5-3: 5.0 mm. OU 1962 NEW FOSSILS TALPIDS REFERENCES GREEN, M. 1956. The Lower Pliocene Ogallala-Wolf Creek vertebrate fauna, South Dakota. Jour. Paleontology, vol. 30, no. 1, pp. 146-169. HIBBARD, C. W. 1953. The insectivores of the Rexroad fauna, Upper Pliocene of Kan- sas. Jour. Paleontology, vol. 27, no. 1, pp. 21-32. MACDONALD, J. R. 1947. A new shrew from the Niobrara River, Upper Miocene of Ne- braska. Am, Jour. Sci., vol. 245, pp. 123-126. 1962. The Miocene fauna from the Wounded Knee area of western South Dakota. Bull. Am. Mus. Nat. Hist. (in press). REED, K. M. 1961. The Proscalopinae, new subfamily of talpid insectivores. Bull. Mus. Comp. Zool., vol. 125, no. 14, pp. 473-494. WILSON, R. W. 1960. Early Miocene rodents and insectivores from northeastern Colo- rado. Vertebrata, Univ. of Kansas Paleontological Contribu- tions, article 7, pp. 1-92. No. 168 BREVIORA ‘x @ cared ordo ‘X GYE “MOTA [BNSUTT ‘SNUB JYSII ‘OXT1G AINND ‘Sisuapapmas sdojposowadsaeY "Oly ‘X C/T “MOTA 1OTLo}S0d ‘snaoumMy 4yStt “~eTOS GINO ‘Sisuauruaypa "q +L ‘Bq ‘X G/T “MOTTA LOLIOJUB ‘snioUINYy FYStL “YCTOS GINO ‘sysuaeuyuapa "Gq “9 “SIA “X CGE “MOTA [BNSUTT ‘snUIBI yFeT ‘ceTEe GINOM ‘sisuawpwaypa “gq -e ‘BIA G GG *x Gg¢g ‘ared otdodsoadteys !MoTA UMOL) “VFO gd “GCLOE AINDD ‘siswaurjpuappa saproumuwmog “pF “Sty a19JS SSNUIRI JUS JO MOTA UMOI) *QITIT CININD ‘sisuvapipnas sdoppososadsafyy *°@ “Sly G ae *x @ cared ordoosoodeqs Ssnwiet 4yoT JO MOA UMO.LD ‘ZETCE GINO ‘sisuampuajpa sapromuwmog 7% “Sty 1G , nw T aLV Id BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. OcToBER 15, 1962 NuMBER 169 NEW RECORDS OF INSHORE FISHES FROM THE ATLANTIC COAST OF PANAMA By IRA RuUBINOFF’ AND RoBEeRTA W. RUBINOFF During February, March, and April, 1961, a collection of in- shore fishes from both coasts of the Isthmus of Panama was made in connection with a study of the effects of geographical isola- tion on fish speciation. This collection consisted of 2095 speci- mens of 136 species. A total of 907 specimens of 47 species was taken from the Pacific coast and 1188 specimens of 89 species were collected on the Atlantic coast. Fourteen species in the latter collection were found to represent extensions of known ranges and some of these are sufficiently abundant to be consid- ered resident members of the Panamanian fauna. The purpose of this paper is to report the new records from this area. Notes are also included on three species represented in previous collec- tions from Panama by only one or two specimens each. All the specimens discussed herein are in the collections of the Museum of Comparative Zoology. Rotenone poisoning of coral pools and lagoons was the princi- pal method of collecting, although occasionally sandy beaches were seined. All of the new records are from coral reef areas, particularly from the reefs bordering the Galeta Point Naval Station near Coco Solo in the Canal Zone. Most of the smaller eels were collected when the incoming tide overflowed poisoned coral pools onto the exposed surfaces of the reefs and into small erevices and burrows in the coral. When the poison reaches these burrows the inhabitants are driven onto the flat of the reef and are easily collected. Many young speci- mens of Muraenidae as well as most of the specimens of Morin- guidae and Echelidae were collected in this manner. 1This research was supported in part by a Public Health Service Training Grant to the Department of Biology at Harvard University, Cambridge, Massa- chusetts, and in part by a fellowship from the Woods Hole Oceanographic Insti- tution during the summer of 1961. Contribution No. 1253 of the Woods Hole Oceanographic Institution Y BREVIORA No. 169 In their classic work on the marine fishes of Panama, Meek and Hildebrand (1923, 1925, 1928) collected 235 species from the Atlantic coast, of which only five were eels. In other collec- tions from this area, the apodes are one of the poorest repre- sented groups. Numerous species are known to range along the Atlantie coast north and south of Panama but have yet to be recorded there. Of the nine species of apodes in our collection seven represent new records for this area. Specimen lengths are standard lengths to the nearest milli- meter in all cases except the eels, for which total lengths are given. The authors wish to thank the following people for making this part of our study possible: Dr. Sydney Galler and Mrs. Helen Hayes of the Biology Branch of the Office of Naval Re- search, Dr. L. P. Schultz of the United States National Museum, and Dr. Martin Moynihan and Mrs. Adela Gomez of the Canal Zone Biological Area. Dr. James E. Bohlke of the Philadelphia Academy of Sciences most kindly identified some of the eels. SPECIES NOT PREVIOUSLY RECORDED FROM THE ATLANTIC COAST OF PANAMA XENOCONGRIDAE Kaupichthys atlanticus Bohlke MCZ 41455-58 Five specimens, 50-205 mm. long, were taken from coral reefs at Galeta Point. These specimens agree with the description of Bohlke (1956). Range: Tropical western Atlantic from Bermuda to south of Jamaica. MORINGUIDAE Aphthalmichthys mayeri (Silvester) MCZ 41459-61 One specimen, 275 mm. in length, was taken from the edge of a reef southwest of Las Palmas mountain about two-thirds of the distance from Maria Chiquita to Porto Bello. This fish had a dis- tinetly pink head and pink-orange body. This color faded to a dull yellow two days after preservation. Three specimens, 150- 330 mm. long, were taken from the exposed surface of Galeta Point Reef. Range: Bermuda, Florida, Puerto Rico. 1962 NEW RECORDS OF PANAMANIAN FISHES 3 ECHELIDAE Myrophis egmontis Jordan MCZ 41462-64 Twelve specimens, 81-231 mm. in length, were obtained from Galeta Point. These specimens may be differentiated from Eche- lidae previously recorded from Panama by the following char- acteristics: origin of dorsal fin behind the vent, and absence of teeth on the vomer. These eels were found in situations similar to those of Aphthalmichthys mayert. Range: Florida, Bahamas, West Indies. MURAENIDAE Enchelycore mgricans (Bonnaterre) MCZ 41465-67 Seventeen specimens of this eel, 81-545 mm. in length, were taken from the reefs at Galeta Point. The slitlike posterior nostril diagnostic of Enchelycore is an unreliable character in separating young specimens from other Muraenidae. Separation of the young of EF. nigricans from Gymnothorax moringa which it closely resembles, was facilitated by two characters. The anterior nostrils of Enchelycore are shorter than those of G. moringa and the upper jaw of Enchely- core has a series of 4-6 long canine teeth medial to the outer row of teeth on both sides. All Gymnothorar moringa which we ex- amined have 1-3 teeth in this series. Range: Bermuda, West Indies. Gymnothoraxr vicinus (Castelnau) MCZ 41468,69 Four specimens, 145-330 mm. in length, were taken at the Galeta Point reefs. Range: Bermuda, West Indies to Brazil, Cape Verde Islands and Africa. Gymnothorax moringa (Cuvier) MCZ 41470-7383 The collection contains nine specimens, 64-185 mm. in total length, taken from the reefs at Galeta Point. Range: Atlantic coast of America from Florida to Brazil, Ber- muda, Bahamas, West Indies, St. Helena. Uropterygius Bohlke (n. sp., in manuscript) MCZ 41475 4 BREVIORA No. 169 One specimen 190 mm. long was collected at Galeta Point. Our specimen was identified by Dr. James Bohlke as a species he is currently describing and it will be designated as a paratype. AULOSTOMIDAE Aulostomus maculatus Valenciennes MCZ 41474 One specimen, 185 mm. long, was taken at Galeta Point. Range: Bermuda, Florida, Gulf of Mexico, Central American eoast of Caribbean, Bahamas, West Indies. HOLOCENTRIDAE Holocentrus coruscus Poey MCZ 41476 One specimen, 45 mm. long, from Galeta Point, has the follow- ing meristics: D-XI, 12; A-IV, 8; gill rakers 9. Color of fresh specimen: dorsal surface of head red; body red and white lateral stripes ; caudal fin red; anal and second dorsal dark red at distal portions; interspinous membranes of dorsal peppermint striped, black spot distally between first three dorsal spines. With the exception of the number of gill rakers this specimen fits the de- scription of H. coruscus by Woods (1955) in his revision of the Western Atlantic species of Holocentrus. Range: Bermuda, Florida, Bahamas, West Indies. APOGONIDAE Apogonichthys stellatus Cope MCZ 41477 Two specimens, 12 mm. and 29 mm. in length, were taken from the edge of a reef southwest of Las Palmas mountain about two- thirds of the distance from Maria Chiquita to Porto Bello. Range: Bermuda, Florida, Bahamas, West Indies. LABRIDAE Thalassoma bifasciatum (Bloch) MCZ 41478-83 Fifty-eight specimens 19-83 mm. in length, were taken at Galeta Point. Although this species has not been previously recorded from Panama it is one of the most abundant representatives of the reef pool fauna. Many more examples were seen than were collected. Range: Bermuda, Florida, Bahamas, West Indies, Honduras. 1962 NEW RECORDS OF PANAMANIAN FISHES 5 CANTHIGASTERIDAE Canthigaster rostratus (Bloch) MCZ 41484, 85 We collected eleven specimens, 19-42 mm. in length, at Galeta Point. Ten of these specimens have the fin formula D-10, A-9; one specimen has D-9, A-9. These specimens agree with the de- scription of Breder (1927). Jordan and Evermann (1898), Evermann and Marsh (1900), and Nichols (1930) report speci- mens with the fin formula D-6, A-8. For a partial explanation of this discrepancy see Breder (1927). Range: Bermuda, Florida, West Indies, Venezuela and Madeira. CLINIDAE Labrisomus kalisherae (Jordan) MCZ 41486, 87 Six specimens, 25-68 mm. in length, were found in Galeta Point reef pools. One 56 mm. specimen taken in the first week of April possessed enlarged ovaries from which ova 0.5 mm. in diameter were obtained. A 54 mm. male also taken at this locality had enlarged testes. Range: Florida to Brazil. Labrisomus nigricinctus Rivero MCZ 41488 One male specimen 45 mm. long was taken at Galeta Point. It was compared with the holotype, MCZ 34150, and with the de- scription given by Springer (1958). Springer (1959) reports the range extension of L. bucciferus and L. guppy? to the Atlantic coast of Panama. L. nuchipinnis reported by Meek and Hildebrand (1928) and our specimens of L. kalisherae and L. nigricinctus bring the total recorded number of Atlantic Panamanian Labrisomus species to five. SPECIES RARELY RECORDED FROM THE ATLANTIC COAST OF PANAMA With the exception of Dinematichthys cayorum these species were not collected by Meek and Hildebrand (1923, 1925, 1928). BLENNIIDAE Rupiscartes atlanticus (Cuvier and Valenciennes) MCZ 41489-91 6 BREVIORA No. 169 Hight specimens, 39-74 mm. in length, from the coral reefs at Galeta Point are in the collection. The previous record of this species from Panama was a single specimen taken by Fowler (1916) at Toro Point. Range: Bermuda, Atlantic and Pacifie coasts of tropical Amer- ica, West Indies. Salarichthys textalis (Quoy and Gaimard) MCZ 41492 Three specimens, 24-42 mm. in length, were taken at Galeta Point. One 31 mm. specimen was previously collected at Cale- donia Bay, Panama, by Breder (1925). Range: Bermuda, Florida, West Indies, Brazil. BROTULIDAE Dinematichthys cayorum (Evermann and Kendall) MCZ 41493-96 Twelve specimens of this species (Ogilbia cayorum of Meek and Hildebrand), 26-51 mm. in length, were collected at Galeta Point. On April 6 some adults were found to contain well developed embryos which could be seen through the body wall. These were extruded in gelatinous strings when a slight pressure was exerted on the abdomens of the females. They were about 5-7 mm. in length and had small yolk sacs. When placed in a bucket of sea- water the embryos were free swimming although apparently pre- mature. Fowler (1916) and Meek and Hildebrand (1928) each found only one example of this species. Range: Bermuda, Florida, Bahamas. LITERATURE CITED BOHLKE, J. E. 1956. A synopsis of the eels of the family Xenocongridae (including the Chlopsidae and Chilorhinidae). Proc. Acad. Nat. Sci. Phila- delphia, 108: 61-95. BREDER, C. M. 1925. Notes on fishes from three Panama localities: Gatun spillway, Rio Tapia and Caledonia Bay. Zoologica, 4 (4): 137-158. 1927. Scientific results of the first oceanographic expedition of the “ pe iy: ST EORTOON . } ai 7 Li . 1” neta} | bad ini Ags ii wat wwerevian & Tih BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. NOVEMBER 16, 1962 NUMBER 170 THE BRAIN OF THE EMU (DROMAEUS NOVAEHOLLANDIAE, LATH)? I. Gross ANATOMY OF THE BRAIN AND PINEAL Bopy” By STANLEY COBB and TimLy EDINGER The histology of the cerebral hemisphere of the emu has been extensively studied by Craigie (1935a, 1935b, 1940) and three diagrams of the hemisphere have been published. Drawings of the whole brain have also appeared in the literature (Strong, 1911; Ktenzi, 1918), and a photograph was published by Anthony (1928). No description of the whole brain, however, is to be found. Since the emu is, next to the ostrich, our largest living bird, and since it belongs to a taxonomically controversial group, it seems of value to describe the brain and compare it with the brains of other birds. Moreover, the emu is considered, by Py- eraft (1900) and many others, to be one of the most primitive of birds. The concept of ‘‘primitiveness’’ will be considered in the discussion at the end of this paper. MATERIAL Three specimens of Dromacus novachollandiae were collected by S. J. J. Davies in November 1960 in Western Australia for Professor Ernst Mayr, Director of the Harvard Museum of Com- parative Zoology. Two of them were kindly given to us by Pro- fessor Mayr for neurological study. The heads had been cut off L'This spelling of Dromaeus is not the one accepted by some newer checklists. but because Dromiceius (an alternative spelling) is the perpetuation of a grapho- logical error (Newton, 1896) and because Dromaius is a less proper Latinization, it seems better to use the older form. 2From the Museum of Comparative Zoology and the Department of Neurology and Psychiatry, Harvard University, and the Laboratory for Psychiatrie Research, Massachusetts General Hospital. This investigation has been aided by grants from the Foundations Fund for Psychiatry and the National Institute of Neurological Disease and Blindness, grant #03429-02. 2 BREVIORA No. 170 and skinned, the eyes had been removed, and the specimens had been fixed in 10 per cent formalin solution in the field. After about two months the heads were packed in moist condition and shipped in cellophane bags to the United States. Here the brains were removed from the skulls after making photographs of vari- ous stages of the dissections. The brains were then fixed in fresh neutral formalin solution (10 per cent) for a month. One brain (that of Emu #85) was divided into its component parts for weighing and special histological studies. The other brain (Emu +104) was removed, photographed, fixed for a month in 10 per cent formalin as above, and embedded whole in celloidin for serial sectioning. Both are brains of adult males. The first (#685) appears somewhat larger and weighed 27.7 grams; the second (3104) weighed 25.1 grams. DESCRIPTION The position of an avian brain within the skull is determined by many developmental factors. The most obvious are the shape of the bill, the size and position of the eves, the habitual posture of the bird, and the size and shape of the brain itself. Starck (1955) has given an excellent discussion of these relationships and emphasizes the importance of the size of the eye and the position of the orbit. One way of describing the position of the brain is to measure the angle between the cerebral axis and the axis of the bill (Cobb, 1959). In the emu this angle is about 27° (see Fig. 1), an angle somewhat smaller than that of the gull (Larus argentatus: 34°) and the grouse (Bonasa umbellus: 36° ), but distinctly greater than that of the cormorant (Phalacrocorar auritus: 15°) which has the straightest (most extended) type of skull and an exceptionally small braim-bill angle. 3esides showing the relation of the brain to the skull, Figure 1 shows the olfactory bulb and membranous sae of the olfactory nasal chamber; the bulb seems to be in direct contaet with the chamber, but closer scrutiny shows that there is a space bridged by the short olfactory nerves. When the light, diploie bone of the bill is removed, the sac which forms the lining of the olfactory chamber is revealed. It is a fairly tough structure containing blood vessels and many nerve fibers. It is crossed anteriorly by a branch of the first division of the trigeminal nerve. The main nerve trunk of this division is seen passing through the orbit, close to the optic nerve and up to a point just below the olfactory bulb. 1962 THE BRAIN OF THE EMU 3 This is the main sensory nerve from the bill, innervating the skin and vibrissae. Its large size suggests that tactile sense in the bill is acute and important. On opening the olfactory chamber, the most posterior of the three nasal chambers, a well-developed turbinal mound (superior or olfactory concha) is seen on the lateral wall. It is covered with a soft, yellowish epithelium, which becomes thinner and _ less yellow as it spreads out over the dorsal and mesial aspects of the chamber. A vertical section through the nasal chambers of the bill at this level (Fig. 3) reveals that the concha is raised to. a height of about 5 mm. and is slightly constricted at its base, but is not folded into a spiral like the conchae of some vultures and albatrosses (Bang, 1960). A specimen for microscopic examina- tion was taken from the dorsal surface of the olfactory chamber ; it shows cells and cilia typical of olfactory epithelium. About 5 em. anterior to the olfactory concha there is a large nostril (Fig. 1), which is the external opening of the anterior nasal chamber. The emu has large eves and the orbits are spacious. As one sees In Figure 1 the brain hes mostly behind the orbit with the olfactory chambers in the bony structures just in front. The optic nerve enters the chiasm and passes directly to the optic lobe of the opposite side of the midbrain (Fig. 2C). The large fascicles of nerve fibers can be seen as they cross. The optie lobe is a large and conspicuous structure (Figs. 1,2B and 2C). In the lateral view only about one-fifth of it is covered by the overlying hemisphere. In Figure 2B (in which the parts of the brain are slightly separated) the relation of the optic lobe to hindbrain and forebrain is emphasized. It is clearly a part of the midbrain. In fact, the optic lobes are homologues of the corpora bigemina of reptiles, and of the anterior corpora quadrigemina of mam- mals. They have taken a ventrolateral position in birds, perhaps because it was easier there to make room for the extraordinary tectal development in this class of vertebrates. The emu brain when viewed from above (Figs. 1A and 2A) impresses one by its triangular shape, with cerebral hemispheres broad posteriorly and narrow anteriorly. The olfactory bulbs protrude, forming the anterior pole of the hemisphere. On the vertex the two sagittal elevations of hyperstriatum stand out con- spicuously and are separated from the lateral parts of the hemis- pheres by a distinct sulcus, the valleeula (Portmann and Stinge- Tima, LO Gis). 4 BREVIORA No. 170 The cerebellum is larger in comparison to the forebrain than in passerine birds. It has a greater diameter dorsoventrally than laterally (Figs. 1 and 2) although the auricles protrude laterally on each side. These lobes, composed of flocculus and nodule, are the only ones that complicate the simple conformation of the cerebellum, the corpus cerebelli being largely a mid-line organ corresponding to the vermis of mammals. Between the anterior surface of the cerebellum (culmen and declive) and the posterior poles of the cerebral hemispheres there is ample space for the pineal stalk and gland. The lateral view of the brain (Fig. 2B) shows the relative sizes of the main subdivisions. For this photograph the forebrain, midbrain, and hindbrain were slightly pulled apart. The hemi- spheres of the forebrain are well developed and extend backwards covering parts of the optic lobe and of the cerebellum. The great- est diameter of the hemisphere is 36 mm. and the greatest diam- eter of the olfactory bulb is 9 mm., giving a ratio of 4 to 1 or 25 per cent. This places the emu among those birds that have large olfactory bulbs (the Gruiformes, Caprimulgiformes, Procel- laruformes, Podicepidiformes, and Apterygiformes). In a list of 47 different species of birds, arranged according to the relative size of the olfactory bulb, the largest at the top, the kiwi would come first and the emu seventh (Cobb, 1960). The anterior end of the hyperstriatum accessorium (sagittal elevation or Wulst ) is close to the olfactory bulb, and the posterior end shades off into the neostriatum before reaching the occipital pole of the hem- isphere. Thus the emu has a large Wulst that reaches well back towards the occipital pole (Figs. 1 and 2) and well forward to a point close to the olfactory bulb. A comparison of the external configuration of the brain of the emu with that of other birds shows that it resembles most some herons and ducks. In comparing it with Stingelin’s (1958) photographs, it is seen to be strikingly similar to the brain of Txrobrychus minutus (see his fig. 21 ‘‘Zwergreiher’’). Seen from below (Fig. 2C) the conspicuous characteristies of the emu brain are: (1) the large, separated olfactory bulbs, form- ing the anterior pole; (2) the flatness of the ventral aspects of the lateral parts of the two cerebral hemispheres; and (3) the pair of big optic lobes shaped like flasks with their necks joined in the optic chiasm. The cerebellum is so narrow that it is almost hidden by the medulla oblongata, only the flocculi showing on each side. The roots of the third, seventh, eighth, ninth, and tenth cranial nerves show in this view. 1962 THE BRAIN OF THE EMU D The brain of emu #85 (after formalin fixation) weighed 27.7 grams; his body weight was 34 ke. The brain of emu #104 weighed 25.1 grams (also after formalin fixation) ; body weight 31 kg. This gives a ratio of brain weight to body weight in emu #85 of 1/1227 and in emu #104 a ratio of 1/1235. Little sig- nificance, however, should be given to these ratios because it is known that a living emu may vary 30 to 40 per cent in weight during a year due to conditions of food, climate and water supply. The first brain was separated into 8 pieces, for weighing, as follows : Olfactory bulb (right) (injured ) Olfactory bulb (left) 0.12 grams Cerebral hemisphere (right ) 8.85 grams Cerebral hemisphere (left ) 8.8 grams Optic lobe (right ) 7.3 grams Optic lobe (left ) 7.3 grams Cerebellum 4.6 grams Brainstem 3.9 grams The brainstem (defined by Portmann, 1946, and named ‘“Stammrest’’) is the basal mass of nerve tissue made up of thalamus, midbrain (with optic lobes removed) and hindbrain (with cerebellum removed). Portmann’s purpose was to choose as his common denominator that part of the brain which varies least in its size relative to the size of the whole bird. That part is obviously the brainstem. He then compares its size to other parts of the brain and, by dividing the weight or volume of the stem into the coresponding value for another part, he obtains his index. This ‘‘index of cerebralization’’ he finds for an emu to be 4.18, obtained by dividing the weight of the ‘‘Stammrest’’ into the combined weight of the two hemispheres. In our emu #85 this index is 17.6/3.9 = 4.5. According to Portmann’s list the figures 4.18 and 4.5 both place the emu far below parrots and ravens, but above loons, grebes, and quail. He believes that this quotient gives an expression of the ‘‘level of integration’’ of the brain for each species. THE PINEAL Bopy In the dissection of emu #104, a large part of the post-central area of the calvarium was left intact and carefully lifted off the brain. The pineal stalk was thus torn away at its attachment to the diencephalon. It is 10 mm. in length and remained attached 6 BREVIORA No. 170 to the pineal body (Fig. +). The body itself is embedded in the dura and lies in a depression of the cranial roof between the an- terior and posterior fossae. The dorsal position of the epiphysis is thus clearly demonstrated ; it hes between cerebrum and cere- bellum at the level of their dorsal surfaces. The stalk leaves the brain at a point just rostral to where forebrain joins midbrain. The pineal body is round and firm, slightly flattened dorso- ventrally. It is vellowish in contrast to the white skull. The fibrous envelope is continuous with dura which has strong bands spreading laterally and anteriorly. Removed from the mem- branes, the pineal body is roughly triangular, 7 mm. long on each side. With stalk attached, it weighs 0.1 em. after formalin fix- ation. DISCUSSION The description of the gross anatomy of the brain of the emu brings up five points for discussion: 1) the size of the brain, 2) the question of primitiveness, 3) the general shape of the brain in relation to the base of the skull, 4) the size and position of the Wulst, and 5) the topographic relations of the pineal body. The size of the brain in relation to body size and ‘‘intelligence”’ has been the subject of much study and many pronouncements. Suffice it to say here that in our opinion the relation of brain weight to body weight (so called cephalization) is a ratio too simple to give information of much significance. Portmann’s (1952) pioneer work in describing an index of encephalization is an advance in the right direction. Body weight in birds is too erossly variable to be used in comparison to the much more stable brain weight. Small birds may show rapid and marked change in weight. There is good evidence that some birds may lose from 30 to 50 per cent of their body weight in 24 hours during a mi- gratory flight (Odum et al., 1961; Helms and Drury, 1960). The emu, being flightless, lives in a fairly uniform environment and does not go through the prolonged exertion of migratory flights. Its ratio of brain weight to body weight might, therefore, be relatively stable. Actually, in Dromaeus novaehollandiae this ratio is approximately 1/1230 (see p. 5). From the weights given by Crile and Quiring (1940) we deduce that the ratio for an ostrich (Struthio camelus massaicus) is 1/2929; for a sparrow (Passer domesticus) it is 1/23; and for a hummingbird (Amazilia teacatl) it is 1/24. This does not mean that the hummingbird has a ‘‘better’’ brain than the emu. It merely indicates that the —~! 1962 THE BRAIN OF THE EMU body controlled by the brain of the hummingbird is just as com- plex as the body of the emu, though much smaller. The question as to which brain is ‘‘better,’’ or more highly evolved, is mean- ingless unless one asks, ‘‘ Better for what ?’’? Obviously, the hum- mingbird’s brain is better for flight and the emu’s better for running. Another factor relative to brain size must be considered. It has been pointed out by Sholl (1956) that small brains are in general more closely packed with nerve cell bodies than large brains which have more glial structures between neurons. Man has 10.5 nerve cell bodies per cubie micron; a mouse has 142.5. In short, the need is to learn what parts of the brain, control- ling what organs, are larger or smaller in each family of birds. With more investigation into the quantitative anatomy of the brain, some of these questions may be answered. Fritz (1949) has estimated the volume of four parts of the striatum in four different species of birds; he found significant differences, but no correlation with Portmann’s cerebral index. Many authors have spoken of the emu, and in fact all ratite birds, as primitive, but their concept of primitiveness is not clear. Some seem to call these birds primitive because they are flightless and have no keel on the sternum (Leach, 1923), others because they have a straight type of skull base (Streckschadel) (Mari- nelli, 1928, p. 156). Stingelin (1958) considers those birds, with a small Wulst which lies neither far forward nor far back, to be the less evolved type. The point would seem to be that one must not apply the term primitive in a general way to the emu (or probably any other bird). One should specify in what respect a given type or family is less evolved (‘‘primitive’’) and in what respect it is more evolved (specialized). Even then, the gaps in our phylogenetic knowledge do not allow us to say whether the ratite sternum is due to a devolution from carinate ancestors or an evolution from cursorial reptiles. The presence of feathers and the avian type of brain suggest strongly a descent from flying ancestors. In respect to running and adaptation to hfe in open plains one feels confident in saying that the emu is highly evolved. Much work has been done on the development of the avian skull. Pertinent to an understanding of the shape of the emu’s brain are three recent lines of investigation. Duym (1951) deseribed the bending of the base of the skull in different birds and specified four types—the stretched or extended type of 8 BREVIORA No. 170 skull and three degrees of bending. Dullemeijer (1960) has re- lated the shape and size of the principal parts of the brain to the amount of bending of the cerebral axis and has described four classes on this basis: 1) stretched skulls with httle bending, 2) bending of 20 degrees, 3) bending of about 70 degrees, +) bending of about 120 degrees. Starek (1955) has shown the great im- portance of the size of the eye and orbit in determining the shape of the brain and skull. In the emu one finds a rather extended type of skull and a very large eye and orbit. Our measurements show that there is an angle of about 27 degrees between cerebral axis and bill axis, and a bending of the cerebral axis of about 13 degrees. Thus the emu falls into class ‘‘1’’ in Dullemeijer’s grouping. We agree with him in emphasizing that ‘‘the position and shape of the brain parts is influenced by the position of the bill and the posi- tion and size of the eye.’’ The development of the bill in birds has been remarkably variable, and with these special develop- ments come variations in the bones of the skull and in the con- formation of the brain. As mentioned on page 4, the general shape of the brain of the emu resembles that of Irobrychus but shape in itself is not very significant phylogenetically or physiologically in comparing birds’ brains. One feature, however, may be of interest: the size and position of the Wulst, because this ganghon is conspicuous on the surface of the cerebral hemisphere and because its size in relation to that of the hemisphere appears to vary. Unfortunately, however, there are not enough quantitative data concerning the size of the Wulst in various types of birds to make any statements about its significance. In the lateral and dorsal views of the emu’s brain the Wulst is conspicuous (Figs. 1 and 2). Its anterior end is almost in contact with the olfactory bulb. The posterior end reaches back to within 4 mm. of the occipital pole of the hemisphere. This rounded ridge is long and les parallel to the interhemispheric fissure — hence, the name used by L. Edinger et al. (1903) ‘*Sagittalwulst,’’ translated as ‘‘sagittal elevation’’ by Portmann and Stingelin (1961). Its position in the emu is like that in the pigeon, a bird whose brain resembles Stingelin’s (1958) ‘* Grund- typus.’’ But the Wulst of the emu in relation to the rest of the hemisphere is both longer and broader than that of the pigeon. Stingelin emphasizes the importance of the position of the Wulst. In his chapter entitled ‘‘Comparison and extent of stri- atal fields’’ there is a comparative description of the striatal 1962 THE BRAIN OF THE EMU i) ganglia in 18 species, with clear diagrams of each. He believes that in the ‘‘highly evolved’’ (‘‘hochevolutiert,’’ p. 38) species there is a tendency to marked frontal enlargement of the hem- isphere. This is achieved in two ways. In developmental line A the frontal pole is largely Wulst, the rostral end of which is in contaet with the olfactory bulb. In developmental line B the frontal pole is made from the neostriatum and ventral hyper- striatum, the Wulst having receded to a position on the vertex by successive caudal shifts. From these observations Stingelin deduces a morphological rank (‘‘Formwert’’) in relation to the basic type (‘‘Grundtypus’’). In developmental line A, crows and owls are considered the more highly developed groups; in line B, the higher ones are snipe, spoonbill (Platalea) and parrot, with a plover considered as ‘‘lower’’ and the lapwing as ‘‘middle.”’ This rank order seems to be entirely based on cerebral anatomy. Reference is made neither to fossils nor to other characteristics such as brain axis or anatomy of skull. As descriptions of the different relationships of one ganghon of the brain to another, the figures and exposition of Stingelin have great value, but taken as indicating evolutionary levels they may be misleading. Until one knows the lines of descent from reptilian and avian ancestors, descriptions of such ** Entwick- lungsgerichtungen’’ and levels of evolution with “‘hoher’’ and ‘‘niederer Formwert’’ are not justified because evolution is the process of phylogenetic transformation, a phenomenon that can- not be observed except in consecutive phases of an ancestral line. The ‘‘ Horizontalmodifikationen’’ (Stingelin’s fig. 32) are cer- tainly of interest as showing differences between the brains of living families of birds, but these modifications are not a basis for conelusions concerning evolutionary ancestry. In short, we doubt if any living bird has a conformation of the brain that we are justified in calling primitive. The data for making such a judgment are inadequate. On the other hand, research into the relative size of various parts of the brain, such as Fritz (1949) has done in Portmann’s laboratory, may give us important leads as to the degree of cerebral developments. Such investigations would be especially useful if correlated with behavior. We wish to emphasize the possibility of drawing erroneous con- clusions when the anatomy of living forms is used as evidence for describing an evolutionary process, disregarding the evidence from fossils. We welcome the opportunity to point to one feature of the emu brain as a graphic argument against a persistent, but erroneous theory. This feature is the position of the pineal body, 10 BREVIORA No. 170 so obvious when one looks at the dorsal surface of the brain. The theory is that absence in birds and mammals of a second epi- thalamie appendage in the pineal area, the ‘‘parietal eye’’ of reptiles, is due to enlargement of cerebrum and cerebellum in the two descendant classes. That concept supposes that the covering over of the diencephalon and mesencephalon by the cerebrum and cerebellum in Aves and Mammalia obstructed the access of epi- thalamic appendages to the brain surface. This process is be- lheved to have caused devolution and loss of the predominantly sensory, stalked, second organ in the pimeal complex which sur- vives only in lizards and the tuatara, the parietal eye. Actually, among birds, a pineal organ reaching to the level of the cerebral and cerebellar vaults, attached to the skull roof, is not an exceptional occurrence. This is found in our Dromaeus and has been previously reported in brains of other ratites (Struthio, Rhea, Apteryx) and, as a button-shaped convexity, on the endocranial cast of the extinet Dinornis (Starck, 1955). The difference between these recently and carefully prepared speci- mens and those figured in the literature is not a real difference, but a matter of preparation. Kitienzi’s (1918) diagrammatie figures of the brains of 36 dif- ferent kinds of birds give the impression that no bird possesses an externally visible pineal organ, as do almost all macroscopic fig- ures of avian brains in the literature, including Strong’s (1911) figures which show Dromaeus. Kiienzi, however, mentions in several places (pp. 28, 52, 89) that the pineal body is too firmly embedded in the meninges to be removed with the brain; he re- ports (pp. 70-71) that the pineal body in all birds studied ocecu- pies a median space between the posterior borders of the hemi- spheres and the front end of the cerebellum, its distal end reach- ing approximately to their dorsal level. Our observations on Bubo, Corvus, Gallus, Columba, and Larus agree with those of Ktienzi; all have pineal bodies extending into the dura. A recent study on ten embryonic and three later developmental phases of Larus (Wetzig, 1961) also clearly testifies against the theory of mechanical suppression in birds of the second (the stalked) organ in the pineal complex of reptiles. The epiphysis extends to the level of the prospective skull roof in an early transitory phase, and again in the last phases of embryogeny. It is then, and re- mains in the adult, fitted into the space between cerebrum and cerebellum. It is club shaped, its apex coalesced with the dura in contact with the roof of the skull. 1962 THE BRAIN OF THE EMU 11 Thus, there is more space available than was assumed. The club shape (with the largest circumference distal) suggests that the form is governed by the space available. While we are well aware that the shape of a predominantly glandular organ (Stam- mer, 1961) has no great significance, we do wish to draw atten- tion to the shape of the epiphysis of the emu. When, as in our emus, the distal expansion is abrupt, its connection with the corpus diencephali a mere stalk (Fig. 4), the avian pineal body strikingly resembles not the reptilian pineal organ, which is more or less sessile, but the reptilian parapineal vesicle with its nervous and vascular stalk —that is, the parietal eye. Krabbe (1961) observed this similarity in an embryo Cygnus. Many species of birds similarly demonstrate that there is no obstruction to the development of a parietal eye. The reason for the absence of a parietal sense organ in birds is obviously not mechanical suppression; it is to be found in the fossil record, namely in pre-avian phylogeny. The presence of a parietal eye is reflected in a corresponding foramen of the roof of the skull. The fossil record of skulls plainly shows that the organ was first present and then lost in innumerable phyletic lines within the classes Pisces, Amphibia, and Reptilia. Birds are an offshoot of the great reptilian subclass Archosauria. Amone the many hundreds of known skulls from its various orders, only two have the parietal foramen. Significantly, both the specimens showing that heritage from Palaeozoic ancestors belong to the earliest forms identifiable as archosaurian, each representing the beginning of a suborder of the order Thecodontia, ‘‘stem archo- saurs’’ (Mesorhinus: Jaekel, 1910, and Erythrosuchus: Huene, 1911). These openings in the parietal bones of Archosauria oc- curred for the last time at the beginning of the Mesozoic era in the earliest Triassic times. It follows that parietal eyes were lost, not within the evolution of birds, but in remote reptilian ancestors some 80 million years before the first, late Jurassic, appearance of feathered animals, and presumbaly more than 100 million years before the modern type of avian brain was evolved. In the Mammalia, likewise, both recent and fossil conditions plainly contradict the assumption that possession of a parietal photoreceptor became impossible because of progressive brain evolution. In various groups of mammals now living, much or all of the midbrain is dorsally exposed in a gap between cerebrum and cerebellum. Actually, it has long been known that in some bats, lagomorphs, and rodents the pineal gland extends into the 12 BREVIORA No. 170 dura mater at the caudal end of the interhemispheric fissure, and this condition has now been found prevalent in studies of a large material from a great number of genera (Pilleri, 1960). The usual absence of a pineal organ in specimens of the rabbit brain is the result of its having been torn off with the tentorium during preparation. Furthermore, endocranial casts show that in most early Tertiary mammals there was a considerable gap between cerebrum and cerebellum. The parietal foramen was obliterated, i.e. the parietal eye had been lost in a pre-mammalian phase of evolution, in this case, in mammal-hke Reptilia shortly before the emergence of the new elass. SUMMARY A description of the gross anatomy of the brain of Dromaeus novaehollandiae is presented on the basis of two specimens from Western Australia. The brain is of the extended type. The olfactory bulbs and sagittal elevations of the forebrain, and the optic lobes of the midbrain are comparatively large. The index of encephalization is 4.5. The brains weighed 27.7 and 25.1 orams, respectively. The pineal body lies in a shallow fossa in the roof of the skull and weighed 0.1 gram. It is pointed out in discussion that there is no good reason for considering this brain to be primitive, and that phylogenetic relationships cannot jJus- tifiably be deduced from the anatomy of the brains of living birds. LITERATURE CITED ANTHONY, R. 1928. Lecons sur le cerveau. G. Doin et Cie (Paris). Austin, ©. lL. 1961. Birds of the world. Golden Press (New York). Bane, B. G. 1960. Anatomical evidence for olfactory function in some species of birds. Nature (London) 188: 547-549. Cops, S. 1959. On the angle of the cerebral axis in the American woodecock. Auk 76: 55-59. 1960. A note on the size of the avian olfactory bulb. Epilepsia 1: 395-402. CRATGING He al 1935a. The cerebral hemispheres of the kiwi and of the emu (Apteryx and Dromiceius). J. Anat. London 69: 380-393. 1962 THE BRAIN OF THE EMU 13 1935b. The hippocampal and parahippocampal cortex of the emu (Dromiceius). J. comp. Neurol. 61: 463-591. 1940. The cerebral cortex in palaeognathine and neognathine birds. J. comp. Neurol. 73: 179-234. CrILE, G. and D. P. QUIRING 1940. A record of the body weight and certain organ and gland weights in 3690 animals. Ohio J. Sei. 40: 219-259. DULLEMEIJER, P. 1960, Shape and size of the brain parts as architectonic factors in the skull of birds. Acta morphol. neerl.-scandin. 4: 96. Duym, M. 1951. On the head posture in birds and its relation to some anatomical features. Proc. Kon. Ned. Akad. Wetensch. (C) 54: 202-211, 260-271. EDINGER, L., A. WALLENBERG and G. HOLMES 1908. Untersuchungen tiber das Vorderhirn der Vogel. Abh. seneken- berg. naturf. Ges. 20: 341-426. EDINGER, T. 1961. Fossil brains reflect specialized behavior. World Neurol. 2: 934-941. FRITZ, WALTER 1949. Vergleichende Studien tiber den Anteil von Striatumteilen am Hemispharenvolumen des Vogelhirns. Rev. suisse Zool. 56: 461-491. Hrums, C. W., and W. H.. Drury, JR. 1960. Winter and migratory weight and fat field studies on some North American buntings. Bird Banding 31: 1-40. HUENE, F. v. 1911. Uber Erythrosuchus, Vertreter der neuen Reptil-Ordnung Pely- cosimia. Geol, Pal, Abh. 14: 1-60. J AEKEL, O. 1910. Uber einen neuen Belodonten aus dem Buntsandstein von Bern- burg. Sber. Ges. naturf. Freunde Berlin: 197-229. KRABBE, K. H. 1961. La glande pinéale. World Neurol. 2: 94-102. KUENZI, W. 1918. Versuch einer systematischen Morphologie des Gehirns der Vogel. Rey. suisse Zool. 26: 17-111. Ibanvha, dh, Xo 1923. An Australian Bird Book. Whitcome and Tombs (Melbourne). MARINELLI, W. 1928. Uber den Schiidel der Schnepfe. Palaeobiologica 1: 135-160. Newton, A. et al. 1896. Dictionary of Birds. Adam and Black (London). Opum, E. P., C. E. CONNELL, and H. L. STopDARD 1961. Flight energy and estimated flight ranges of some migratory birds. Auk 78: 515-527. 14 BREVIORA No. 170 PILLERI, G. 1960. Beitrige zur vergleichenden Morphologie des Nagetiergehirns. Acta Anat. 42: 1-88. PORTMANN, A. 1946. Etudes sur la cérébralisation chez les oiseaux. Alauda 14: 1-20. 1952. Die allgemeine biologische Bedeutung der Cerebralisations- Studien. Bull. schweiz. Akad. med. Wissensch. 8: 253. PoRTMANN, A. and W. STINGELIN 1961. Biology and Comparative Physiology of Birds. Marshall, A. J. (ed.). Academic Press (New York), volume 2: 1-36. PYCRAFT, W. P. 1900. On the morphology and phylogeny of the Palaeognathae (Rati- tae and Crypturi) and Neognathae (Carinata). Trans. Zool. Soe. London 15: 149-290. SHOLL, D. A. 1956. The Organization of the Cerebral Cortex. Methuen and Co. (London). STAMMER, A. 1961. Untersuchungen tber die Struktur und die Innervation der Epiphyse bei Vogeln. Acta Univ. Szeged., Acta Biol. N.S. 7: 65-75. STARCK, D. 1955. Die endokraniale Morphologie der Ratiten, besonders der Aptery- gidae und Dinornithidae. Morph. Jahrb, 96: 14-72. STINGELIN, W. 1958. Vergleichend-morphologische Untersuchungen am Vorderhirn der Vogel auf cytologischer und cytoarchitektonischer Grundlage. Helbing and Lichtenhahn (Basel). Srrone, R. M. 1911. On the olfactory organs and smell in birds. J. Morphol, 22: 619-658. Wetzic, H. 1961. Die Entwicklung der Organe des Zwischenhirndaches (Epiphyse und Plexus choroideus anterior) bei der Sturmmove, Larus canus, L. Morph. Jahrb. 101: 406-431. THE BRAIN OF THE EMU 1962 Bee) ST OL sue [tq - Ulead *‘[PNyYs FO ISLC WO nyts ul ‘Tuo JO ULL JO MOLA [BLdPET FO SUIMBIG Vall ONG TY “© wo Gl Zl 6 9 ¢ fe) ea a a a Oa Carle ae | Pa ad (ees ee oe Pe] ee 9qo7 214d0 aAdAN Ido AAAON |OUIWabBlay bYO™ JTONV 1118 —Nivud —_——_—- ql aliite SCY Y/ a90dS |04!q10 ajouny tp” wn|jaqaiad rast MG SS aiaydsiway \ Se qing \ ~ ISIN AKsOpods]O \ TAS saqwoyy) ~—. A104905/0 ~ 16 BREVIORA No. 170 Figure 1B. Photograph, life size, of dorsal view of same dissection. 1962 THE BRAIN OF THE EMU WA FIG 24 DORSAL FIG 2° LATERAL FIG 2° VENTRAL Figure 2. Three views of the brain of the emu (Dromaeus novaehollan- diae) #104, Life size. F, forebrain hemisphere, H, hindbrain, M, mid- brain, showing optic lobe (7) and optie chiasm (C), V’, vallecula, W, Wulst or hyperstriatum accessorium. 18 BREVIORA No. 170 FIG 3 FIG 4 Figure 3. Section through the nasal chambers cut in frontal vertical showing the olfactory (or posterior) chamber (P) into which protrudes the olfactory concha (C) covered with yellow olfactory epithelium. Below is seen part of the middle nasal chamber (M). The two chambers are divided by the septum. They connect anteriorly with the anterior chamber and the external nostril. Life size. Figure 4A. Photograph, life size, of pineal body (PB) lying in shallow cavity of the calvarium, posteroanterior view. The stalk (S) protrudes downward. Figure 4B. Ventral view, looking upwards at under surface of calvarium. The stalk (S) is bent backwards. ra) BREVIORA Museum of Comparative Zoology CAMBRIDGE, MAss. DECEMBER 14, 1962 NUMBER 171 NOTES ON AMPHISBAENIDS (AMPHISBAENIA; REP- TILIA), 6. REDESCRIPTION AND RANGE EXTENSION OF AMPHISBAENA SPURRELLI BOULENGER. By CarL GANS Department of Biology, The University of Buffalo, Buffalo 14, New York In 1915 Boulenger (p. 659) described the new species of Amphisbaena spurrelli, characterized primarily by the presence of tubercular or subconical segments on the dorsal surface of the tail. The two syntvpes were collected at ‘‘Anda Goya [Colombia], at the junction of the R. Condoto and San Juan.’’ The only subsequent record of the species (Burt and Burt, 1931, p. 40) is the citation of a single specimen (A.M.N.H. 18261) from the neighboring locality of ‘‘Boca de la Raspadura,’”’ Colombia, without supplementary description. The present note is based upon a re-examination of these three and of two additional specimens, one of which extends the range of A. spurrelli into Panama. The original description has been amended and rewritten according to the standard pattern (Gans and Alexander, 1962). Simple, non-idealized illustrations are included in the present paper. It is a pleasure to acknowledge the support of the National Science Foundation (NSF G-9054, G-21819). Examination of the types was made possible by assistance from the estate of Leo Leeser. Specimens were examined through courtesy of C. M. Bogert, The American Museum of Natural History (A.M.N.H.), Miss Alice G. C. Grandison, British Museum (Natural History) (B.M.), R. F. Inger and H. Marx, Chicago Natural History Museum (C.N.H.M.), and E. E. Williams, Museum of Comparative Zoology (M.C.Z.). I am particularly grateful to Dr. Federico Medem who made the C.N.H.M. speei- men available, and to Miss C. Rhodes for technical assistance. 2 BREVIORA No. 171 AMPHISBAENA SPURRELLI Boulenger, 1915. Amphisbaena spurrelli Boulenger, 1915, p. 659. Terra typica: ‘‘ Anda Goya, at the junction of the R. Condoto and San Juan,’’? Colombia. LECTO- TYPE: B.M. 1915.10.21.9 (by present designation). PARATYPE: B.M. 1915.10.21.8. Diagnosis: A form of Amphisbaena without fusions of head scales; with 4+ oval [not round| precloaeal pores; and with the dorsal and lateral surfaces of the caudal tip covered with conical or tubercular segments. Specimens have 218 to 222 body annuli; 18 to 20 caudal annuli; 16 to 18 dorsal and 16 to 18 ventral see- ments per midbody annulus; and one row of postgenial and one row of postmalar chin shields. There is no visible autotomy con- stricton of the tail. Autotomy takes place after the seventh annulus. Notes on the types: Boulenger (1915, p. 659) illustrated the smaller of his syntypes, which has here been chosen as a lecto- type. The types, still extant and in good condition, suggest that his illustrations were idealized, and several of his counts [shown S RASPADURA SS ANDAGOYA 6 Fig. 1. Amphisbaena spurrelli. Map showing localities mentioned in text. Anda Goya and Boea de la Raspadura are actually closer together than can be indicated on a map drawn to this seale. 1962 AMPHISBAENA SPURRELLI 3 in brackets in the table] erroneous. The errors do not affect the validity of the species. Description: Meristic characters are listed in the table. Figure 3 shows the head sealation, Figure 4 the segmentation of cloaca and tail, Figures 5 to 8 photographs of head, midbody pattern and tail. Preserved specimens are various [faded] shades of brown dorsally, somewhat lighter ventrally. The darker dorsal color is in part produced by a darkening of the rectangular center of each segment, the contrast with the hghter segmental margins giving the impression of dark spots. The fully dark dorsal spots descend the sides to approximately the third ventral below the lateral groove on each side. Ventrad from this the dark center shrinks drastically or may fade out entirely. The anterior fifth (M.C.Z. 39784), or the head alone, lacks the dark colored seg- ments. The head sealation shows some variability and no major fu- sions. An azygous rostral barely visible in dorsal view is fol- lowed by three pairs of enlarged cephalic shields in contact along the dorsal midline. The nostrils pierce the first pair (nasals). The second pair (prefrontals) are the largest segments of the head. There are three supra- and two and a half infra- labials, as the third infralabial extends considerably beyond the angulus oris. The supralabials are large, the second much the largest. The C.N.H.M. specimen has this segment subdivided differently on both sides. The second infralabials are the largest segments on the lower jaw. Small segments lie beyond the angulus oris in the position of fourth supralabials. The mental is shaped like a truncated wedge with a posteriorly con- vex tip. The postmental is hexagonal and elongate. It les in lateral contact with the medial edges of the second infralabials, as well as the anterior portion of the medial edges of the rela- tively short, wedge-shaped malars. There are one to two rows of postgenial segments, followed by a single postmalar row, the lateral segments of which are shghtly enlarged. The head is relatively blunt, flattened slightly dorsoventrally and oval in cross-section. The lower jaw is but shghtly shorter than the upper. The sides of the head would, if extended, inter- sect some distance anterior to the rostral tip, even in adults 4 BREVIORA No. 171 27 * 25 E Er io3 <£ e)) ce Cc be le 5 Je] ® :S —_ 19 ° 17; 20 22 24 26 28 30 Snout - Vent Length - cm Fie. 2. Amphisbacna spurrelli. Seatter diagram showing plot of tail length versus snout-vent length for the available specimens. The lectotype is shown as a hollow cirele. in which the bulge of the temporal musculature changes the out- line. The attachment of the skin to the crest of the skull pro- duces a coneave dishing of the interfrontal suture, particularly in adult specimens. The first body annulus curves forward to contact the frontals. Its dorsalmost segments may be somewhat enlarged, and one of the specimens has an intercalated dorsal half-annulus. The second through fifth annuli are narrowed, and the fourth marks the level of the head joint or the point at which the bulge of the temporal musculature returns to normal. 1962 AMPHISBAENA SPURRELLI 5 Fig. 3. Amphisbaena spurrelli. Dorsal, lateral and ventral views of the head of A.M.N.H. 18261 from Boea de la Raspadura, Colombia. The line equals 1 mm to scale. (V. Cummings, del.). No. 17 BREVIORA POET] 2222 CCCCLORDD) YY» CCRKCATDD YD)» CCKCEABDIDDIYD)) CCACEAAD)D DDD) Sa CCCCECCADDD DDD) DDD CAMutennapyasee, (COOCCECIDE (Ca CCC TF) CCCI) (CCCACURED) ((CaCCCAE TD) MCU TY] WUC LTT MMU Wee ese WO WG COU RT COCCeCanay: OT TTT Td cceaaneuss: A Colombia. The ventral a double row of and ventral views of the taspadura, Cummings, del.). The middorsal segments are (Ove d by fine Soca de la I 4 Il de Dorsal, lateral 4 61 from | » 18 = and are we — a —S The line equals 1 mm to seale. —— Amphisbaena spurrelli. 4, lateral grooves start about one and a half head lengths be- The dorsal groove is only indicated on the head. roove is indicated mainly as a gap between aligned segments. The Fig. cloaca and tail of A.M.N.H, almost twice as long as wide, the midventral segments are almost Note the extent of cone formation on the tail and the flattened, slit-shaped twice as wide as long. hind the head joint, triangular segmental fragments. precloacal pores, oO =) -I 1962 AMPHISBAENA SPURRELLI The oval precloacal pores lie in a single uninterrupted row of normal sized segments anterior to the precloacal shield. The precloacals are characterized by a central group of four some- what elongate segments. The postcloacals, slightly greater in number, characteristically have a set of two to four midventral and enlarged segments and, flanking these, several split segments entering the cloacal sides. The cloaca may be entirely prolapsed. The tail becomes gradually wider posterior to the cloacal slit and somewhat higher as well. The ventral surface appears plane and an extension of the precloacal region. The terminal third of the tail shows reduction, with the tip about twice as high as wide. The segments of the dorsal and lateral surfaces are stronely tuberculate or cone-shaped. This character facilitates diagnosis of specimens with intact tails. Caudal autotomy takes place behind the seventh posteloacal annulus (cf. Vanzolini, GSE pe 25)! Range: Lowland river valleys of northwestern South America, from extreme northern Colombia (Choco) to southern Panama. Distribution records: COLOMBIA: Choco Province: Anda Goya, mouth of Rio Condoto (Boulenger, 1915); B.M. OP Ae* CATA DYE 19S T0219 (EH CLOLYVE HN): C.N.H.M. 130988 [E. R. Dunn? leg. per F. Medem]. Boea de la Raspadura (Burt and Burt, 1931) ; A.M.N.H. 18261. PANAMA: Tucuti branch, Tuira River [H. C. Clark, leg.| M.C.Z. 39784. LITERATURE CITED BOULENGER, GEORGE ALBERT 1915. Descriptions of a new Amphisbaena and a new snake discovered by Dr. H. G. F. Spurrell in southern Colombia. Proc. Zool, Soc. London, (1915), pp 659-61. Burt, CHARLES E. AND MAy DANHEIM Burt 1931. South American lizards in the collection of the American Mu- seum of Natural History. Bull. Amer. Mus. Nat. Hist., vol. 61, art. 7, pp. 227-395. GANS, CARL AND ALEXANDER ALLAN ALEXANDER 1962a. Studies on amphisbaenids (Amphisbaenia, Reptilia), 2. On the amphisbaenids of the Antilles. Bull. Mus. Comp. Zool., vol. 128, no. 3, pp. 65-158. VANZOLINI, PAULO EMILIO 1951. Amphisbaena fuliginosa, Contribution to the knowledge of the Brasilian lizards of the family Amphisbaenidae Gray, 1825. 6. On the geographical distribution and differentiation of Amphisbaena fuliginosa Linné. Bull, Mus. Comp. Zool., vol. 106, no. 1, pp. 1-67. BREVIORA No. 171 oe Fie. 5. Amphisbaena spurrelli, Lateral view of the head of the topotype, C.N.H.M. 180988, from Anda Goya, Colombia. Note the irregular sub- division of the second supralabial. 1962 AMPHISBAENA SPURRELLI 8) Fig. 6. Amphisbacna spurrelli. Dorsal (left) and ventral (right) views of A.M.N.H. specimen at midbody. Note the darkening of the dorsal seg- mental centers. Fic. 7. Amphishaena spurrelli. View of caudal tip cross-lighted to empha- size the knobbed nature of the terminal segments. 10 BREVIORA No. 171 Fic. 8. Amphisbaena spurrelli. Ventral view of the cloaca and tail of the A.M.N.H. specimen. Note the onset of autotomy at annulus seven, also the contrast between the plane ventral and conical laterodorsal caudal segments. qs AMPHISBAENA SPURRELLI 1962 6I+203 o1-9-F 1-8-3 81/91 0G G/F 136 Bot+0Es €1-9-F orp) 91/91 61 ¢/F GGG [gt/stl 83 [FIs] Got8Ge 0-F L-3 SI/81-91 SI G 81 [sSa+ozc¢ [st/9tl [Tz] [tz] LO+L66 9-F LE-§ 61/81 61 if 613 Io+c33 G1L-9-F L-% SI-L1/9T 61 i GG qysuo'yT 2)VOT,) “ULdOG UL, ) “Ud A /*S10(] [LQ ‘7eT Xpogd SLNAWDAS L'IONNV Yjatinds nuavgsiydup Jo suouttoods day 1oF VAL F8L6E ZOW 8860ET INN YD | GL6L ‘teduojnog | GT OVST6L Wa [G16 ‘tesuo;nog | SIS OUsit6l Wa [9c8T HN’ V LOqunu pur WOT}Iod|[OL) BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 14, 1962 NuMBER 172 A NEW SPECIES OF THE RODENT PIPESTONEOMYS FROM THE OLIGOCENE OF NEBRASKA By Raymonp ALF Webb School of California, Claremont In 1956 John C. Donohoe described from the Pipestone Springs formation of Chadronian age in the Montana Oligocene two jaw fragments of a new rodent, Pipestoneomys bisulcatus, that he placed in the family Aplodontidae. The purpose of this paper is to record the occurrence of Pipo- stoneomys in the Chadron formation of Nebraska, to suggest that it be placed tentatively with the castorids rather than with the aplodontids, and to propose a new species of the genus. North of Crawford, Nebraska, well known exposures of Oligo- cene sediments occur along the Pine Ridge escarpment. For several years, through the kindness of Frank Arner of Crawford, Nebraska, the Webb School of California has had the privilege of collecting fossils on the Arner ranch. In See. 26, T. 33N., R. 53W., Sioux County, Nebraska, are several harvester ant mounds in the Chadron formation, and from these have been collected a rich microfauna including Pipestoneomys. I wish to thank W. D. Turnbull of the Chicago Natural History Museum for the loan of the type specimen of P. bisulcatus for study, and Bryan Patterson of Harvard University, and Dr. Mary Dawson of the National Science Foundation for advice. The new species is named in honor of Professor Bryan Patterson. The drawings are by Nick Strekalovsky. The abbreviations refer to the following institutions or collections: C.N.H.M., Chicago Na- tural History Museum; M.C.Z., Museum of Comparative Zoology ; R.A.M., Raymond Alf Museum, Webb School, Claremont, Cali- fornia. 2 BREVIORA No. 172 Family CASTORIDAE? PIPESTONEOMYS PATTERSONI Nn. sp. Type: M.C.Z. no. 7113, fragment of left maxilla with M12. Hypodigm: The type and M.C.Z. nos. 7110, RP*; 7106, LP#; 71108, dbP4 57102, LUM oft RM: 7104 uP, -71OTe Re aean LOS, RM, ; 7109, RM, ; 7112, RM, ; 7101, LM, ; and R.A.M. nos, 3072, RM?; 1683, LM?; 671, LM?; 1283, RP, ; 2105, RP, ; 2104, RM,. Horizon: The base of the Chadron formation along Pine Ridge is marked by a basal conglomerate resting on the weathered sur- face of the Pierre shale. The top is characterized by the upper purple-white layer, a continuous purple-tinted white limestone designated by Schultz and Stout (1938) as the boundary be- tween the Chadron and the Brulé formations of the White River group in this area. About half way between this and the basal conglomerate there is a second limestone lens, which is referred to as the lower purple-white layer. The specimens come from well below the lower purple-white layer. Localtiy: See. 26, T. 33N., R. 53W., Sioux County, Nebraska. Diagnosis: Differing from P.. bisulcatus as follows: paracone and metacone of M1-? not rounded, paracone and anteroloph not sharply separated, antercexternal and anterointernal lakes sepa- rate, relatively small; M, smaller, more rectangular in outline. Description: The paracone and metacone of M1", though dis- tinct and higher than the remainder of the crown, are not the rounded cusps that they are in P. bisulcatus. In P. bisulcatus the paracone is separated from the anteroloph by a deep groove com- munieating with the anteroexternal lake, whereas in P. pattersont the separation is very faint. In P. bisulcatus the anteroexternal and anterointernal lakes communicate by a slight groove. In P. pattersoni the corresponding lakes are completely separate and are also relatively smaller and are more delicately outlined. There is no trace of a mesostyle. P+ is larger than either M! or M?. A deep groove separates the anteroloph from the rest of the tooth and communicates openly with a deep valley that separates the paracone-metacone loph from the hypocone. On the anteroloph there is a lake below the anterocone and an embayment projecting inward to the protocone. As on the upper molars, a mesostyle is lacking. The lower first molar of P. pattersoni differs in shape from the corresponding tooth in P. bisulcatus, being much more rectangu- lar. It is also smaller and more delicate. Py is larger than My. 1962 NEW OLIGOCENE RODENT 3 Fig. 1. Pipestoneomys pattersoni. A, M.C.Z. no, 7113, left maxillary frag- ment bearing M12, type. B, M.C.Z. no. 7110, RP*. C, M.C.Z, no, 7108, RM). D, M.C.Z. no. 7107, RP4. X 25. 4 BREVIORA Nost72 The anterolophid, metalophid, and posterolophid are well devel- oped and separated by deep grooves that terminate at the meta- conid and entoconid. There is a lake posterior to the metaconid and one on the posterolophid adjacent to the hypocones. Discussion: Pipestoneomys, especially in the light of the new evidence reported here, does not fit well into the Aplodontidae. The upper Eocene Eohaplomys, the earlest known aplodontid, Q: has the dental formula es (Stock, 1985; McGrew, 1941). This formula is retained to the present. The presence or absence of P% is indeterminable in the holotype of P. bisulcatus, but a Fig. 2. Pipestoneomys bisuleatus. Right M13, crown view, C.N.H.M. no. UM 409, type. small fragment of the maxilla anterointernal to a P# of P. patter- sont (M.C.Z. no. 7106) shows no sign of the presence of P?. P4 are large relative to the molars. The anterior margin of the mas- seteriec fossa in P. bisulcatus extends to beneath the middle part of Py, and the ramus is relatively thick and deep. These charac- ters are not typical of the early aplodontids, but are quite lke those occurring in castorids. The cheek tooth structure seems to me to accord better with that of castorids than with that of aplo- dontids. A lower molar of P. pattersoni (M.C.Z. no. 7112) was ground down to show the pattern of a well worn tooth. The median external groove (hypoflexid) persists but its medial portion be- comes transformed into a lake, forming a pattern similar in gen- eral to that found in, e.g., Paleocastor (cf. Fig. 3 and Stirton 1935, Fig. 30). The incisor enamel of Recent Castor, Miocene Paleo- castor, and Oligocene Agnotocastor is characterized by an orange 1962 NEW OLIGOCENE RODENT 5 color (Wilson 1949), that of aplodontids is not pigmented. Among the many incisors picked out of the ant mounds are a few with orange color, and those of appropriate size could well belong to P. pattersoni. Fig. 3. Pipestoneomys pattersoni. M.C.Z. no. 7112, RM, ground down to show pattern near base of crown. X 25. The past and present distribution of aplodontids is also of some interest in this connection. Not only is the range of Recent Aplo- dontia limited to a strip along the west coast of North America, but no unquestioned fossil form has been found east of the Great Basin.! If this picture of geographic distribution is adequate, we may have another item of evidence that argues against aplodontid affinities for Pipestoneomys. Pipestoneomys seems to me to fit more reasonably into the Cas- toridae than into the Aplodontidae, and I know of no other family of appropriate range to which it might be referred. If this tentative assignment is correct, the genus is the earliest beaver so far known, although its position in castorid phylogeny is un- certain. REFERENCES DONOHOE, J.C. 1956. New aplodontid rodent from Montana Oligocene. Jour. Mam- mal., 37: 264-268. McGrew, P. O. 1941. The Aplodontoidea. Geol. Ser., Field Mus. Nat. Hist., 9: 1-30. 1Shotwell, in his interesting study of the aplodontids (1958), mentions Pipes- toneomys only in passing, noting that P. bisuleatus occurred well to the east of the other known fossils. 6 BREVIORA No. 172 ScHuLTZ, C. B. and T. M. Stout 1938. SHOTWELL, 1958. StirtTon, R. 1935. Stock, C. 1935. WILSON, R. 1949. Le Classification of Oligocene sediments in Nebraska. Bull, Univ. Nebr. State Mus., 4: 15-52. J. A. Evolution and biogeography of the aplodontid and mylagaulid rodents. Evolution, 12: 451-484. A. A review of the Tertiary beavers. Univ. Calif. Publ. Bull. Dept. Geol. Sei., 23: 391-458. New genus of rodent from the Sespe Eocene. Bull. Geol. Soc. Amer., 46: 61-68. W. Early Tertiary rodents of North America. Carnegie Inst. Wash- ington, Publ. 584: 71-164. TABLE 1 Pipestoneomys. Upper Dentition Measurements in millimeters pattersoni j P. bisulcatus C.N.H.M. UM 409, type Crown length M1 M.C.Z. 7102 1.62 1.78 M.C.Z. 7113 1.47 M? M.C.Z. 7111 1.48 1.6 M.C.Z. 7113 1.23 R.A.M. 671 1.48 R.A.M. 1683 1.62 R.A.M. 3072 1.44 Pt M.C.Z. 7106 2.16 M.C.Z. 7108 2.07 M.C.Z. 7110 1.98 Greatest width M1 M.C.Z. 7102 offal 1.95 M.C.Z. 7113 1.67 M? R.A.M. 1683 1.62 1.81 R.A.M. 671 1.44 IMEC Ze lait 1.35 IMEC Zales 1.67 R.A.M. 3072 1.35 Pt M.C.Z. 7106 1.98 M.C.Z. 7108 2.09 M.C.Z. 7110 1.94 1962 P. pattersoni Crown length My M.C.Z. P4 M.C.Z. M.C.Z. M.C.Z. R.A.M. M.C.Z. M.C.Z. R.A.M. R.A.M. Greatest width My M.C.Z. P4 M.C.Z. M.C.Z. M.C.Z. R.A.M. M.C.Z. M.C.Z. R.A.M. R.A.M. 7101 7103 7109 7112 2104 7104 7107 1283 2105 7101 7109 7112 7103 2104 7104 7107 1283 2105 NEW OLIGOCENE RODENT TABLE 2 Pipestoneomys. Lower Dentition Measurements in millimeters 1.44 1.48 74: 1.89 1.62 1.62 1.51 1.80 iegal 1.51 1.62 1.62 1.35 1.53 1.35 1.42 1.48 1.62 P. bisulcatus C.N.H.M. UM 408 1.83 dP, 1.78 dP, 1.18 “J ; 7 7 7 . ‘dM darlin wae LY c mat be os ¢ a BREVIORA Museum of Comparative Zoology CAMBRIDGE, Ass. DECEMBER 24, 1962 NUMBER 173 NEW SPECIES OF LAND MOLLUSKS FROM THE REPUBLICA DOMINICANA By WILLIAM J. CLENCH We are deeply indebted to Drs. Stanley Rand, Clayton Ray and Juan Rivero for two important collections of land mollusks collected in the Reptiblica Dominicana. Both collections were made in areas hitherto unknown as far as the land mollusks were concerned, Of the four species herein described, two represent a subfamily and a genus not previously known from Hispaniola, Probably one-third of the species are known only from their type localities, the remaining two-thirds from a very few locali- ties. For this reason it is impossible to give significant distribu- tional patterns of any one species, and but few generic patterns. Our present knowledge of the land and freshwater mollusks of Hispaniola is probably only about ten percent, compared with that of Cuba. Much of this is due, of course, to the fact that there are no local interested persons in Hispaniola. Cuba has had a host of highly trained naturalists, such as Juan Gundlach, Charles Wright, Filip Poey, Rafael Arango, Carlos de la Torre, P. J. Bermudez and C. G. Aguayo. All have made extensive collecting trips throughout most of Cuba and their interest stimulated many others to collect. As a consequence, there are comparatively few areas in Cuba where the fauna still remains unknown. Conversely. Hispaniola has had none of the local stimulation, and what is known of the area has been gathered by outsiders on a few expeditions such as those made by Auguste Sallé in 1847-1851, D. F. Weinland in 1857, Justus Hjalmarson in 1858, Paul Bartsch in 1917, 1920, and 1929, W. J. Kyerdam Mm 1927... K. Oreutt im 1929-1930"): CC. Pease m- 1932 and W. J. Clench, R. A. McLean and H. D. Russell in 1937. Many others have contributed material, much of it secondary to other zoological or botanical pursuits (see Crosse 1891, and Bartsch 2 BREVIORA No. 173 1946). There remains much territory on this island about which nothing is known, particularly the central mountain system, the Cordillera Central. HELICINIDAE HELICINA JULIAE, new species Plated, ficuner2 Description. Shell reaching 8 mm. in greater diameter, im- perforate, subglobose, smooth and shining. Color a bright pea- green to yellow with the first 1144 post-nuclear whorls with a broad band of brownish purple, as well as a peripheral band of the same color. Columellar area white and occasionally mar- gined with blue. There may be one or more axial bands which occur only from the peripheral band to the suture above. Whorls 4 and convex. Spire depressed and obtuse. Aperture subtri- angular in shape. Outer lip reflected, white, and with a mod- erate depression behind. Parietal area thinly glazed. Columella very short. Seulpture consisting of very fine growth lines. [leieht Width 5.9 min. 8.0 mm. Holotype 5.1 mm. 7.0 mm. Paratype Types. The holotype is in the Museum of Comparative Zool- ogy, no. 168267. from Colonia Ramfis. 20 km. W of San Cristobal. Republica Dominicana. Dr. Juan Rivero collector, March 19°7. Paratypes from the same lceality in the Museum of Comparative Zoology and the Academy of Natural Sciences of Philadelphia. Remarks. This species differs from Helicina viridis Lamarck to which it is probably related, by being much smaller and by having several color phases in addition to the axial brown line or lines which extend from the peripheral band to the suture above. H. viridis reaches 14 mm. in width and 9.5 mm. in height and is a pure pea-green in color. The two species are the same in shell outline. PROSERPINA MARCANOI, new species Plate 1, figure 3 Description. Shell reaching 4.7 mm. in width, depressed, im- perforate, smooth and shining. Color a hght whitish green. Whorls 5 and convex. Spire depressed. Aperture semicircular. 1962 LAND MOLLUSKS FROM HISPANIOLA 3 Outer lip simple. Parietal wall glazed and supporting a high and thin lamella. Columella short and supporting a second lamella which is about one-third as high as the parietal lamella. Umbilical area slightly depressed. Suture defined but not im- pressed. Sculpture consisting of exceedingly fine growth lines. Height Width 2.4 mm. 4.7 mm. Holotype 2.2 mm. 4.4 mm. Paratype 2.1 mm. 4.5 mm. ie Types. The holotype is in the Museum of Comparative Zool- ogy, no. 188911, from Colonia Ramfis, 20 km. W of San Cristobal, Reptabhca Domimicana. ‘wo paracypes are in the Museum of Comparative Zoology from the same locality. Dr. Juan Rivero collector, March 1957. Remarks. This is the first species of the Proserpininae to be recorded from Hispaniola. The subfamily has been known elsewhere only from Jamaica and Cuba. In relationship, it appears to be nearest to P. depressa d’Orbigny, a species which is widely distributed in Cuba. It differs from depressa by being smaller in size, proportionally higher, and in having a columellar lamella nearly twice as high as the Cuban species. This species is named for Eugenio Mareano of the Instituto de Investigaciones Botanicas y Zoologicas de la Universidad de Santo Domingo, who collected with Dr. Juan Rivero when he was at Colonia Ramfis in Mareh 1957, TRUNCATELLIDAE GEOMELANIA (MERRILLIANA) RIVEROI, New species Plate 1, figure 4 Description. Shell reaching 7.6 mm. in height (truncated), extended, limperforate and strongly sculptured. Color a dull white. Whorls 8 and moderately convex. Aperture holostoma- tous and attached to the body whorl. Parietal and palatal lips reflected. Columella short and shghtly oblique. Suture im- pressed. Sculpture consisting of numerous axial costae with exceedingly fine spiral threads between the costae. Operculum unknown. Height Width 7.6 mm. 2.0 mm. Holotype 4 BREVIORA No. 173 Types. The holotype is in the Museum of Comparative Zool- ogy, no. 230505, from Colonia Ramfis, 20 km. W of San Cristo- bal, Reptiblica Dominicana, collected by Dr. Juan Rivero in March 1957. Remarks. This is the second known species of Geomelania from Hispaniola. It differs from Geomelania haitiensis Wein- land in being twice as large and in having the axial sculpture coarser and by havine 8 whorls, G. haitiensis having only 5. In relationship, it appears nearest to Geomelania elongata Pfeiffer from Oriente, Cuba, and differs from it by being a little larger, having one more whorl and in being a little more coarsely sculptured. SAGDIDAE ZAPHYSEMA RANDI, hew species Plate 1, figure 1] Description. Shell large, reaching 67 mm. in height, smooth, rather thin and globose. Color probably a yellowish brown. Whorls 414 and strongly convex. Outer lp simple, parietal wall with a thin glaze. Suture slightly imdented. Columella long, sinuous and somewhat oblique. Sculpture consisting of very fine axial growth lines. Height Width 65.0 mim. 67.0 mm. Holotype 67.0 min. 67.5 mim. Paratype Types. The holotype is in the Museum of Comparative Zool- ogy, no. 2305038, from Cueva de San Francisco, Cerros de San Francisco, Mun, Pedro Santana, San Rafael, Republica Domini- cana. Collected by Stanley Rand and Clayton Ray, August 1958. There is single paratype from the same locality. Remarks. This species is placed provisionally in the genus Zaphysema, a genus heretofore known only from Jamaica and Navassa islands. It is about three times as large as the largest Jamaican species, Zaphysema macmurrayr C. B. Adams. Both specimens were dead when found in the cave and appear to be quite old as they are white and completely devoid of perios- tracum. This species is as large as the largest Polydontes gigantea Scapoli from Hispaniola and Zachrysia petitiana d’Orbigny from the Trinidad Mountains of Cuba. Its large size and thinner shell makes it more capacious than either of these two species. 1962 LAND MOLLUSKS FROM HISPANIOLA 5 This is by far the largest species in the family Sagdidae, a family which occurs in the West Indies, southern United States,! and south through Central America and northern South Amer- ica. Its greatest generic development is centered in the island of Jamaica. REFERENCES Barrscu, Paun 1946. The operculate land mollusks of the family Annulariidae of the Island of Hispaniola and the Bahama Archipelago. Bull. U.S. Nat. Mus. 192: 1-3. CrossE, H. 1891. Faune malacologique terrestre et fluviatile de 1’[le de Saint Dominge. Jour. Coneiyl. 39: 69-95. TURNER, R. D. 1960. Land shells of Navassa Island. Bull. Mus. Comp. Zool., 122: 233-244. 1A single genus, Microphysula, with two species, ingersolli Bland and cookei Pilsbry extends north through the Rocky Mountain states to southern British Columbia. PLATE Fig. 1. Zaphysema randi Clench. Cueva de San Francisco, Mun. Pedro Santana, Reptblica Dominicana (about natural size). Fig. 2. Helicina juliae Clench. Colonia Ramfis, 20 km. W of San Cristobal, Reptiblica Dominicana (7.4X ) Bigs 3: Proserpina marcanoi Clench. Colonia Ramfis, 20 km. W of San Cristobal, Reptblica Dominicana (13.8X). Fig. 4. Geomelania (Merrilliana) riveroi Clench. 20 km. W of San Cristobal, Republea Colonia Ramfis, Dominicana (about 9X). BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 24, 1962 NUMBER 174 A NEW ARCTOCYONID FROM THE PALEOCENE OF WYOMING By Bryan Patterson and Paul O. McGrew ! Since 1959, the Department of Geology of the University of Wyoming and the Museum of Comparative Zoology, Harvard University, have been carrying on a joint program of field and laboratory work in the later Mesozoic and earler Cenozoic verte- brate-bearine formations of Wyoming, paying special attention to those deposits amenable to washing techniques. Much of the work accomplished thus far has been done in the Shotgun member (Keefer, 1961) of the Fort Union formation in the northern part of the Wind River Basin. Within this member is a bone bed, some twelve to twenty-four inches in thickness, that is extraordinarily rich in vertebrate re- mains, mostly single teeth of mammals, crocodiles and small sharks. The bed occurs at the base of a sandstone (umt 20 of Keefer’s section at Twin Buttes) 217 feet above the base of the Shotgun member, which is 1265 feet in thickness at this locality. Keefer has shown that overlying the lower Fort Union in this region are two essentially contemporaneous, interfingering rock units, the lacustrine (or marine?) Waltman shale and the mar- einal Shotgun member. The mammalian fauna from the bone bed in the Shotgun is a rich one. From Keefer’s sample collection, Gazin (in Keefer, 1961) has recorded: Ptilodus, Mimetodon, Ec- typodus, Anconodon, Eucosmoedon?, Catopsalis, Pecradectes?, Gelastops, Diacodon?, possibly Aphronorus, Pentacodon, possibly Zanycteris, cf. Pronothodectes, Clacnodon, Tricentes, Peripty- chus, Anisonchus, Promioclacnus, Litomylus?, Gidleyina and Pantolambda. Our extensive collections will add a number of forms as the work of identification proceeds. The Shotgun mem- ber at Twin Buttes has vielded very little in the way of vertebrate material outside of the bone bed.2 Thus far we have found only a 1 Department of Geology, University of Wyoming. 2 During the 1961 season a concentration similar to but less rich than that of the bone bed was found some 90 feet lower in the member. 2 BREVIORA No. 174 few fragments of a medium-sized pantodont. We propose that the assemblage be known as the Shotgun local fauna, from the name of the member in which it occurs. The age, so far as can be deter- mined at present, appears to be early Tiffanian. At Shoteun Butte, the type locality, where the member has a thickness of 2.850 feet (Keefer, 1961), Phenacodus sp. and Plesiadapis sp. cf. P. cookei, have been found in the upper part of the sequence (Keefer and Troyer, 1956). We are greatly obliged to Dr. William R. Keefer for his help in the field. For the opportunity to examine comparative material we are indebted to Dr. C. Lewis Gazin, U. S. National Museum, Dr. Malcolm C. MeKenna, The American Museum of Natural His- tory, and Dr. Craig C. Black, Carnegie Museum. During the field season of 1960 we were assisted by James A. Jensen, Lee A. Wooderson, John Zamecnik, Floyd Andrews, Clyde T. Williams. Richard P. Timmermeyer, Charles P. Lyman, Jr., and Robert IF. Wallin. The photographs are the work of Miss Linda Loring. The following abbreviations are used: A.M.N.IH1., American Museum of Natural History; M.C.Z., Museum of Comparative Zoology ; U.W., University of Wyoming. ARCTOCYONIDAE ! Murray OXYCLAENINAE Matthew COLPOCLAEN US “gen. nov. Type species: C. keefert sp. nov. Known distribution: Paleocene, early Tiffanian, Wyoming. Diagnosis: Enamel of molars stronely wrinkled, all crests cren- ulated, accessory cuspules numerous. Principal cusps of upper M Ingh, massive, tightly grouped; central basin small; protocone nearly central in position with lone lingual slope; conules large, blunt, separated by grooves from principal cusps; hypocone small on Ml?, rudimentary on M*; cingula strong, not continuous around protocones; external cingulum interrupted by labial con- tinuation of sharp cleft between paracone and metacone ; upper M wide relative to length. Trigonids of lower M high, short ; para- conid internal in position, closely appressed to metaconid, para- conid crest well developed: posterior crest connecting protoconid 1QOne of us (B.P.) proposes to transfer the Aretocyonidae from the Creodonta to the Condylarthra. ‘This transfer and the various questions that it raises will be discussed in a later paper. 2 Kolpos, a fold and claenus ; in allusion to the wrinkled enamel of the molars and. in particular. to the curiously folded trigonids. ~~ 1962 NEW PALEOCENE ARCTOCYONID and metaconid present; protoconids and metaconids with short, centrally situated crests running lingually and labially, respee- tively, forming a third, transverse trigonid crest ; deepest portion of talonid basin near lingual side; hypoconulid of Ms lone, broad, high, cuspidate. COLPOCLAENUS KEEFERI! sp. nov. Type: M.C.Z. no. 8355, LMs. Hypodigm: Type and U.W. nos. 1931, LM. ; 1932, RMs ; 1933, LMt; 1934, RM; 1935, LM.; M.C.Z. nos. 8356, RM; (much worn) ; 8357, LM?; 8358, RM». Horizon and locality: Shotgun local fauna, Shotgun member of the Fort Union formation; NE, SE1,, see. 31, T. 6 N., R. 3 E., 34 mile SW of the more northerly of the Twin Buttes, Fremont County, Wyoming. Diagnosis: Sole known species of the genus diagnosed above. DESCRIPTION Colpoclacnus keeferi is notable for the massive, high, principal cusps of the upper molars, the elevated trigonids and the extreme- ly rugose enamel. The latter feature disappears after very little Wear, as is demonstrated by a moderately worn Ms (M.C.Z. no. 8398). There can be, we believe, no doubt that the various lower molars represent the same species, and their association with the uppers would appear to be demonstrated by excellent occlusion between M.C.Z. no. 8357, LM?, and U.W. no. 1935, LMs, as well as by the general structure. The upper molars are considerably wider than long, \I!> being essentially quadrangular, M® suboval in outline. The protocone is the largest of the three principal cusps; its apex is a little lin- cual to the center of each tooth of the series, and its hnegual face is very long and gently sloping, rather shorter in M*? than in Mt. The paracone is more labial in position than the metacone and is slightly larger and higher, being approximately the same height as the protocone. Paracone and metacone are grooved on their basinward faces and are separated by a well marked cleft. The large blunt proto- and metaconules are separated from the primary cusps by grooves and are united by crests to the anterior and posterior cingulum respectively; the protoconule is smaller 1For Dr. W. R. Keefer who discovered the Shotgun bone bed in the course of his work on the Fort Union of the Wind River Basin. 4 BREVIORA No. 174 Figure 1. Colpoclaenus keeferi gen. et. sp. nov. Upper molars: a, LM, M.C.Z. no. 8357: b, LM!, U.W. no. 1933: c, RM?, U.W. no. 1934. X83. Stereoscopic views. 1962 NEW PALEOCENE ARCTOCYONID 5 than the metaconule on M12, about equal to it in size on M®. The rather shallow basin enclosed by these cusps and conules is situ- ated in the labial half of the tooth and bears various poorly de- fined cuspules on its floor. A hypocone is differentiated on M1l?, on which it is considerably smaller than the conules, but is barely distinguishable on M*. Apart from an interruption on ihe exter- nal face, where the cleft between paracone and metacone eon- tinues on to the labial margin, a strong, crenulated cingulum runs from the hypocone around the tooth to the antero-internal corner, at which point the cingulum is broken in M!. M? shows no proto- stylar enlargement here and M® a very slight one. With the ex- ception of a sheht rise in the cingulum at the parastylar site on M®, there is no indication of external styles. Anterior and pos- terior cingula extend farther lingually on M? than on the preced- ing teeth and the two are almost united by a median lingual cuspule. Several cuspules are present on the posterior face of M®? above the cingulum. Ms is shehtly constricted at the Junction of trigonid and talo- nid but is otherwise nearly quadrangular in outline; the trigonid and talonid are approximately equal in Jeneth and width. The trigonid is somewhat higher than the talonid and narrows apically. The protoconid is a large massive cusp that forms nearly half of the trigonid. The heavy, cuspidate paraconid crest runs from the apex of the protoconid to the lingual side of the tooth, where the paraconid is poorly differentiated. The blunt metaconid is but little smaller than the protoconid and equal to it in height; the apices of the two cusps are connected by a papillate crest that is more lightly built than the paraconid crest. Blunt, wrinkled crests run directly lingually and labially down the facing slopes of the protoconid and metaconid. These abut (U.W. no. 1935) or fuse (M.C.Z. no. 8358) to form a third, central transverse crest. A sinuous, antero-posterior cleft partially separates the trigonid apex into labial and lingual halves. All this compheated structure is shallow and would rapidly be obliterated by wear. The pos- terior face of the trigonid is sloping and bears various small, papillate crests; that behind the metaconid is the most prominent but there is no trace on it of a metastylid. The very large, blunt hypoconid nearly equals the protoconid in size and makes up nearly half of the talonid. The cuspidate crista obliqua is low and fades away toward the base of the talonid. Two minor crests run forward from it to the trigonid base, isolating a very small fossette. The hypoconulid is low but relatively large, and in the 6 BREVIORA No. 174 ) Figure 2. Colpoclaenus keeferi gen. et sp. noy. Mo: a, U.W. no. 1935; b, M.C.Z. no. 8858. X3. Stereoscopic views. unworn U.W. no. 1935 is tricuspidate and set off from the adja- cent cusps by shallow grooves. As is shown by M.C.Z. no. 8358, it is obliterated as a distinct cusp after only a little wear. The entoconid is higher but searcely larger than the hypoconulid and, like it, at least in U.W. no. 1935, is trieuspidate. From it a cuspi- date ridge runs down to the base of the trigonid. The talonid basin is deepest in its anterolingual portion, and the floor bears a number of poorly defined cuspules. A strong, papillate cimeu- lum runs from a point beneath the paraconid around the labial side of the tooth to the hypoconulid; in U.W. no. 1935, but not in 1962 NEW PALEOCENE ARCTOGYONID if SUM ios Je Rie c, U.W. no. 1932. X3. Stereoscopic Views. Figure 3. Colpoclaenus keeferi gen. et sp. nov. Ms: a b, M.C.Z. no. 8355, type; ’ 8 BREVIORA No. 174 M.C.Z. no, 8358, this rises to a cuspule in the sight notch between trigonid and talonid. The third molar is shghtly narrower and rather longer than the second; it tapers bluntly toward the rear and, as in Mos, trigonid and talonid widths are approximately equal. The trigonid is shorter and conspicuously higher than the talonid, notably more so than in Mos, tapers apically as on that tooth and is slightly oblique to the long axis, being longer on the lingual than on the labial side. The crown of the trigonid resembles that of Ms in general, differing in the lesser development of the central trans- verse crest, which results in the formation of a shallow basin. The posterior face is nearly vertical, with wrinkled enamel. The hypo- conid is the largest cusp of the tooth; it is low, blunt, elongate and bears a crest that is bowed laterally. The crista obliqua is lacking in U.W. no. 1931 and is represented only by vestigial, incomplete crestlets in the type and U.W. no. 1932; the hypoconid crest con- tinues on to the base of the protoconid. The broad, flat-crowned hypoconulid is the highest of the talonid cusps and is almost as large as the hypoconid. It is set off by shallow grooves and is itself varyingly grooved and cuspidate. The entoconid resembles that of Ms both in size and in structure. As on My, the talonid basin is deepest antero-lingually and its floor is vaguely cuspidate ; it is open anterior to the entoconid due to the absence of a crest on the posterior slope of the metaconid, which on the preceding tooth, together with the entoconid crest, forms a slight dam at this point. The external cingulum runs from the anterior face of the trigonid to the hypoconuhd, showing some tendency toward formation of a cuspule between hypoconid and hypoconulid. The tooth is very slightly constricted at the junctions of the trigonid and talonid and of the hypoconulid with the adjacent cusps. DISCUSSION We have been able to compare the material referred to Col- poclaenus keeferi with specimens representing most of the de- scribed aretoeyonids. Colpoclaenus is a very distinct genus; the combination of characters it presents — summed up in the diag- nosis — separates it sharply from all other forms, although cer- tain of them do approach it in one character or another. Anacodon resembles it in havine accessory cuspules and strongly wrinkled enamel, and also in the possession of three transverse crests on the trigonid, as shown by an unworn Ms of A. ursidens (A.M.N.H. 1962 NEW PALEOCENE ARCTOCYONID 9g no. 92). Claenodon has moderately wrinkled enamel and com- parable crests on the molar trigonids (Gazin, 1956, pl. 7, fig. 5). Other forms — Thryptacodon, Tricentes, Mimotricentes, Para- doxodon — have the enamel wrinkled to varying degrees but do not otherwise resemble Colpoclaenus. In Protogonodon the proto- cone has a fairly long lingual slope. The trigonid is high in Oxy- claenus, Chriacus, Spanoxryodon, Prothryptacodon, and Metachri- acus. There is a tendency in some of these, especially in Prothryp- tacodon, for the trigonid height to decrease from M, to Ms; in Colpoclaenus the trigonid is higher on Ms than on Ms. Colpo- claenus thus presents a curious combination of primitive charae- ters, such as high principal cusps in the upper molars, high trigo- nids in the lowers, and advanced ones, such as a degree of wrink- ling of the enamel and proliferation of accessory cuspules that is exceeded only in Anacodon. As the family is currently sub- divided, Colpoclaenus must be placed in the Oxyclaeninae, but its acquisition of arctocyonine-like wrinkling perhaps gives added point to Simpson’s suggestion (1937, p. 172) that the Arctocyo- ninae may be a partially artificial assemblage composed of several lines that were independently following similar adaptive trends. The last lower molars of Colpoclaenus are quite primate-like in general appearance and in some structural details. When our knowledge of the animal was confined to these teeth and to the shghtly worn Ms we were persuaded that we were dealing with a primate astonishingly large for the Paleocene!. Recognition of the upper molars rendered such an identification less likely and the finding of the unworn Ms seems to us to have ended all pos- sibility of it. We wryly sympathize with Cope and with Osborn and Earle who also at times believed an oxyelaenine (Chriacus ) to be a primate. Measurements in mm. U.W.-no. 1933 M.C.Z. no. 8357 U.W. no. 1934 M! M? M2 L fell 8.0 6.4 W alse = 8.8 U.W. no. M.C.Z. Type, M.C.Z. U.W. no. U.W. no. M.C.Z. 1935 no. 8358 no. 8355 1931 1932 no. 8356 Mo Mo Ms Ms Ms Ms L 8.8 9.1 112 10.5 10.4 10.9 W 7.0 7.6 6.4 6.3 6.2 6.3 1 Unfortunately this enthusiasm found fleeting expression on page 9 of the Society of Vertebrate Paleontology News Bulletin No. 61, February, 1961. 10 BREVIORA No. 174 REFERENCES GAZIN, C. L. 1956. Paleocene mammalhan faunas of the Bison Basin in south-central Wyoming. Smithson. Mise. Coll., 131, No. 6: i-iv, 1-57. KEEFER, W. R. 1961. The Waltman shale and Shotgun members of the Fort Union formation (Paleocene) in the Wind River Basin, Wyoming. Bull. Amer. Assoc. Pet. Geol., 45: 1310-1323. KEEFER, W. R. and M. L. TROYER 1956. Stratigraphy of the Upper Cretaceous and Lower Tertiary rocks of the Shotgun Butte area, Fremont County, Wyoming. U.S. Geol. Sury., Chart OC 56, Oil and Gas Inv. Ser. SIMPSON, G. G. 1987. The Fort Union of the Crazy Mountain field, Montana, and its mammalian faunas. Bull. U.S. Nat. Mus., 169: i-x, 1-287. eS BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 24, 1962 NUMBER 175 A PICRODONTID INSECTIVORE(?) FROM THE PALEOCENE OF WYOMING By Pauu O. McGrew ! and BRYAN PATTERSON Among the interesting materials found in the course of recent field work in early Tertiary deposits in Wyoming, conducted jointly by the University of Wyoming and the Museum of Com- parative Zoology, are several fragments of maxillae and man- dibles and a number of isolated teeth of a picrodontid. Only two other records of these problematical little mammals are known: Picrodus silberling’ Douglass 1908 from the Fort Union (Lebo) of Montana and Zanycteris paleocena Matthew 1917 from the Tiffany of Colorado, the former based on lower, the latter on upper teeth. We confidently assume that our upper and lower molars are referable to the same species. On the evidence of the upper molars this species cannot be placed in Zanycteris and on that of the lowers it cannot be excluded from Picrodus. We suspect it to be new, but the differences from, and the parts in common with, P. silberlingt are so few that we refrain for the present from any attempt at diagnosis. Gazin’s tentative record of Zanycteris from the Shotgun member of the Fort Union for- mation in the Wind River Basin (1961, p. 51, and in Keefer, 1961) is almost surely based on this form. The acknowledgments made in a previous paper (Patterson and McGrew, 1962) apply equally to this one, except that the photographs printed here were taken by Mr. John F. Cutler. Abbreviations are as follows: U.W., University of Wyoming; M.C.Z., Museum of Comparative Zoology. 1 Department of Geology, University of Wyoming. y BREVIORA No. 175 INSECTIVORA(?) PICRODONTIDAE Simpson PICRODUS Douglass Picropus sp. ef. P. SILBERLINGI Douglass Material: U.W. no. 1780, fragment of left maxilla with M?t?, incomplete alveolus of M?; M.C.Z. no. 8363, fragment of right maxilla with M!, incomplete alveolus of M*; U.W. no. 1781, four complete and three partial M,; M.C.Z. no. 8422, three M, ; M.C.Z. no. 8423, Mo. Horizon and localitics: Paleocene, early Tiffanian. U.W. no. 1780 is from the Bison Basin Saddle locality, Fort Union for- mation, E1/, sec. 28, T. 27 N., R. 95 W., Fremont Co., Wyoming (Bell, MS.; Gazin, 1956). All the other specimens are from the Shotgun loeal fauna, Shotgun member, Fort Union formation, *, mile SW of the more northerly of the Twin Buttes in the northern part of the Wind River Basin, NE 14, SE 1%, see. 31, T.6N., R. 3 E., Fremont Co., Wyoming (Keefer, 1961; Patter- son and MeGrew, 1963). All specimens were encountered in the course of washing operations. DESCRIPTION The upper molars of the Wyoming form agree with those of Zanycteris paleocena (Matthew, 1917; Simpson, 1935) in’ basic structure vet differ considerably in detail, especially as regards M?. This tooth is nearly quadrangular, rather than triangular, in outline, an outpocketing of the posterior cingulum forming a postero-internal angle. The protocone is more anterior im posi- tion and more directly internal to the paracone. The protoloph, within which the protocone is completely subordinated, is a heavy, transversely aligned crest that extends almost to the in- ternal base of the paracone, from which it is separated by a shallow notch. The anterior face of the tooth slopes gently up- ward; the anterior cingulum (double in U.W. no. 1780) is strong compared with that of Zanycteris. The antero-external corner of the tooth is rounded and bears no stvlar projection. A faint, irregular crest extends, with interruptions, from the base of the protocone to the metacone, It is situated almost wholly within the central basin, of which the lingual and postero- lingual margins are formed by the cingulum. The enamel within 1962 A PALEOCENE PICRODONTID ment of left maxilla with M! b Fig. 1. Picrodus sp. ef. P. silberlingi Douglass. a, U.W. no. 1780, frag- b, M.C.Z. no. 8363, fragment of right max illa with M!, Approximately X 8. Stereoscopie views. 4 BREVIORA No. 175 the area enclosed by the cingulum and the aforementioned low crest is fully as wrinkled as that over the main portion of the central basin. Picrodus sp. is rather more advanced than Z. paleocena in the essentially complete incorporation of this ¢in- eular area into the basin and in the presence of the small postero- internal outpocketing already mentioned.’ Paracone and meta- cone are mere elevations at the ends of the lone, nearly straight eetoloph. An external cingulum is present posterior to the para- cone, where it forms a stout, rather wide and featureless shelf that passes back into the postero-externally projecting meta- stylar area; a crest connects metastyle and metacone. M? is quite similar to the corresponding tooth of Z. paleocena, differing only in a few points. The tooth is shghtly smaller rela- tive to M!, the postero-internal angle is somewhat more squared, and the antero-external style is rather more prominent and con- nected by a low inconspicuous crest to the tip of the paracone. The adjacent styles of M12 abut to form an apex from which the outer marein of the cheek tooth series falls away anteriorly and posteriorly. The first lower molar differs only in detail from that of P. silberlingt. The paraconid is usually higher and shehtly more independent and is invariably larger than the metaconid. In several teeth the latter cusp has almost or quite lost its identity in the oblique trigonid crest. Trigonid and talonid are con- sistently better demarcated externally. The crest that forms the external border of the talonid curves inward at its anterior extremity to reach the center of the posterior face of the trigonid a short distance below the protoconid rather than abutting against the base of the trigonid externally, as is the rule in P. silberlingi. A narrow, shallow groove les between the trigonid face and the incurvinge position of the crest. The main portion of the talonid crest, that forming the lateral and posterior borders of the basin, appears to be more cuspidate than in the Lebo sample; the cuspules number from four to nine rather than from two to three. The two ecuspules on the lingual side of the talonid are extremely variable in the Shotgun sample, amount- ing in some specimens to little more than irregularities on the talonid margin. An interradicular crest is present. Ms. resembles the corresponding tooth of P. silberling: more closely than does M,; Simpson’s description (1937, p. 138) applies practically verbatim to our single specimen. 1 The posterior cingulum of M1 of Zanycteris paleocena is not as distinct from the dentral basin as Simpson’s figure (1935, fig. 6) would suggest. 1962 A PALEOCENE PICRODONTID D Fig. 2. Picrodus sp. ef. P. silberlingi Douglass. a, U.W. no. 1781, RM), approximately X 20. b, M.C.Z. no. 8422, RM,, approximately X 11. ¢ M.C.Z. no. 8423, RMo, approximately X 11. Stereoscopic views. ’ 6 BREVIORA No. 175 In both upper and lower molars, the enamel at the base of the erown extends out far beyond the necks of the teeth, the external and posterior cingula of M?! and the postero-external portion of the talonid of M, being especially projecting. The term exoedae- nodont has been apphed to this condition, which, in the Insee- tivora, is encountered especially among the Dimylidae ( Hiirzeler, 1944). Hurzeler and also Saban (1958, p. 896 n) believe that exoedaenodonty is an indication of a malacophagous diet, Saban citing in support the numerous shells of Anodonta that occur in deposits that yield remains of Dimylinae. However this may be for dimylids with their strongly cusped molars, we do not believe that it indicates any such habit for the picrodontids.! As Matthew recognized in his description of Zanycteris, low crowned more or less flattened molars with finely wrinkled enamel of the sort that occur in the family almost certainly in- dicate a frugivorous diet. The small fragments of maxillae and mandibles add nothing to knowledge except to suggest that the anterior root of the zyzoma may not have been as stout as in Zanycteris. MEASUREMENTS IN MM. U. W. 1780 M.C.Z. 8363 Length Mt 2.85 2.95 Width M! 2E3O 2.40 Length M= E35 = Width M? 1.63 = N OR M o Vi Oblique maximum diameter? M, 7 240 2385 2566) s=205il RS) SEONG 5.07 £1.35 Width talonid My LO 10 O14 OP aC S03 86s lo i= 0 ll ON Oe Length Mo 1.90 Width My 1.10 1 Nor, a fortiori, for such forms as soricids and various microchiropterans that are exoedaenodont to varying degrees. 2 As measured by Simpson (1987, p. 138). —l 1962 A PALEOCENE PICRODONTID DISCUSSION Matthew, in his description of Zanycteris paleocena, took no account of Picrodus silberlingi, and indeed there was nothing in Douglass’ description and figures to invite comparison between the two forms. It remained for Simpson, with new material of P. silberlingt and first-hand knowledge of Z. paleocena, to show that they were closely related. As he pointed out (1937, p. 136), the species were distinct, but in the absence of comparable parts it could not certainly be stated that this was also true of the genera. We now have lower molars indistinguishable generically from Picrodus found with upper molars clearly distinet from those of Zanycteris. On the face of it, then, the two previously described species would appear to belong to distinet genera. However probable, this cannot yet be considered as certain. The highly specialized lower molars may perhaps prove to be rather stereotyped within the family and hence unreliable for generic discrimination. Although quite unlikely, it is thus still coneeiv- able that P. silberlingi and Z. paleocena may represent one genus, and P. sp. another. Finds of associated upper and lower teeth of the previously described forms will be required before all doubt on this score can be set at rest. As regards the broader question of picrodontid affinites, we are in complete agreement with Simpson (1937) that the family cannot be referred to the Chiroptera. The enlarged lower incisor, small upper canine, small premolars, lone and slender muzzle, and origin of the anterior root of the zygoma opposite M12 are definitely non-chiropteran characters. Since Matthew’s work, it has always been stated that the molars resembled those of the specialized Phyllostomatidae, members of the subfamilies Stur- nirinae, Phyllonycterinae and Stenoderminae. Thanks to the excellent collection of bats in the Museum of Comparative Zool- ogy, we have been able to make comparisons with nearly all members of these three groups. We are quite unimpressed by the resemblances between their molars and those of the picro- dontids. Essentially these are limited to wrinkled enamel and pointed trigonids, and are more than offset by numerous dif- ferences. Sturnira and the Phyllonyeterinae have smooth enamel. In Sturmra, the upper molars have deep, antero-posteriorly run- ning central valleys and no traces of lophs; the lowers have the trigonid and talonid basins confluent, the metaconid widely sep- arated from the protoconid, and the trigonid of M, neither com- pressed nor elevated. In the phyllonycterines, the upper molars 8 BREVIORA No. 175 are triangular and lack lophs and a stylar area; the lowers have the trigonids searcely elevated above the talonids, and all cusps, save the protoconid of M,, incorporated in the rims of the talo- nid basins. The Stenoderminae have wrinkled, often strongly wrinkled, enamel, but the molar structure is very different from that of the picrodontids. The upper molars are short in com- parison with their widths, M? is large (except in Pygoderma), frequently larger than M!, and the paracones and metacones are high and trenchant (in Pygoderma the paracones only), standing well above the flattened lingual portions of the teeth. The trigonids are elevated and compressed to points, but the compression does not involve the whole trigonid as it does in the picrodontids. The metaconid, when not subordinated in the crest running to the apex of the protoconid, is a distinct element situated low on the crown. The pointed protoconids of both molars and premolars give every appearance of being involved in the same morphogenetic gradient. The few resemblances in molar structure between picrodontids and specialized phyllostomatids seem clearly to be of the sort brought about by convergence. The Microchiroptera can be elim- inated as possible relatives of the Pierodontidae, and we can see in the latter nothing suggestive of the Megachiroptera. On the positive side, we can offer no really useful suggestion as to the ordinal affinities of the picrodontids. Simpson has remarked that ‘‘reference to the Primates is merely a possibility, with no positive evidence to commend it.’? The possibility, of course, exists, and such picrodontid characters as the small size of the trigonid compared with that of the talonid and the tight group- ing of the trigonid cusps could be cited as resemblances, how- ever vague, to early members of that order. Resemblances of this sort scarcely constitute positive evidence, however. Refer- ence to the Insectivora is equally unsatisfactory. Our queried placement of them there is strictly faute de mieux, due largely to reluctance to use the Primates as a serap-basket order. REFERENCES BELL, W. G. MS. The geology of the southeastern flank of the Wind River Moun- [1955]. tains, Fremont County, Wyoming. Pp. 1-204. On deposit in the library of the Department of Geology, The University of Wyoming. 1962 A PALEOCENE PICRODONTID 9 GAZIN, ©: I. 1956. Paleocene mammalian faunas of the Bison Basin in south-central Wyoming. Smithson. Mise. Coll., 131, no. 6: i-iv, 1-57. 1961. Occurrences of Paleocene Mammalia in Tertiary basins of Wvy- oming. Wyo. Geol. Assoc. 16th Ann, Field Conf, 47-52. HURZELER, J. 1944. Beitrige zur Kenntnis der Dimylidae. Abh. Schweiz. Pal. Ges., 65: 1-44. Kererer, W. R. 1961. The Waltman shale and Shotgun members of the Fort Union formation (Paleocene) in the Wind River Basin, Wyoming. Bull. Amer. Assoc. Pet. Geol., 45: 1310-1323. MarrHew, W. D. 1917. A Paleocene bat. Bull. Amer. Mus. Nat. Hist., 37: 569-571. PATTERSON, B. and P. O. McGrew 1962. A new arctocyonid from the Paleocene of Wyoming. Breviora, Mus. Comp. Zool., No. 174: 1-10. SABAN, R. 1958. Insectivora in Traité de Paléontologie, edited by J. Piveteau, 6 vol. 2: 821-909. Paris: Masson et Cie. SIMPSON, G. G. 3 1935. The Tiffany fauna, upper Paleocene, I.—Multitubereulata, Marsupiaha, Insectivora, and ?Chiroptera. Amer. Mus. Novit., no. 795: 1-19. 1937. The Fort Union of the Crazy Mountain field, Montana, and its mammalian faunas. Bull. U. S. Nat. Mus., 169: i-x, 1-287. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 27, 1962 NUMBER 176 ON THE RACKS OF KINIXYS BELLIANA GRAY By R. F. LAuRENT Museum of Comparative Zoology When I noticed the peculiarities of AKinixys belliana from northeastern Congo, I suspected that these populations might represent a race already described by Ruppell under the name of schoensis, but I did not have Riippell’s description. Dr. Mer- tens kindly gave me the necessary information and it became evident that these populations were different from schoensis, and I therefore described the race mertensi (1956). Dr. Mertens (in litt.) declared that schoensis was in his opinion a valid race, but he apparently changed his mind, since the rehabilitation of scho- ensis has never appeared (Wermuth and Mertens, 1961). In preparation for a checklist of the turtles of Africa, I have now reinvestigated the validity of schoensis and also the range of mertenst toward the east and northeast. Acknowledgments. I wish to thank Miss A. G. C. Grandison (British Museum of Natural History) and Mr. A. Opdenbosch (Technician in the Musée Royal de l’Afrique Centrale, Tervuren) who very kindly gave me invaluable information on specimens in their charge. My thanks are due also to Prof. G. G. Simpson for advice on statistical matters and to Dr. E. E. Williams for read- ing the manuscript and helpful criticisms. This study has been supported by National Science Foundation grant NSF G 17144. KINIXYS BELLIANA Gray This paper limits itself to the examination, with the aid of the ratios already used in the description of mertensi, of two ques- tions: (1) Is schoensis Riippell recognizable as a northeastern race, and (2) does mertensi connect with the typical form by a smooth celine in which no objective limits could be formulated for the eastern distribution of the race? bo BREVIORA No. 176 Measurements, additional to those taken for the description of mertenst (1956) (i.e. on other specimens), were taken on the MCZ specimens; Miss A. G. C. Grandison, with her customary kindness, sent the pertinent measurements for specimens from northeastern Africa and Uganda. Mr. A. Opdenbosch furnished also data on a large series of K. belliana from the Congo. Admittedly, the method of relying on other persons to take measurements has serious drawbacks since it can introduce bias in the data. For amphibians the discrepancies are very great indeed, but as far as turtles are concerned it has been hoped that the measurements are generally so easily definable that these dis- erepancies would be almost negligible. However, the comparison of the ratios already published (Laurent 1956) with those eal- culated on Opdenbosch’s measurements disclosed a considerable bias in the breadth of the carapace (maximum breadth, and breadth on level of lateral ends of humero-pectoral sutures). Even disregarding these data, the variation of belliana is such that the validity of schoensis appears untenable on present evi- dence. [Of course, other characters may prove some day that the northeastern populations or even some others from east or southern Africa are subspecifically differentiated.] Thus, in re- gard to the four ratios used in my previous work (Laurent 1956) the situation is as follows. 1. From the data at hand in 1956, the general shape of the carapace seemed definitely different in mertensi and ‘‘schoensis”’ and belliana. The carapace breadth (at the level of the lateral ends of the humero-pectoral suture) in per cent of the maximum breadth (back part of the carapace) was 87.2 to 97.8 (m = 92 for nine specimens) for belliana, 78.7 to 90.3 (m = 85.67 for fifteen specimens) for mertensi and 83.8 for the type of schoensis. The overlap was slight enough to make unnecessary any test for sig- nificance of the difference. However, the data now at hand are definitely less conclusive. The difference seems to be still statis- tically significant between mertensi and the complex of popula- tions from eastern and southern Africa which we eall belliana but which surely cannot be considered as homogeneous. A|so, because bias is obvious in these rather difficult measurements and because other differences are more clearcut, it has not been found worthwhile to make any statistical calculations. The specimens from Sudan, Abyssinia and Somaliland do not appear to be significantly distinct from those from East Africa, though more 1962 KINIXYS BELLIANA GRAY 3 consistent measurements taken on more numerous specimens could some day disclose a valid difference. 2. In my 1956 work the ratio between the median gular suture plus the median humeral suture and the breadth of the plastron at the level of the lateral ends of the humero-pectoral sutures was seen to show a very clearcut difference between belliana and mertenst — the figures being 56.7 to 73.3 (m = 66.87) for belli- ana (N=9), 72.9 to 91 for mertenst (N=15). The new data are ana (N=9), 72.9 to 91 (m=84.92) for mertenst (N=15). The new data are still reliable as the measurements are not likely to suffer from individual bias. The ratios now obtained are as fol- lows (expressed in percentage) : a) belliana (N = 87) 49.5-82.8 m=66.45 v=—12.51 b)) mertenst. (N = 22) (57)2 75-94 m—8418 v=12-47 G)) eschoensis: s(N— 8) 551-1 m—6913 9 — 10:30 d) nogueyt (N = 8) 65.4.73-83.1 The variation coefficients are high because the samples are heterogeneous from the point of view of size and consequently of age. In the mertensi-belliana and belliana-‘‘ schoensis’’ com- parison, Student’s ¢ is respectively 8.45716 and 8.25106, which shows highly significant differences in view of the small number of specimens. 3. The ratio between the pectoral suture and the sum of the gular and humeral sutures gives the following figures: a) belliana (N = 87) 23.6-69 i ON alten b) mertenst (N = 21) 0-31 m 19 ileal @)) sschoensis’? (N = 2) g-4-42.3) m—34.50) v— 986 d) nogueyr (N = 3) 27.8-27.8-39.1 Notwithstanding an enormous variation coefficient in mertensi, t shows a significant difference between mcrtenst and ‘‘schoen- sis’’ (t = 5.82) and mertensi and belliana (t = 12.71) but not between belliana and ‘‘schoensis.’’ 4. The ratio between the pectoral and the abdominal sutures gives the following figures: a) belliana (N = 86) 21.3-60 m—39.89 v—25.81 b) mertens: (N = 21) 0-31.2 m=18.71 v—42.98 ce) “‘schoensis” (N = 8) 294-374 m=33.5 v= 8.62 d) nogueyt (N = 3) 22.4-36 1 The low figure 57 comes from a juvenile. 4 BREVIORA No. 176 Only the difference between ‘‘schoensis’’ and mertenst proves to be significant with a t of 5.03. Two other ratios not previously used but combining the same measurements have proven useful. 1. The ratio between pectoral suture and the maximum width of the carapace: a) bellana (N = 87) 11.3-24 lol a —slornO b) mertenst (N = 21) 0-138.8 m= 8.95 v=41.87 eG) “schoensis--9 (ON, — 7) 1 balos nn — 12862 ay) S83 Student’s ¢ is significant between belliana and mertensi (9.48), and between ‘‘schoensis’’ and mertensi (6.65), dubiously signifi- eant between belliana and ‘‘schoensis’’ (3.11). The same relation between belliana and ‘‘schoensts’’ becomes, however, significant if analvzed by regression lines (t = 4.26). 2. The ratio between the width of the plastron at level of the lateral ends of the humero-peetoral suture and at the abdominal suture : a) belliana (N = 86) 110-200% m=145.45 v=11.80 b) meriensi (N = 21) 1034479) m= AN 42, y= 34 Student’s ¢ (8.05) is highly significant. As a result of these computations, mertenst appears definitely valid, but ‘‘schoensis’’ seems to be so only for two ratios: 1. the ratio of the sum of humeral and gular sutures to the breadth of the plastron at the level of the humero-pectoral suture and 2. the regression line for the correlation between the length of the pec- toral suture and the maximum breadth of the carapace. From a taxonomic point of view, it does not seem advisable to revive ‘‘schoensis’’ on such slender evidence — more espe- cially as the belliana sample is geographically highly heteroge- neous. A complex clinal variation from Rhodesia to Ethiopia is possible, since we have no data from the populations between Kenya and Ethiopia or Eritrea. Some records from Eritrea (Sordelli 1901, Calabresi 1927, Scortecci 1928) disclose a short- ening of the pectoral suture, as in mertensi. On the other hand, no east-west cline exists between mertensi and belliana: the British Museum specimens from Entebbe and Mount Elgon are clearly mertensi. The two questions propounded have thus the following an- swers: (1) A race ‘**schoensis’’? cannot now be revived for the north- eastern populations of HKinirys belliana. 1962 KINIXYS BELLIANA GRAY a (2) The distribution of Kinirys belliana mertensi includes Uganda, but its northern limit remains unknown. These conclusions are still provisional. More material could prove not only that ‘‘schoensis’’ is valid but that other subspe- cies can be recognized, or on the contrary that even mertensi merges in belliana through Sudanese and Abyssinian popula- tions. Material examined Kimeys belliana belliana Gray MUSEUM OF COMPARATIVE ZOOLOGY : Sudan: Torit (1). Kenya: Golbanti (1), Ithanga Hills (1), Kibwezi (1), Voi (5). Tanganyika: Amboni (1), Kilosa (1), Kiponda to Mitungu (1), Kitaya (1), Mikindani (4), Morogoro (1), Simo near Tabora (2), Turiani (1), Ujiji (2). Zanzibar Island: Zanzibar (1). Nyasaland: Cholo Mtn. (1), Mtimbuka (5). Northern Rhodesia: Isoka (1). Southern Rhodesia: Birchenough Bridge (1), Bula- wayo (1), Hot Springs (2), Lumani (1), Selinda Mtn. (3), Umtali (1). Transvaal: Naauwpoort (1). Katanga: Kapiri (1), Lukafu (1). BRITISH MUSEUM (NATURAL HISTORY): Sudan: Ka- dugli (1). Hthiopia (1), Anseba (2). Somaliland: Berbera (1), near Berbera (1). MUSEE ROYAL vz L’AFRIQUE CENTRALE (Tervuren, Belgium). Kenya: Kibwezi (1), Voi (1). Northern Rhodesia: Abercorn (3). Katanga: Kabambaie (1), Kabinda (1), Kakanda (3), Kansenia (12), Kapiri (8), Lofoi sources (1), Lukafu (2), Lukonzolwa (4), Mwera (1), Ste. Walburge (2). Kivu: Ma- kunega (2). Kinizys belliana mertensi Laurent MUSEUM OF COMPARATIVE ZOOLOGY: Jtuwri: Mahagi- Bort, Gl): BRITISH MUSEUM (NATURAL HISTORY): Uganda: En- tebbe (1), Mt. Elgon (2). MUSEE ROYAL pve L’AFRIQUE CENTRALE (Tervuren, Bel- gium). Congo, without locality (4). Uele: Dika (3), Mauda (1), Niangara (1). Jturz (1): Abimva (1), Gangala na Bodio (3), Mahagi-Port (3). Stanleyville District: Avakubi (1). 6 BREVIORA No. 176 Kinizys belliana nogueyi Lataste. MUSEUM OF COMPARATIVE ZOOLOGY : Dahomey: Bassila (1). Togo: Tohoun (1). Sierra Leone: Kabala (1). Two specimens from Lukolela (western Congo: M.A.C. 4648)? and from the Kwango District (southwestern Congo: M.A.C. 10736) suggest that a differentiated population exists in the lower Congo region, which somewhat resembles mertenst in hav- ing a short pectoral suture and a long abdominal suture. Other specimens are, of course, needed. Key to the races of Kinixys belliana A. Forelimb with 4 claws — Range: Western and northern Cameroon, west towSene galery eis ee ae a end ok oe tae de ae Ae ae K. b. nogueyi Lataste B. Forelimb with 5 claws (occasional specimens have 4 claws) 1—Ratio between the median gular suture + the median humeral suture and the breadth of the plastron at the level of the lateral ends of the humero-pectoral sutures: 75 to 94% (less than 75 in juveniles). Ra- tio between the pectoral suture and the sum of the gular and hu- meral sutures: 0-31% Range: Northeastern Congo and Uganda (presumably also Répub- lique Centre Africaine and eastern Cameroon). See eae) GEER ett a aos rere me ae Ce otra K. b. mertensi Laurent 2—These ratios, respectively, 49 to 83% and 25 to 69%. Range: Sudan east to Eritrea and Somaliland, south to Natal, north- west to Angola and southern Congo. ces ...K. b. belliana Gray BIBLIOGRAPHY CALABRESI, E. 1927. Anfibi e rettili raccolti nella Somalia dai Proff. G. Stefanini N. Puecioni. Atti. Soc. Ital. Sci. Nat. Milano, 66: 14-60, pl. I. LAURENT, R. F. 1956. Contribution 4 l’herpétologie de la région des Grands Lacs de 1’Afrique Centrale. I. Généralités. II. Chéloniens. III. Ophi- diens. Ann. Mus. Congo, Zool., 48: 1-390, pls. I-XXXI. SCORTECCI, G. 1928. Rettili dell ’Eritrea esistenti nelle collezioni del ’Museo Civico di Milano. Atti Soc. Ital. Sei. Nat. Milano, 67: 290-339, figs. 1-8, pls. VIJT-IX. SORDELLI, F. 1901. Anomalia in una testuggine (Cinixys belliana) del Sudan Orien- tale. Atti Soc. Ital. Sci. Nat. Milano, 39: 111-114, figs. 1-2. WerMuTH H. and R. MERTENS 1961. Schildkroten. Krokodile. Briickenechsen. G. Fischer, Jena. 1M.A.C.: Musée Royal de l’Afrique Centrale (Tervuren). . Prem i +f.) ae : (Le 2Cr Fy _ ried _ 7 W 2 Te; S _ | Py - ‘ 6a Pa - ive s ee. - mh ty 0s - _ ocd marty "7 r oO4 7 7 a Lae i 4 year? whic 7 bs 7 PF ) “A. ie x 7 0 a ee a? » a aa. oa; "ae 4 v é - i] 41> Areal i = ' f ; a : a = mT ba 7 ies | ' awit § ae Se Ey Mitel aus § | . ; Voges pe ° + ’ ( | i? Ve ‘. ; at i coer 5 ) vo id v =e ei a By LAY, , ‘ , j F a 7 oe ae a ' e : 7 r% 2 whe tte e a | A ee : : : y > pt fhage Decali ct. 7 j ; as Cc = 1 Serena’ th A Retinal? evel l= Wie |) om call, 5 hee a phi teed 5 oth (p4) Tai Os> tae) Mis Wa - 7 =e 7 a leds PMmvee” (ON 7 hte, ) Se > Bigeea @ ay tpt! es ace r ae OF Hi nlbie en te as esha V4, (hi> lee ar’ ms 5 Vee ee 7 =#gge ; " ats Ut PRyws maps a ‘4h ome a ee | pes alk snl O° Oire, Arie & Da. ore ae : E: : *? BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 27, 1962 NUMBER 177 RHIPIDISTIAN CLASSIFICATION IN RELATION TO TEE ORIGIN OF THE THERAPODS By Keiru 8. THOMSON Department of Biology and Museum of Comparative Zoology, Harvard University INTRODUCTION In an extensive study of the nasal region of the lower gnatho- stomes, Jarvik (1942) concluded that the four families of rhipi- distian Crossopteryeli represent two distinct stocks (superorders according to Lehman, 1959) — the ‘Porolepiformes’ (Porolep- idae and Holoptychidae) and the ‘Osteolepiformes’ (Osteolep- idae and Rhizodontidae). Of major interest to students of vertebrate evolution was his conclusion that the structures seen in the snout of ‘Porolepiformes’ were closely comparable to those seen in the recent Urodela, and those of ‘Osteolepiformes’ were comparable to those of the recent Anura, and that here was evidence of a biphyletic origin of the tetrapods. His work was based upon detailed studies of two genera, Porolepis and Eusthenopteron (a rhizodontid). Jarvik’s material was unusual in that it showed details of the endocranial part of the snout which it is not usually possible to study in fossil crossopterygi- ans; in recent years, however, more such material has been described (Vorobjeva, 1959, 1960a, 1960b; Kuleyzeki, 1960) and Srvig (1957) has published a full treatment of the inter- relations of the Rhipidistia on the basis of the structure of the scales. For the last year or so I have been engaged in a study of the ethmoid region of the osteolepids Megalichthys and Ectosteorhachis, continuing the work started by Romer (1937, 1941). Briefly, the newly available evidence does not support all of Jarvik’s original conclusions. Because of the relation of this 2 BREVIORA No. 177 work to the problem of the origin of the tetrapods and particu- larly to the currently disputed theories of the ancestry of the recent Amphibia, I have decided to publish this short review of the more pertinent points in advance of a more thorough treat- ment. This may also be useful since the work of Vorobjeva, be- ing in Russian, may not be readily available to everyone. This paper forms part of the work to be submitted to the De- partment of Biology, Harvard University, in fulfillment of the requirements for the degree of Doctor of Philosophy; it 1s my pleasure to acknowledge the constant help and guidance of Pro- fessor A. S. Romer during all this study; he and Dr. E. E. Wil- liams and Professor B. Patterson read and criticized the manu- script. I am also grateful for the friendly encouragement of Drs. Jarvik and 4rvie in Stockholm; the former has allowed me to see an as yet unpublished manuscript on this subject (Jarvik, in press). My friend Mr. Simon Karlinsky provided the translations from the Russian. My studies have been sup- ported by the award of N.A.T.O. Science Studentship 3/60/955 by H. M. Department of Scientific and Industrial Research, London, and by the Jeffries Wyman Scholarship at Harvard University. MATERIAL It is necessary to establish the taxonomic status of certain of the rhipidistians concerned in this study; these are: Megalichthys Agassiz 184! (including Ectosteorhachis Cope 1880). Carboniferous and Lower Permian of Europe and North America. Platycephalichthys Vorobjeva 1959. Upper Devonian of WES: S-Re Panderichthys Gress 1941. Upper Devonian of U.S.S.R. Porolepis (P. ex grege posnaniensis [Kade 1858]). Lower Devoman of Poland. Megalichthys-Ectosteorhachis. Ectosteorhachis was originally described by Cope (1880) from material from the Lower Per- mian of Texas; it was later (1891) referred by him to the Car- boniferous genus Megalichthys. In recent years it has been sug- gested several times (see, for example, Romer, 1941) that the two genera are, in fact, distinct and that the Lower Permian material will have to be referred back to Ectosteorhachis. No formal diagnosis of this has been given; however, in advance of 1962 RHIPIDISTIAN RELATIONSHIPS 3 such a diagnosis I shall follow precedent and refer to the Per- mian fish, for convenience, as ‘i ctosteorhachis.’ Megalichthys and Ectosteorhachis are generally acknowledged to be members of the Osteolepidae. Berg (1958) unites them with Owen’s Parabatrachus to form the separate family Para- batrachidae; however, available evidence, e.g. Bystrow (1950), shows that Megalichthys is closely related to Osteolepis itself. Porolepis. Kuleyzcki’s (1960) material of Porolepis is with- out doubt correctly referred to that genus. Platycephalichthys and Panderichthys. These genera are in- volved in a lone series of taxonomic shufflings concerning the genera Cricodus, Dendrodus and Polycodus; they may be con- sidered most conveniently together. The best way to review the situation seems to be to start with the description by Rohon (1889) of several specimens from the Upper Devonian of Russia, some of which he assigned to Dendrodus biporcatus Owen (Ro- hon, 1889, plate 1, figures 1, 5, 9, a skull, and 2,7 8, a tooth and two scales), while others he named Cricodus wenjucovi (Rohon, 1889, plate 1, figures 4, 6, a skull from the River Ojatj, and 3, 11, a skull and lower jaw from the River Sjass). Since these specimens are not given numbers by Rohon, I shall refer to them, for convenience, by the number of the figure by which they are illustrated. Gross (1933) placed those specimens figured in plate 1, figures 1, 5, 4, 5, 6, 9, together with new material from the Baltie Old ted Sandstone, in the genus Pclyplocodus Pander 1860, as the new combination Polyplocodus wenjucovi. Jarvik (1937), how- ever, put ‘‘figures 1, 3, 4, 5, 6, 9, 11’’ in Eusthenopteron as E. wenjucovtl. He also coneluded that the names Cricodus, Den- drodus, and Polyplocodus should not be assigned to any fresh material because of the fragmentary and enigmatic nature of the originals. Hence Gross (1941) named a new genus Pan- derichthys in order to reassign material named by him (1930) Polyplocodus (Cricodus) rhombolepis (see also Gross 1933, 1936). Now there enters a possible source of confusion because Gross (1941) named a second new species — Panderichthys bystrovi — for the material from the Baltic Old Red which he had named Polyplocodus wenjucovi in 1933 (see above), while leaving the Rohon material in Eusthenopteron wenjucovi. Vor- objeva (1960a) named a third species — Panderichthys stol- bovi. 4 BREVIORA No. Lie In 1959 Vorobjeva described a new genus Platycephalichthys, with the type Platycephalichthys bischoffi, based on recently collected material from the Upper Devonian of Russia, and in- cluded in this genus the Rohon specimen from River Ojatj (‘‘fig- ures 4 and 6’’) as a new species Platycephalichthys rohon. Later (1960b) she referred the remaining Rohon material to Eusthenodon Jarvik 1955 as Eusthenodon wenjucovr. Thus the two genera now comprise: Panderichthys P. rhombolepis Gross 1941 P. bystrovi Gross 1941 P. stolbovi Vorobjeva 1960 Platycephalichthys P. bischoffi Vorobjeva 1959 P. rohoni Vorobjeva 1959 From the structure of the scales and teeth, Platycephalichthys is a member of the Rhizodontidae. Panderichthys was assigned by Gross (1941) to the Rhizodontidae but there seems little doubt (Vorobjeva 1960a, and Orvig 1957) that it should be placed with the Osteolepidae. SUBDIVISION OF THE RHIPIDISTIA Jarvik (1942, p. 489) lists a series of differences between Porolepis and Eusthenopteron which he considers to be repre- sentative of a basic split within the Rhipidistia. He also (1942, pp. 417, 495) discusses the connections between these fishes and the recent Amphibia. In general, the same characters are in- volved in the two arguments. In the next few pages I shall review recent findings which indicate that certain of these char- acters no longer support Jarvik’s distinctions. These are all characters which may readily be determined in the fossils; this is in contrast with some of Jarvik’s points involving the passage of nerves and vessels, the refutation of which could be as diffi- cult as their interpretation, Some of Kuleyzcki’s (1960) state- ments may be disputed, as I shall mention later, on the grounds that his material was insufficiently well preserved. This criti- cism cannot apply to Vorobjeva’s work, particularly since the most important of the characters she has described are, as already mentioned, easily ascertained. The recognition, in certain anatomical characters, of resem- blanees between specific rhipidistians and recent amphibians, without the supporting evidence of a fossil lineage (lacking in 1962 RHIPIDISTIAN RELATIONSHIPS 5 the case of the Urodela and Anura) becomes, to a certain ex- tent, a matter of subjective judgment. In the following pages I shall consider primarily the evidence for a basic distinction between ‘Osteolepiformes’ and ‘Porolepiformes,’ for it is upon the validity of this supposed dichotomy that all subsequent phylogenetic hypothesis must rest. The nature of the anterior palatal fenestrae. According to Jarvik (1942, p. 489, ete.), an important difference between the ‘Osteolepiformes’ and the ‘Porolepiformes’ lies in the nature of the paired palatal recesses present between the anterior edges of the vomers and the posterior rim of the premaxillae. In Eusthenopteron this region has the form of a shallow, partially sub-divided groove, the ‘prenasal groove,’ limited posteriorly by the edges of the vomers; in Porolepis there is a pair of ‘pits’ extending backwards between and separating the vomers. These palatal recesses which are described by three names — ‘fossae apicales,’ ‘anterior palatal fenestrae,’ and ‘pre-nasal pits’ — were assumed by Jarvik to have contained glandular organs homologous with the various intermaxillary glands of recent Amphibia. It was further proposed that the condition in Hus- thenopteron foreshadowed that of recent Anura, and the condi- tion in Porolepis that of Urodela. It had already been suggested, however, that these recesses served solely to receive the points of large tusk-like teeth set in the tips of the lower jaws (Holmgren and Stensi6, 1936; Romer, 1937; and now also Kuleyzeki, 1960). The material at my disposal shows quite clearly that this latter explanation is the true one for the Osteolepidae. Plate I shows the tusk fitting into the palatal recess, leaving no room for any glandular strue- ture. That this is also the case in other ‘Osteolepiformes’ may be deduced from the similar large tusks of the lower jaws — for example, in Panderichthys (Gross, 1941). In ‘Porolepiformes’ the situation is very similar, but in this case, as Jarvik has shown, the teeth concerned are a pair of tooth whorls. This extremely interesting discovery provides, incidentally, positive indication that the familiar Onychodus, long known from such tooth whorls, is, in fact, a ‘porolepiform’ rhipidistian (Jarvik, in press). The presence of the paired tooth whorls speaks, indeed, for the unity of the ‘Porolepiformes,’ but the morphological rela- tionship between the type of palatal fenestra and the lower jaw dentition completely precludes any phylogenetic relationship 6 BREVIORA No. 177 between the fenestrae in Rhipidistia and the glandular organs of recent Amphibia. Evidence from the cranial cavity. A point of resemblance, apparently possible of interpretation as evidence of relationship, between Porolepis and the Urodela is that in both ‘*. . . the in- ternasal wall is broad and does not form any nasal septum. It lodges the ethmoid part of the cranial cavity’’ (Jarvik, 1942, p. 417). This, moreover, is supposed to be in direct contrast with the ‘Osteolepiformes’ (and Anura) —by extrapolation from the situation in HLusthenopteron in which the internasal wall is relatively narrow and also solid. Kuleyzeki (1960) noted that his material of Porolepis did not show any forward extension of the cranial cavity between the nasal sacs; his material was perhaps not as good as might be desired fully to substantiate this view, since it consisted entirely of natural casts and not true bony remains. At the same time that Kuleyzeki’s work was published there appeared the work of Vorobjeva. Her material consisted of several portions of the skulls of Platycephalichthys and Panderichthys, both of which are undoubtedly ‘osteolepiform’ and both of which, she states, have an ethmoid extension of the cranial cavity. Her deserip- tion of Platycephalichthys is interesting; ‘*. . . a wide inter- nasal portion (of the braincease) with a cavity stretching forward almost to the front edge of the skull’? (Vorobjeva, 1960a, trans. ). In the face of evidence that a rhizodontid and an osteolepid have a ‘pars ethmoidalis cranialis’ and at least one species of Porolepis may not, one is forced to conclude that this character is of no significance in any attempt to distinguish supra-familial eroupines within the Rhipidistia. One may further bear in mind that the extent of the cranial cavity need have no relation to the extent of the brain contained therein. A dramatic demon- stration of this is afforded by the coelacanth Latimeria (Millot and Anthony, 1958), in which a large cranial cavity contains but a small brain. Presence or absence of the pars ethmoidalis cranialis is thus a rather labile character in the Rhipidistia and, having no great anatomical or functional basis, no phylogenetic speculation may reasonably be drawn from it. Nasal apertures and the nasal cavity. All known ‘Porolepi- formes’ are characterized by the presence of two external nares and a choana. Further, the endocranial opening for the poste- rior naris is confluent with that for the choana. In Husthenop- teron, on the other hand, there is but one external naris and this =~] 1962 RHIPIDISTIAN RELATIONSHIPS is separated from the choana by the lamina nario-choanalis of the endoskeletal nasal capsule. Until the work of Vorobjeva this was believed to hold true for all ‘Osteolepiformes’ but Pan- derichthys is deseribed (Vorobjeva, 1960a) as having both an anterior and a posterior external naris. From her figures it is not possible to determine whether the described confluence of the posterior naris with the fenestra endochoanalis is due merely to a defect in the preservation or not. This is indeed a strange situation for a fish which is in all major respects to be consid- ered ‘osteolepiform. ’ Jarvik has other speculations based on the detailed configura- tion of the nasal cavity itself, but these seem to be more open to dispute in connection with phylogenies spanning 300 million years. Ectosteorhachis certainly lacks most of the ridges, grooves and depressions described in Lusthenopteron and supposed to be typical of all ‘Osteolepiformes.’ Jarvik described in Eusthenopteron a foramen in the post- nasal wall which he states is the posterior endonarinal fenestra (Jarvik’s terminology). This is also present in Ectosteorhachis, but the presence of an anterior naris in Panderichthys may pos- sibly indicate that the single external naris of ‘Osteolepiformes’ is homologous with the posterior naris of Porolepis. In this case the foramen in the post-nasal wall, which Jarvik considers to be the forerunner of the tetrapod naso-lachrymal duct, may pos- sibly have to be interpreted in some other fashion, but consider- ably more evidence is needed to settle this point. Other structures. There are several other points, noted by Jarvik as indicative of a division within the Rhipidistia, which are contradicted by the anatomy of Ectosteorhachis: 1) Jarvik (1942, p. 492) states that a difference between ‘Osteolepiformes’ and ‘Porolepiformes’ is that in the former the vomers are in mesial contact, while in the latter they are separate from each other. Although in the sectioned material at my dis- posal the vomers are shghtly displaced, it seems that these bones are not in mesial contact. Further, they lack the pos- terior extension passing back on either side of the tooth-bearing part of the parasphenoid, which has also been stated to be typi- cal of *Osteolepiformes.’ The vomers are thus much more simi- lar to those of Porolepis than to those of Eusthenopteron;: they lie entirely anterior to the tooth-bearing part of the parasphe- noid. BREVIORA No. 177 CD 2) Jarvik states (1942, p. 492) that in ‘Osteolepiformes’ the parasphenoid is narrow and in ‘Porolepiformes’ it is broad. But in Ectosteorhachis (an osteolepid) only the tooth-bearing part is narrow — the parasphenoid is continued forward and _ later- ally as a broad, if thin, film of bone fused to the ventral surface of the endoecranium (cf. Romer, 1937, p. 19). 3) Jarvik states that lack of the external parietal foramen is typical only of ‘Porolepiformes’; both Megalichthys and Ecto- steorhachis lack it, however. Kuleyzeki further criticizes Jarvik’s description of the snout anatomy of Porolepis, especially the detailed description of the nasal capsule and the canals for nerves and vessels (see espe- cially Kuleyzcki, 1960, pp. 81-94). On the other hand, Jarvik states (in press) that in most details the new material he has of Glyptolepis fully bears out his description of Porolepis. DISCUSSION AND CONCLUSIONS The nature of the fossil remains, by which the Rhipidistia are known, is extremely variable, rangine from the excellent and fairly plentiful material preserved ‘in the round’ of Eus- thenopteron, Ectosteorhachis and Glyptolepis (Jarvik, in press, and in preparation), to the fragments of skulls, isolated teeth, and seales typical of most genera. It is not surprising, there- fore, that most phylogenies and taxonomic studies have been based on the histology of teeth and scales. The latter approach is, of course, open to some doubts and reservations, but the re- cent work of Orvig seems particularly important (see Orvig, 1957, p. 409 for phylogeny ). It is especially interesting to note the positions assigned by Orvig to Panderichthys and Platycephalichthys. According to Jarvik’s interpretations of crossopterygian anatomy, the pres- ence of the pars ethmoidalis cranialis in both genera and the two external nares in Panderichthys would ally them with the Poro- lepis lineage. All other evidence, however (Grvig and Vorob- jJeva), opposes this view and maintains their ‘osteolepiform’ status. One is forced to conclude that these two characters are more labile than was formerly supposed; they fail to show cor- relation with other, diagnostic, characters. Schmalhausen (1959) rejected Jarvik’s theories on the grounds that the two ‘types’ of skull merely reflected the relative pro- portions of the skull, Porolepis being broad-snouted and HLus- thenopteron narrow-snouted. Schmalhausen was referring par- ticularly to the difference between the two types of anterior 1962 RHIPIDISTIAN RELATIONSHIPS 9 palatal fenestra, the nature of which we have already seen to depend on the type of lower jaw dentition. Although Schmal- hausen’s idea is an attractive one, it does not explain the dis- crepancy in the occurrence of the pars ethmoidalis cranialis. The ‘Osteolepiformes’ contain both broad and narrow snouted forms, but the presence of the pars ethmoidalis cranialis is 1n- dependent of this factor: Pars ethmoidalis Snout cranialis ease | Ectosteorhachis Broad Absent Seal ae | Panderichthys Narrow Present BGO aiaae \ Eusthenopteron Narrow Absent ; ~ | Platycephalichthys Broad Present This character is not even related to the relative width of the internasal wall, which seems to be dependent on the relative size of the nasal capsules rather than on the external proportions or intrinsic ‘osteolepiform’/‘ porolepiform’ nature of the snout. I can think of no explanation of the apparently random oc- currence of these characters except that the Rhipidistia form a fairly close-knit group within which comparable variations can occur in all families. There is no doubt that a distinction of some sort can be drawn between a porolepiform and an osteolepiform assemblage (al- though the former is a much more compact group than the lat- ter). The controversy lies in the status to be assigned to each. The two groups have been interpreted by Jarvik (1942, 1955, 1960) to represent a fundamental split within the Rhipidistia and, by extrapolation, within the recent Amphibia also. There are two major objections to the arguments which are presented to substantiate this hypothesis. Firstly, only one genus of ‘Osteo- lepiformes’ (Eusthenopteron) was available for consideration in any detail, and secondly, all the characters dividing the two groups are chosen with reference to the proposed relation to the Urodela or Anura and not at all with reference to the Rhipidis- tia in general. Taking into consideration the evidence presented above and bearing in mind other more detailed points such as are disputed by Kuleyzcki and to which I hope to return in a future work, it seems that the two groups of Rhipidistia are more closely related than has been stated. Furthermore, those characters which might reflect a relationship with particular 10 BREVIORA No. 177 recent Amphibia are present in both groups of rhipidistians and are not, therefore, justifiably so considered. From my own studies of the Rhipidistia, | personally favour the view propounded by Berg (1958) that there are three groups of Rhipidistia: the Porolepiformes, the Osteolepiformes and the Rhizodontiformes. I would allow each group no more than super-familial rank. Literature cited Bere, L. S. 1958. System der rezenten und fossilen Fischartigen und Fische. 310 pp. Deutscher Verlag, Berlin. Bystrow, A. P. 1939. Zahnstruktur der Crossopterygier. Acta. Zool., 20: 283-338. 1950. Microscopic structure of the bones and teeth of the Carbonifer- ous crossopterygian fish Megalichthys (family Osteolepidae). Dokl. Akad. Nauk. U.S.S.R., 34: 119-121. (In Russian.) Copg, E. D. 1880. Second contribution to the history of the Vertebrata of the Permian formation of Texas. Proc. Am. Phil. Soc., 19: 38-58. 1891. On the character of some Palaeozoic fishes. Proce. U.S. Nat. Mus., 14: 447-463. Gross, W. 1930. Die Fische des mittleren Old Red Sud-Livlands. Geol. Paliont. Abh., N.F., 18: 121-156. 1983. Die Fische des baltischen Devons. Palaeontographica, 79, Abt. Ate 2. 1956. Beitrage zur Osteologie baltischer und rheinischer Devon-Cros- sopterygier, Palaont. Z., 18: 129-155. 1941. Uber den Unterkiefer einiger devonischer Crossopterygier. Abh. Preuss. Akad. Wiss., math.-nat. Kl., no. 7: 1-51. HouMGREN, N. and E. A. STENSIO 1986. WKranium und Visceralskelett der Akranier, Cyclostomen und Fische. Jn L. Bolk et alia, Handbuch der Vergleichenden Ana- tomie der Wirbeltiere, vol. 4: 345-353. JARVIK, E. 1937. On the species of Husthenopteron found in Russia and the Baltic States. Bull. Uppsala Geol. Inst., 27: 63-127. 1942. On the structure of the snout of crossopterygians and lower gnathostomes in general. Zool. Bidrag, 21: 237-675. 1955. The oldest tetrapods and their forerunners. Sei. Monthly, 30: 141-154. 1960. Théories de l’évolution des vertébrés. 104 pp. Masson et Cie, Paris. 1962 RHIPIDISTIAN RELATIONSHIPS 11 (in press). On the Porolepiformes and the origin of the urodeles. (Unpub- lished, read before the International Colloquium on Evolution, Paris, May, 1961.) KULCYZCKI, J. 1960. Porolepis (Crossopterygii) from the Lower Devonian of the Holy Cross Mountains. Acta Palaeont. Polonica, 5 (1): 65-106. LEHMAN, J. P. 1959. L’évolution des vertébrés inférieures. 188 pp. Dunod, Paris. MIuLotT, J. and J. ANTHONY 1958. Anatomie de Latimeria chalumnae. Vol. I. Squelette, Muscles et Formations de soutien, 122 pp. Centre National de la Rech- erche Scientifique. Paris. Orvie, T. 1957. Remarks on the vertebrate fauna of the lower Upper Devonian of Escuminae Bay, P.Q., Canada, with especial reference to the porolepiform crossopterygians. Arkiv. Zool., 10 (6): 367-426. ROHON, J. V. 1889. Die Dendrodonten des devonischen Systems in Russland. Mem. Acad. Imp. Sci. St. Petersburg, (7) 36 (14): 1-52. Romer, A. S. 1937. The braincase of the Carboniferous crossopterygian Megalich- thys nitidus. Bull. Mus. Comp. Zool., 82 (1): 1-67. 1941. Notes on the crossopterygian hyomandibular and braincase. J. Morph., 69 (1): 141-160. SCHMALHAUSEN, I. I. 1959. Concerning monophyletism and polyphyletism in relation to the problem of the origin of the land vertebrates. Bull. Mosk. Obshech, Isp. Pri. Biol., 64 (4): 15-33. (In Russian. ) VOROBJEVA, E. I. 1959. A new genus of crossopterygian fish — Platycephalichthys from the Upper Devonian of Lovat. Palaiont. J. Akad. Nauk. U.S.S.R., 1959, no. 3: 95-106. (In Russian.) 1960a. New facts about the genus Panderichthys from the Devonian of the U.S.S.R. Paldont. J. Akad. Nauk. U.S.S.R., 1960, no. 1: 87-96. (In Russian. ) 1960b. Concerning the systematic position of Husthenopteron wenju- covi (Rohon). Palaont. J. Akad. Nauk. U.S.S.R., 1960, no. 2: 121-129. (In Russian.) 13 BREVIORA Non i? anterior palatal fenestra premaxilla dentary tusk Plate 1. Megalichthys from the Seottish Carboniferous. M.C.Z. 8941. Tip of the snout in anterior view, showing premaxillary teeth, dentary tusks, and the anterior palatal fenestrae exposed by a natural break in the specimen: A. Photographed immersed in water. B. Semi-diagrammatic sketch of A, emphasizing the pertinent features. BREVIORA Museum of Comparative Zoology CAMBRIDGE, Mass. DECEMBER 27, 1962 NuMBER 178 ON A NEW SPECIES OF THE EARTHWORM GENUS TRIGASTER BENHAM 1886 (OCTOCHAETIDAE)! By G. E. GATES University of Maine, Orono, Me. The presence, in Cuba, of an apparently endemic Trigaster is of such zoogeographical importance as to contra-indicate further and perhaps indefinite delay in publishing available data. TRIGASTER CAVERNICOLA Nn. sp. b ‘Cueva de Colon,’’ Punta Caguanes, Yaguajay, Las Villas Provinee, Cuba, 0-0-1. G.S. Taboada. (Type in the Museum of Comparative Zoology. ) Katernal characteristics. Size, 143 by 5 mm. Segments, 276. Prostomium, epilobous, tongue open posteriorly but crossed by a deep transverse furrow near anterior margin of 1. Setae, AB < CD < AA which is ca. = or slightly < BC, DD ca. =1,C. First dorsal pore, at 8/9. Clitellum, faintly indicated, possibly saddle-shaped and reach- ing down only to A, on (xii?) xiv-xix(xx?). Spermathecal pores. minute, superficial, midway between A and B, two pairs, equa- torial in vill-ix. Female pore, at A and midway between 13/14 and eq/xiv, on left side. Male and prostatic pores, as well as apertures of setal follicles in the male area, unrecognizable. Seminal grooves, deep and wide, about at B, between eq/xvui and xix/eq. Genital markings, unpaired, transversely placed, in AA, each with a deep transverse, central groove, primarily postsetal on xvi, primarily presetal on xx and xxi but perhaps crossing sites of 19/20 and 20/21. Internal anatomy. Septa, 6/7-10/11 muscular and opaque, 11/12 membranous but with discrete muscular strands. Pigment, 1From research financed by the National Science Foundation. 2 BREVIORA No. 178 red, recognizable only in region of mD, in or associated with the circular musculature. Gizzards, large, in v and vi. Esophagus, deeply constricted at insertions of 11/12-13/14, in xi-xiii with a bifid ventral typhlosole and numerous low, rounded, irregular ridges that are gorged with blood, without discrete calciferous glands. Intestinal origin, apparently in xv. Caeca, intestinal typhlosoles and supra- intestinal glands, lacking. Dorsal blood vessel, single, complete, bifurcating under the brain, the branches uniting over subpharyngeal ganglion to be- come the ventral trunk. Extra-esophageal trunks, blood filled and recognizable alongside the ventral trunk in three segments, posterior portions empty and invisible. Supra-esophageal trunk, recognizable only in x-xiii, Subneural trunk, presumably lacking as no trace of one was found. Hearts, of x-xii are apparently lateral as no bifurcations to supra-esophageal trunk were seen, hearts of xiii latero-esophageal. Nephridia, in at least six longitudinal ranks posteriorly, those of the median rank on each side larger than the others (or median rank composite ?). Male funnels, rather small and thick, without folds, in x-xi. Seminal vesicles, finely acinous, soft, in xii, none recognizable in coagulum of xi. Prostates, 3-4 mm. long, an ectal portion about one half mm. long and with a muscular sheen presumably being the duet. Penial and copulatory setae, not found. GM elands, apparently lacking. Spermathecae, fairly large, adherent to posterior faces of septa and to parietes. Diverticulum, nearly spheroidal, sessile on the duet just below the ampulla. Ovaries, fan-shaped, each with several quite lone egg strings. Reproduction. Spermatozoal iridescence is lacking in the opaque coagulum of spermathecal diverticula and probably also on the male funnels. Ovaries are fully mature but the elitellar epidermis is only slightly tumescent. Postsexual regression may have been under way and if so biparental reproduction is, per- haps, to be anticipated. Ingesta. Soil, brownish. Remarks. The type was received in May 1958 for identification. Better material, fully clitellate if possible, was requested from the collector. Directions for preservation were not followed and subsequent specimens from the type site (received in August, 1958) were not identifiable as to genus or family. A search in 1962 NEW SPECIES OF TRIGASTER a the cave during the dry season of ‘‘Freedom Year’’ (1959) by the same person was fruitless. Collecting in the immediate vicin- ity of Cueva de Colon at the appropriate season was then recom- mended as the species is unlikely to be confined to caves. Unfortunately, receipt of further collections in the near future now seems to be as unlikely as during the past three years. Though the characterization above leaves much to be desired the species is distinguishable from its congeners by location of the spermatheeal pores. Key to species of Trigaster IE MAGiZZaTdsreuwOl acer Ae ye ace neta arc ore Ee OR go PE 2 Gizzards three@rcns ye ee ae ny ih Sas eae en on Chere 3 2. Spermatheeal pores at (or slightly behind?) Gntereéeniental fur- rows, spermathecae adiverticulate ... cS Re a. .tolteca (1) Spermathecal pores equatorial, spermathecaeraiverticilatele a0) eerie: ele eee ee cavernicola 3. Pigmentation, dense, red, resistant to aleohol and formalin. ..rufa (2) IPNeainernir, IEA ONE TOE BSNS IIN OR oo soc bse sae netoeateszese 4 4. Spermathecae adiverticulate and with pores in AB, gizzards in Ti cibce © eM es oe See Seer arenes _lankesteri. (3) Spermatheeae diverticulate and with pores at (unpaired or close to mV, gizzards in v-vil ............ _... intermedia (4) (1) T. tolteca Eisen, 1900, Proe. California Acad. Sci., (3) 2, p. 208. Known only from the original deseription of a single specimen from Toluca, Mexico, at 8000 feet. Type (lost in San Francisco earthquake?), abnormal, preservation ‘‘indifferent.’’ Testes are said to be present in x. (2) T. rufa nom. noy. for Trigaster sp. Gates, 1954, Bull. Mus. Comp. Zool., Harvard College, 111, p. 244. Known only from the very meager description of two macerated specimens from Luquillo Forest, at 1800 feet, Porto Rico. (3) T. lankesteri Benham, 1886, Quart. Jour. Micros. Sei., 27, p. 94. Known only from the original description of the anterior portion of a single specimen from St. Thomas, West Indies. Whereabouts of the type, if still extant, is unknown. Errors in text as well as figures and/or abnormality are suspected. Testes, male funnels and seminal vesicles were not found (lost from sections?) but the amputee was clitellate and the record of sperma- thecal sperm (the only one for the genus) enables an assumption that repro- duction is biparental. Michaelsen assigned Benham’s form the rank of typical subspecies, and named two additional subspecies. (4) T. lankesteri intermedia and calwoodi Michaelsen, 1900, Das Tier- reich, 10, p. 235. Erected on three (or more?) specimens from St. Thomas, one juvenile, one clitellate but stretched and presumably macerated, and one that may have been aclitellate and not quite so much stretched. Types, if extant, presumably in the Hamburg Museum but information as to their fate and condition as yet unavailable. Such information about the two taxa J ie 4 BREVIORA No. 178 as was vouchsafed by Michaelsen is in the definitions of his Tierreich mono- graph. Differences between intermedia and calwoodi now seem unimportant but specific distinction from lankesteri does seem possible if Benham’s characterizations of gizzards, spermathecae, clitellum, ete., are correct. DISCUSSION In Trigaster a simple digestive system without calciferous and supra-intestinal glands, intestinal typhlosoles and caeca but with two or three esophageal gizzards is associated with a meroie excretory system possibly much like that of other octochaetid genera such as Hutyphoeus and Hudichogaster. Four prostates and four spermathecae, supposedly ancestral in the classical phylogenies, have been retamed throughout but wunassociated with penial and copulatory setae. Although testes were said to be present in x of one species (ef. notes above), absence of all seminal vesicles except those of xii suggests that each species is functionally metandrie.* None of the trigasters seems to be common and all may be restricted to more or less remote habitats difficult of aecess from routes usually travelled by man. None of the species is known to have been transported and all are believed to be endemic. The generic range is from Mexico through Cuba? and Porto Rico to the American Virgins. The new Cuban species appears to to be more closely related to the Mexican one. The Porto Rican species seems to be closer to those of St. Thomas. Karthworms (of course after exelusion of the transported forms), because of their confinement to, breeding in, and slow migration through the soil, long have been believed to be second in paleogeographical importance to no other group of animals. Trigaster, perhaps more than any other North American genus, now seems likely to provide especially interesting data if the native faunas of the Mexico-Virgin axis can be studied before they are too profoundly affected by human interference with their habitats. 2 Male funnels of x, in other octochaetid genera, frequently are retained after the testes of that segment ceased to mature sperm. 3 Little is known of the oligochaete faunas of the West Indies, including His- paniola. Indeed, nothing at all is known of the oligochaetes of Santo Domingo- Harvard MCZ Librai wi 302 720 PAAEC AIAT AIR Ass oA ee onm—mie ~ eae ma = aes ~ pa crate ~ pers pane ~ ~ - = “>: a“ ~a ee ere rs os ~ = rears ae on a . pool ~ ~ “> Be ~ as ae never - ~ oe. - . Z = + * ab ~ x Lanne wv - rpery z z n ~ aoe cr a eceeret ~ ree as ~ - Se = a ao. = — rs : a rey vy Aen Aa RASA A hat OCA s Abe ORT ins Heer thie eens geri atten tH) hen Timp etal! en Aine has ngs wee ells rao eg an emee, ea S aepbestnetinnaginehinn teteetineememtticen toes ah the tetinnest ee eee a ee es hs Na NE I ate AA ENA cl AR Hi A EARP ALOACEAE LADD Ot tA Ph AP eid a tte OG AS eae A APOE DOE AEE Ae bye pa any ny petioes Re i nee hea ee Sooua ncowtan- ime mathe! men gapastprnnruasaten. -atiaapkn vi etedeeteaee ee : . r . “ rm ar 7 ; Pocoeatoatintcetietctarte prvanedi-etritntnaatragrnein nineties ee i i ac a ent mye a Ba pea ES 1G No Oak Oo nell CA Ny Rape mee kk Rivesoaer-sooiery seeni ve temsenereosrrtsaipeaeementnsoveenster enenedoormmeae weeretmetnencenr nan careceoemeroeeneoeertepeenborunmicmte wenieaeuremenmmn ramen meme ronan margin “~ ~ 7 = ¥ ~" ~ ae . oe im paveunmnery Devore venete Dyas a A i “a