Fa age le BA tn it meme ere thee He DPC AS ir at

Pevassiea : pire 2 eS get ake

aon Stas

DOES NOT CIRCULATE

BREVIORA

MUSEUM OF COMPARATIVE ZOOLOGY AT

HARVARD COLLEGE, IN CAMBRIDGE

NuMBERS 1-66 1952-1956

CAMBRIDGE, MASS., U.S.A. 1957

Cave oh, et) ty on . vie e

ie Sanaa a eas. 2 5 : are dy) 6 5

Koa

Pw

Now il. Non 2: Nov 3 No. 4 No: 5 No. 6 No. 7 No. 8 No. 9 No. 10

CONTENTS BREVIORA

MuSsEUM OF COMPARATIVE ZOOLOGY

NuMBERS 1-66 1952

A new Panamanian tree frog. By Epwarp H. Taytor. 3 pp., 1 pl. February 8.

A staurotypine skull from the Oligocene of South Da- kota. (Testudinata, Chelydridae). By Ernest WIL- LIAMS. 14 pp., 2 pls. February 8.

Notes on siphonophores. 38. Nectopyramis spinosa n. sp. By Mary Sears. 4 pp. May 23.

A unique remopleuridid trilobite. By H. B. Warirtrne- TON. 10 pp., 1 pl. June 9.

Tricholimnas conditicius is probably a synonym of T'rt- cholimnas sylvestris. (Aves, Rallidae). By James C. GREENWAY, JR. 4 pp. August 29.

Contributions toward a reclassification of the Formicidae. [. Tribe Platythyreini (Hymenoptera). By WILLIAM L. Brown, JR. 6 pp. August 29.

On the nomenclature of the Pacific gray whale. By W1L- LIAM E. ScHEVILL. 3 pp. September 29.

A new species of the cyclostome genus Paramyzine from the Gulf of Mexico. By Henry B. BickELow and W1t- uIAM C. ScHROEDER. 10 pp. October 21.

New species of earthworms from the Arnold Arboretum, Boston. By G. E. Gates. 3 pp. October 21.

On the earthworms of New Hampshire. By G. E. GatTEs. 3 pp. December 29.

No.

mB ol

eli

1953

Characters and synonymies among the genera of ants. Part I. By Wiuutam L. Brown, Jr. 13 pp. March 20.

A eave faunule from western Puerto Rico with a diseus- sion of the genus Isolobodon. By Tuomas E. Reyn- oups, Kart F. KoopMan and ERNEST 1B WILLIAMS. 7 pp., 1 pl. April 23.

Fossils and the distribution of chelyid turtles. 1. ‘“Hydraspis’’ leithii (Carter) in the Eocene of India is a pelomedusid. By Ernest WiuuiamMs. 8 pp., 3 pls. April 23.

Record of a hermaphroditic horseshoe crab, Limulus polyphemus L. By Joun P. Baptist. 2 pp., 2 pls. May 28.

further notes on the earthworms of the Arnold Arbore- tum, Boston. By G. E. Gates. 9 pp. May 15.

Notes on the races of Micrurus frontalis (Duméril, Duméril and Bibron). By BENJAMIN SHREVE. 6 pp. May 28.

A new Ordovician trilobite from Florida. By H. B. WHITTINGTON. 6 pp., 1 pl. May 28.

Characters and synonymies among the genera of ants. Part II. By Wiuuram L. Brown, Jr. 8 pp. Septem- ber 23.

Lower Cretaceous nautiloids from Texas. By BERNHARD KumMEL. 11 pp., 2 pls. September 23.

Lower Triassic Salt Range nautiloids. By BERNHARD KummMeEL. 8 pp., 2 pls. September 23.

The ancestry of the family Nautilidae. By BERNHARD KuMMEL. 7 pp., 1 pl. September 23.

Notes on the ant, Leptothorazr obliquicanthus Cole (Hy- menoptera: Formicidae). By Rosert E. Greee. 3 pp. October 13.

A new species of Hypognatha from Panama. By ARTHUR M. Cuickerine. § pp. October 23.

No.

No.

No. 35.

24.

eA

. 30.

voile

. 32.

. 33.

. 34.

1954

A new family, a new genus, and two new species of batoid fishes from the Gulf of Mexico. By Henry B. BIGELow and WILLIAM C. SCHROEDER. 16 pp. Janu- ary 27.

A new Miocene species of Pelusios and the evolution of that genus. By Ernest Wiuutams. 7 pp., 4 pls. Janu- ary 28.

A preliminary list of the earthworms of northern New Jersey with notes. By H. Davies. 13 pp. February 3.

Anterior regeneration in a sexthecal species of lumbri- cid earthworm. By G. E. Gates. 5 pp. February 5.

Clemmydopsis Boda a valid lineage of emydine turtles from the European Tertiary. By Ernest WILLIAMS. 9 pp. February 8.

Absence of mesoplastra in a Pelomedusa (Testudines, Pelomedusidae). By Ernest WiLuiAms. 4 pp.. 2 pls. February 8.

Aestivation in a Permian lungfish. By ALFRED S. RoMER and Everetr C. Ouson. 8 pp., 1 pl. February 8.

New freshwater gastropod mollusks of the African genus Lanistes. By T. Pan. 4 pp. March 3.

Fossils and the distribution of chelyid turtles. 2. Addi- tional reputed chelyid turtles on northern continents: Palaeaspis conybearu (Owen) —a pelomedusid. By ERNEsT WILLIAMS. 6 pp. March 12.

Systematic and other notes on some of the smaller species of the ant genus Rhytidoponera Mayr. By Wiuuiam L. Brown, Jr. 10 pp. May 14.

A review of the coxalis group of the ant genus Sticto- ponera Mayr. By Witu1am L. Brown, Jr. 10 pp. July 20.

New or rediscovered pelomedusid skulls from the Tertiary of Africa and Asia (Testudines, Pelomedusidae). 1. Dacquemys paleomorpha, new genus, new species from the Lower Oligocene of the Fayum, Egypt. By Ernes? WiuuiaMs. 8 pp., 1 pl. July 28.

No.

No.

No.

36.

noo:

. 40.

41.

. 43.

. 44.

. 46.

47.

Present knowledge of the snake Elachistodon wester- manni Reinhardt. By Cart Gans and ERNEsT E. WiuuiamMs. 17 pp. August 6.

On the evolution of an oriental earthworm species, Pheretima anomala Michaelsen 1907. By G. E. GATEs. S pp. August 18.

Onnia (Trilobita) from Venezuela. By H. B. Wurr- TINGTON. 5 pp., 1 pl. November 28.

New or redescribed pelomedusid skulis from the Terti- ary of Africa and Asia (Testudines, Pelomedusidae ). 2. A podoenemide skull from the Miocene of Moghara, Egypt. By Ernest Witiiams. 8 pp., 2 pls. Novem- ber 24.

Some mollusks from the continental slope of northeast- ern North America. By ArTHUR H. CLARKE, JR. 11 pp. November 29.

Comments on the classification of rodents. By ALBERT E. Woop. 9 pp. December 17.

1955

A new salamander of the genus Parvimolge from Mexico. By Gerorce B. Rass. 9 pp. February 28.

Speed-induced skin folds in the bottle-nosed porpoise, Tursiops truncatus. By FRANK 8. EssaPIAN. 4 pp., 9 plss April i:

A new Murex from Matanzas, Cuba. By WILLIAM J. CLENCH. 2 pp., 1 pl. April 8.

Palaeotaricha oligocenica, new genus and species, an Oligocene salamander from Oregon. By RicHarp vAN FRANK. 12 pp., 3 pls. June 13.

Cave-fossil vertebrates from Camaguey, Cuba. By Karu F. KoopMan and Ropoutro RurBAu. 8 pp. June 24.

A new species of whiptailed lizard (genus Cnemido- phorus) from the Colorado plateau of Arizona, New Mexico, Colorado, and Utah. By Cuaries H. Lowe, Jn. Ipp. July 12.

No.

No.

48.

149:

. 00.

54.

. 60.

Notes on American earthworms of the family Lum- bricidae. I-II. By G. E. Gates. 12 pp. October 10.

Three new shark records from the Gulf of Mexico. By Henry B. Bicgetow, W. C. SCHROEDER, and STEWART SPRINGER. 12 pp. November 18.

New frogs of the genera Asterophrys and Oreophryne from New Guinea. By ArtrHuR LovertpaGr. 5 pp. November 30.

1956

A small mustelid from the Thomas Farm Miocene. By STANLEY J. OLSEN. 5 pp. January 27.

Remarks on some Miocene anurans from Florida, with a description of a new species of Hyla. By WALTER AUFFENBERG. 11 pp. April 6.

Food-finding by a captive porpoise (Tursiops trun- catus). By WiuuiAM E. ScHrvini and BarBara Law- RENCE. 15 pp. April 6.

A revision of the genus Brachymeles (Seincidae), with descriptions of new species and subspecies. By WALTER C. Brown. 19 pp. June 6.

Rediscovery of. Hyla dorsalis and Lechriodus papuanus in New Guinea. By ARTHUR LOVERIDGE, 4 pp. June 6.

Notes on the Jamaican and Cayman Island lizards of the genus Celestus. By PENNY NORSEEN COUSENS. 6 pp. June 15.

Three new species of Mimetidae (Araneae) from Pan- ama. By Artuur M. Cuickerina. 14 pp. June 29. Sequence of Passerine families (Aves). By E. Mayr

and J. C. GREENWAY, JR. 11 pp. June 29. A new subgenus of Chamaeleo from Rhodesia and new

race of Mabuya from Kenya Colony. By ArTHUR LOVERIDGE. 4 pp. September 12.

A new species of Agriognatha from Jamaica, B.W.I. By ArtHur M. CHICKERING. 7 pp. September 12.

No.

No.

No. 6

No

GD)

. 66.

On regeneration by earthworms of a species of the lum- bricid genus Dendrobaena Eisen 1874. By G. E. GATES. 6 pp. September 14.

A third leaf-nosed species of the lizard genus Anolis from South America. By JaAmMEs A. PETERS and Gus- ‘pavo OrcEs-V. 8 pp. October 3.

New bathyal Isopoda from the Caribbean with observa- tions on their nutrition. By Roperr J. Menzies. 10 pp. October 11.

Rare species of Copepoda, Calanoidea, taken from the Izu Region. By Oronrro Tanaka. 8 pp. October 11. A new species of Agriognatha (Araneae, Argiopidae )

from Panama. By ArtHuR M. CHICKERING. 7 pp. November 20.

The Caninae of the Thomas Farm Miocene. By StTan- LEY J. OLSEN. 12 pp. November 27.

INDEX OF AUTHORS BREVIORA

MUSEUM OF COMPARATIVE ZOOLOGY NuMBERsS 1-66

1952-56 No. PSERENBERG, WALTER «2: oie Sc cldn skies °. 52 EP ISA ROLUN Me 3) atehes t 1 o oats te She BRINE, «A .. 6 Sng 14 BiegELow, Henry B. and WILLIAM C. SCHROEDER ......... 8, 24 BigELow, Henry B., W. C. ScHROEDER and STEWaRT RIBRUN GER aes ais a oh. pee lee tes eee OG. say aan 49 RON MATER Co pcisc.)-5 ane See 2 ery = ee Os! ISROW NE WVIGLTAMG Ii) Ral. 0 2. veut tat se) 6, dl 18188. 84 WHICKERING/ ARTHUR DE. .(: 5.1.5 «tee. sates ee 20, Di 0009 CLARKE, ARTHUR H., JR. .... 40 (INC ET paNV TITAN elie we ots. 2s. 2 2) fp eh aa cl nde cea rie ie 44 WONSENS SO ENN WNORSEEN ... ..:. ha ve ine). slo cane 56 Daviss, H. 26 FISSAPTAN, URANEK (93.05... -.-.- + « ie Chis aera" hs as a 3 Gans, CaRL and Ernest E. WILLIAMS .................. 36 Golmigorgye 161) SRN Ue IU oe ee 9, 10, 15, 27, 37, 48, 61 SREMNAUVAEE AIMS COUN el Rey sc. 2 ase > Padawan ohepciares See 5 GEEMINWAY.t- CdR. and H. MAYR.% 3; hee: 58 PREGGO MIROBERT (Hi eile ag. ose ae 99 Koopman, Karu F., and RopoLro RUIBAL ... 46 Koopman, Karu F., THomas E. REyYNoups and ERNEst BL WV RUBIA MS 24S boro ae ope 12 PO MMEE ISERNEARD (0% 052725 wjsirge wedtae hf aol. ee 19, 20, 21 LAWRENCE, BARBARA and WILLIAM E. SCHEVILL ..... 53

LGVERIDGEA AG TUR caer ese e Sem eede 50, 55, 59

Own CHAR ES Bil. cman choo: eee gee eee Fe amo oe 47 MAwR Hy sand: Jit CaGREbINpwaAty, c) Rad) <7 geee ene ete he ce eer eae 58 MENZIES) GORI Tee oe See. aes oe eee oe ie), Se heel 63 (ONES TOTS ea SUW UNb) a as MAN cei or CMR eny ah Aco) Ct Aa 51, 66 - OLsoN, Everett C. and ALFRED S. ROMER ................ 30 Orcrs-V., Gustavo and Jams A. PETERS .:............¢. 62 1 PUN eid NR rca Stat bon ES I ey, 1. a ee, Sete 8 31 PETERS, JAMES Apand:GuSTAVO ORGHS-V ols)... ..22s507e 506 62 RARBG RORGE DO aaer te a2 22 sin tie REI arn Sok 2a eee 42 REYNOLDs, THoMAsS E., Karu F. KoopMAN and ERNEsT EK.

Wigs TAUNIGREPIERE ee ds if We ee ER tod Ake a 12 Romer, Auennp is, and HveRErTr ©. OUSON 02.2.) 2.4 seee 30 RUIBAL; RopoLro and Karu F’. KoOPMAN ..............-- 46 SCHEV wa EbaNAME JH), 22k). ss ces nee Gata oc ee 7 SCHEVILL, WILLIAM E. and BARBARA LAWRENCE .......... 5S

ScHROEDER, WILLIAM C. and Henry B. BIGELow ........ 8, 24

ScHRoEDER, W. C., Henry B. BigELow and STEWART

SPRINGIR 08 [eso ccs ace eee ee 49 SEARS. MAR Wi. ceStet s. 5 ec ER: Geen ee hat ee tae 3 SEREVE, (SENUAMING: . . sl:+ oat. Meera cho, eo eee eee ee 16 SPRINGER, STEWART, Henry B. BiaELow and W. C.

SCHROEDERS —c4. 54 Sone Ce Eee el. ee ee 49 iDanacas) OPOBITO: 2. 3 ...1 J serve eae te coat ae eee 64 WAviOR.. MDWARD: J” .%.- .,: secaateigeat oils serene eee il WAN GM RANK? (RIGEDARD: |< 5.0. \. 2 dey eke, cig Set 8 ee aes ee 45 VEERING DON Ely Bern 2 pain apne naee teeta ls rata 4,17, 38 NV PANS HIRST o2.05 24).2 3 2). yee 2, 13, 25, 28,29 S2e some WaburAmMs, HRNESsT EH. and Caru,GANS 2°34 45-25 ee ee 36

WiuiAMs, Ernest E., Kart F. KoopMAN and THomas E. IRE VNOMDS. “sos Moun ch. A ic a One LO ee eee a

WioODTALBERT Bs avis. sass 2 See eee eh ge ete eel 41

- i

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass.

Frsruary 8, 1952

NUMBER |

A NEW PANAMANIAN TREE FROG

By Epwarp H. TayLor Department of Zoology, University of Kansas, Lawrence, Kansas

While studying Central American frogs in the Museum of Com- parative Zoology, Harvard College, a small, presumably undescribed frog was noted, and Mr. Arthur Loveridge, with his characteristic kindness, suggested that I study the form and describe it if it proved new. The most characteristic of the differential features of this di- minutive frog is the very considerable enlargement of the horny spines on the nuptial swelling of the first finger. It is presumed that it is a rivulet species, living in the neighborhood of small streams, rather than a bromeliad species.

After my study of the specimen and comparison with other known species of southern Central America, | conclude that the species is new and I present herewith its description.

Hy La SHREVEI sp. noy.

Type. M.C.Z. No. 26769 o. La Loma, Chiriquicito, Reptblica de Panama; EK. R. Dunn and C. Duryea, collectors.

Diagnosis. A diminutive frog (25.2 mm.) with a broad, rather flat head; skin smooth; outer fingers with a trace of web, and well-developed dises; canthus rostralis wanting; nostrils nearly terminal; tympanum large directed strongly upward, its diameter at least half of eye length;

2 BREVIORA No. |

eye directed obliquely forward; vomerine teeth in two rounded fasci- culi, lying almost completely behind posterior level of large choanae; no web between two inner toes; others one third, to one half webbed; first finger of male with 19-22 large horny spines.

Description of the type. Head flattened, broader than body, its width (10 mm.) greater than its length (9.1 mm.); tympanum large, semi- transparent, pigmented, facing more upward than outward, the upper edge obscured by a fold arising at eye and curving back to jaw angle, its diameter (1.5 mm.) equal to half length of eye opening (8 mm.); distance between tympanum and eye 2.8 mm.; eye prominent, directed somewhat forward; length of snout less than eye length (2.85 mm.); canthus absent, loreal region concave, sloping obliquely to lip; nostrils nearly terminal, the areas about them strongly swollen with a de- pression between; no notch in upper lip; groove from angle cf jaw touches tympanum; choanae subcircular, the distance between them 2 mm.; diameter (transverse) of choana .65 mm.; vomerine teeth in two somewhat circular fasciculi almost completely behind the posterior level of choanae, the teeth in somewhat curving rows; they are equally separated from choanae and from each other, a fasciculus as large as a choana; tongue rather thick, broader than long, not or scarcely notched behind; vocal slits very large, lateral; openings of the palatal gland forming a series of separate slitlike openings arranged in a broad V-shape about the middle of palate.

Arm very short, the wrist not reaching tip of snout; first finger short, greatly expanded at base by a very large nuptial swelling and a large metacarpal tubercle; swelling covered with a black, horny plate bearing 19-22 large horny spines; dises of fingers smaller than tympa- num, that on first finger only little smaller than those on outer fingers; a small trace of webbing between outer fingers; subarticular tubercles single; supernumerary tubercles numerous; palmar tubercle partly divided; a slight axillary web present; heel reaches forward to edge of eye; tarsal fold represented by a series of tubercles growing more distinct on the proximal end of tarsus; a large inner metatarsal tubercle and a small distinct outer; no web between first and second toes; one fourth webbed between second and third; approximately half webbed between the third and fourth and two fifths webbed be- tween the fourth and fifth, the web forming a slight ridge to dises on sides of digits; subarticular and supernumerary tubercle distinct. Skin on dorsum smooth, slightly wrinkled or corrugated on sides; chin smooth; breast, venter, part of the ventral surface of thighs and region lateral to vent, strongly granular; a rather elongate anal flap, the vent opening midway between upper and lower surface of thighs, followed

1952 NEW TREE FROG

by a groove bordered by small granules and on ventral surface by a pair of distinctly larger granules; fold above tympanum curves down to jaw angle.

Color. In preservative, light brown above with a brownish-white shade below; the pigment is in minute chromatophores, that can be clearly seen under a lens; no pigmentation on under surfaces except on edge of jaw, under surface of thigh, part of under surface of tibia and some scattered flecks under hand and foot; upper eyelids appearing dark from the dark covering of the eyeball; tip of snout somewhat darker than dorsum.

Measurements in mm. Snout to vent 25.2; width of head 10; length of head 9.1; diameter of tympanum 1.5; length of eye opening 3; snout length 2.85; arm 13; hand 7.2; leg 38; tibia 12; foot and tarsus 15.

Remarks. The species is probably related to Hyla zeteki. There are numerous similarities in structure, and the measurements are similar but the most striking differences appear to be in the remarkable nuptial asperities on the first finger, the position and direction, and the larger size of the tympanum (14 instead of 14 of eye); the absence of the spectacle-like markings about eyes. The characteristic anal decoration of zeteki is absent, and the strong granulation on under surface of arm, and on sides is absent.

The species is named for Mr. Benjamin Shreve of the Museum of Comparative Zoology, Harvard College.

4 BREVIORA No. |

Hyla shrevei sp. nov. M.C.Z. No. 26769, La Loma, Chiriquicito, Reptblica de Panama. KE. R. Dunn and C. Duryea, collectors. (Type o X 2.) 1. Dorsal

view. 2. Lateral view of head. 3. Enlargement of nuptial spines on inner digit.

BREVIORA

Museum of Comparative Zoology CAMBRIDGE, Mass. Fepruary 8, 1952 NuMBER 2

A STAUROTYPINE SKULL FROM THE OLIGOCENE OF SOUTH DAKOTA

(Testudinata, Chelydridae)

By ERNEST WILLIAMS

INTRODUCTION

The modern representatives of the Staurotypinae (two genera, Staurotypus and Claudius, and three described species) are restricted to Southern Mexico and Central America. No fossils belonging to these living genera are known. A form known only from a single perfect shell from the Oligocene (Chadron) of South Dakota, Xenochelys formosa Hay, seems to be the only previously described fossil record of this subfamily.

Recognition of a staurotypine from shell material as good as that of the unique type of Xenochelys (A.M.N.H. No. 1097) is quite easy. Staurotypines differ from chelydrines in having only 23 instead of 25 marginals and 21 rather than 23 peripherals. The same feature dis- tinguishes them from dermatemydids. They differ from kinosternines in possessing an entoplastron. In these characters Xenochclys is clearly staurotypine. In the elongation of the first vertebral scute and in its precise shape Xenochelys closely resembles Staurotypus. The neural series is much as in Staurotypus. The plastron of Xenochelys has a reduced number of shields, the pectorals and abdominals having ap- parently fused. This again is a staurotypine or chelydrine feature. The general form and height of the shell are very Stawrotypus-like. There is also some suggestion of the tricarinate condition found in Staurotypus in the carapace of Xenochelys.

2 BREVIORA NO. 2

In other respects the shell of Xenochelys is more primitive (more dermatemydid-like) than any other member of the chelydrid series. The plastron is relatively large (although the bridge is quite narrow as in chelydrids, not as in dermatemydids). There is a xiphiplastral notch. The nuchal scute is small. There is a trace of nuchal indentation.!

In shell characters, therefore, Xenochelys is a good structural inter- mediate between Dermatemys and Staurotypus, and it is also a temporal and phylogenetic intermediate between the complex of fossil forms called dermatemydid and the staurotypine section of the Recent Chelydridae.

It should be noted that Hay was not under any misapprehensions as to the affinities of Xenochelys. He quite explicitly cited Staurotypus and Claudius as ‘‘the nearest living relatives of Xenochelys’’ (1908, p. 282), and in his assignment of the form to the Dermatemydidae he merely followed Boulenger’s 1889 catalog of the Recent turtles in which the Staurotypinae were associated with the Dermatemydidae. When he published his 1908 monograph he had probably not seen Siebenrock’s 1907 monograph of the “Cinosternidae” in which the close affinities of the Staurotypinae and Kinosterninae were for the first time made clear; he thus missed an opportunity to point up more clearly the apparent ancestral position of the South Dakota genus.

In my 1950 classification of the testudinate order I united the chelydrines, staurotypines and kinosternines in the family Chelydridae as understood in a broad sense. To do so somewhat obscures the ex- treme closeness of relationship of staurotypines and kinosternines, which differ in the simultaneous loss of an entoplastron and acquisition of more or less of a box turtle habitus by the latter. The chelydrid series is surely a natural group, but within that group the staurotypines and kinosternines stand very much closer to one another than to the chelydrines.

In the Oligocene this specialized section of the Chelydridae sensu lato was evidently fully distinct in shell characters, if still somewhat primitive in a few features. It has, however, not previously been known to what extent the skull had evolved concomitantly with the shell.

In the collection of the Department of Geology, Princeton Uni- versity, I have now found a skull (No. 13686), likewise from the Chadronian Oligocene of South Dakota, which surely belongs to a member of the staurotypine-kinosternine section of the Chelydridae. It is distinctly more specialized in a number of ways than any previ-

1 Most of these characters might also be counted as kinosternine resemblances.

1952 A STAUROTYPINE SKULL 3

ously known staurotypine or kinosternine skull. Although incomplete it merits extended description and discussion. In the section which follows, the skull is described in detail and compared with Claudius augustatus, Staurotypus salvinii, and Sternotherus carinatus.

DESCRIPTION OF THE PRINCETON SKULL

The skull is complete as far as the postorbital bar. Behind this, however, only the parietals, pterygoids, basisphenoid and basioccipital are retained, all of them somewhat fragmented and incomplete. Sutures are rather difficult to make out because of fractures in critical areas.

The profile of the face is strikingly like that of Stawrotypus or Sternotherus. The prefrontals project dorsally above the narial opening but laterally are somewhat retracted, so that in lateral view the nostril is seen as a distinct angular indentation. The premaxillae unite in the formation of a distinct median beak, while posterior to this median projection the contours of premaxilla and maxilla form a smoothly sinuous curve, which, however, is doubly incurved in the fossil in contrast to both Stawrotypus and Sternotherus. The depth of the premaxilla is markedly greater than in Stauwrotypus. The orbit is rela- tively smaller than in Stawrotypus and even more distinctly lateral (in contrast to dorsolateral) than in that form. The maxilla below the orbit is marked by a distinct groove running down to the second incurving of the lateral festooning of the jaw. The postorbital bar is rather wide, half the rostro-caudal length of the orbit rather than between 14 and 14 as in Staurotypus.

In lateral view the prootic is seen to project far forward about the pterygoid in a very exceptionally developed “‘crista praetemporalis”’ (Siebenrock 1897). Only a narrow channel separates these two bones. The dorsal margin of the prootic is continued also in a ridge on the parietal. A similar forward projection of prootic is seen in Staurotypus (and in Graptemys) but is far less extreme.

The “‘crista praetemporalis” which is the feature exaggeratedly de- veloped in the fossil and in Staurotypus and Graptemys serves to modify and increase the leverage of the jaw muscles (Zdansky 1924, pp. 101-104). All three genera have widened alveolar surfaces of the maxilla with strong tendency to formation of a secondary palate. In the case of Graplemys the wide alveolar surfaces are known to be associated with a malacophagous diet. Probably in the case of the fossil a strengthening of the jaw action and a similar diet are to be inferred.

4 BREVIORA No. 2

The dorsal planum of the parietal continues the remarkably flat dorsal margin of the face.

In anterior view the nostril is very small, much as in Sternotherus, not as in Staurotypus. The prefrontals are anteriorly distinctly convex, posteriorly are very flat. Behind them the frontoparietal area rises somewhat more sharply than in Staurotypus.

In palatal view the maxillae have united in the formation of an extensive secondary palate, complete in front with a pronounced median ridge but incomplete in the midline behind. In this secondary palate the palatines share to about the same extent as in Staurotypus and to a significantly greater extent than in Sternotherus. A striking and unique feature is the very impressive dorsomedial slope of the palatal roof. To a very slight extent this condition is prefigured in the other genera, particularly Stawrotypus, but the difference is very con- siderable: in this respect no other genus is at all close. There are no ectopterygoid processes and the pterygoids also are bowed dorsally at the midline, so that their lateral flanges are very strong and high, though spreading wide apart. Anteriorly the premaxilla has the deep pit for the tip of the lower jar characteristic of chelydrids. In the specimen it breaks through into the narial region. As in Stawrotypus and Claudius (differing in this from all examined kinosternines and chelydrines), foramina incisiva appear to be lacking in the fossil. At the postorbital bar the palate, and thus the whole outline of the skull, is very expanded from the side: this form must have been decidedly brachycephalic. The waist of the pterygoids, however, is only moder- ately broad, as in Sternotherus, not very broad as in Staurotypus or narrow as in Claudius. The basisphenoidal suture is not clearly dis- tinguished from breaks in this region, but it seems probable that the exposed portion of this bone was very short and broad, not tending to be elongate craniocaudally as in Staurotypus. The infracondylar depression, so marked in Staurotypus, is less distinct in this form, as also in Sternotherus, but not so weak as in Claudius.

In dorsal view the great breadth of the skull at the postorbital bar is again evident, along with the considerable incurving of the skull contours just in front of the orbits. The origin of the supraoccipital spine is indicated by a triangular plane surface with well defined lateral margins, as in Staurotypus and Sternotherus, not as in Claudius.

The sutures bounding the frontals are somewhat obscured by breaks, but it is extremely probable that as in other Chelydridae the frontals occupy a very small area and do not enter the orbits.

1952 A STAUROTYPINE SKULL 5

SKULL CHARACTERS IN THE CHELYDRIDAE AND THE ALLOCATION OF THE FOSSIL

For the comparisons made in the course of the description just given I have had available the skeletal collections of the Museum of Com- parative Zoology and of the American Museum of Natural History. Included in the M.C.Z. collection are a young and an adult skull of Staurotypus (the young specimen, M.C.Z. No. 4989, is figured), while the A.M.N.H. collection has furnished for study a skull of Claudius (A.M.N.H. No. 65865).

Because the fossil is a fragment only, comparisons must remain incomplete and portions of the skull which might be diagnostically significant are unavailable. In this circumstance and in the absence of more complete knowledge of variation and difference within the genera Kinosternon and Sternotherus (valuable information which we may hope to learn in the forthcoming revision of these genera by Dr. Norman Hartweg) I do not attempt to discriminate too finely the affinities of the fossil.

However some rather general discussion is possible. We may first consider what characters define a skull as chelydrid in the broad sense, then what features are chelydrine, staurotypine, or kinosternine, and finally what provisional allocation of the fossil skull is possible and expedient.

There are six genera of living chelydrids (Chelydra, Macroclemys, Staurotypus, Claudius, Kinosternon, Sternotherus), and of these Sterno- therus might quite properly be relegated to the synonymy of Kino- sternon. The fossil record adds a few more (Acherontemys, Chelydrops, Chelydropsis, Xenochelys). In contrast the Testudinidae has about 30 living genera while the fossil record brings the count up to about 50. It is not surprising, therefore, that the Chelydridae seem a more closely knit group than do the Testudinidae, even if the rather isolated Platysternon is omitted from the latter assemblage.

If the skulls of chelydrids (all living forms North America, a few Tertiary fossils European) are compared only with the skulls of North American or European testudinids very clear distinctions are evident. If, however, the comparison is extended to the very rich testudinid fauna of Southeast Asia where, so far as known, chelydrids have never occurred, some of the forms to be found there bridge over the differences which were thought to be significant. Even so astute an observer as Baur, and one so familiar with the testudinate order, found it possible

A discussion of the anatomy of Claudtus is in preparation.

6 BREVIORA NO. 2

to refer Adelochelys (= Orlitia) to the ‘‘Chelydroidea’”’ when he had the skull only, though the shell would have placed the genus without question in the Emydinae.

The distinctions between the Chelydridae and Testudinidae are wide enough, when all parts of their anatomy are taken into account and the trends within them are considered, that there is no doubt that, although related, they have long been separated, perhaps since the Cretaceous, certainly since earliest Tertiary. Thus there are differ- ences in cervical formula (Williams 1950), in the form of the eighth cervical vertebra, in degree of development of the costiform processes of the nuchal, in the presence versus absence of inframarginals, in the characteristic reduction of the plastron or its elements in the Chely- dridae, in the proximal end of the femur, and in the absence in chely- drids of gaudy or bright patterns.

However, in a way which seems very characteristic of turtles, few of these features hold good with complete fidelity in all cases. In cervical formula there is a striking difference between the two families in that the eighth cervical is biconvex in the Testudinidae, procoelous in the Chelydridae. A few individuals of the Testudinidae (mostly advanced tortoises) vary in the direction of the condition of the Chelydridae, but no chelydrids are known to vary in the direction of the testudinid condition. The eighth cervical vertebra tends to differ in the two families, the Testudinidae generally showing three ventral crests on the centrum, though the lateral ones may be barely indicated, the Chelydridae showing a single median crest which may divide into two (some Chelydra, kinosternines).

The costiform processes of the nuchal are typically much developed in the Chelydridae, relatively little developed in adult testudinids, but the kinosternine section of the chelydrids approaches the condition of the testudinids, and young emydines have this process rather strongly developed. Inframarginals are never normally present in most testu- dinids, but there is an exception in the case of Platysternon, and inframarginals do occur as individual variations in Chrysemys picta (A.M.N.H. specimens to be reported on by Samuel McDowell). The plastron is never reduced in testudinids; it is strikingly reduced in protective efficiency in chelydrines and staurotypines; in some species of the kinosternines it regains its complete coverage of the ventral surface, but prior to this redevelopment it had lost one of the bones normally present in unreduced plastra. In the testudinid femur a fusion of the trochanters tends to limit the intertrochanteric fossa to a shallow dorsal pit, though a number of emydines (and Platysternon)

ry

1952 A STAUROTYPINE SKULL

have the juncture barely suggested. In chelydrids as in most turtles the intertrochanteric fossa is a widely open groove. The lack of bright patterns in the Chelydridae, while characteristic enough of the rela- tively few genera and species involved, is not consistently contrasted with the presence of such patterns in the testudinids (see, for example, the uniform pigmentation of Galapagos tortoises and of some of the Asiatic emydines.)

Nevertheless, in spite of these exceptions, these postcranial charac- ters, external and internal, permit in combination a clear discrimination of the two families.

Several characters distinguish the skulls of chelydrids and testu- dinids, but here even more than in postcranial characters exceptions reduce the utility of single characters.

Chelydrids usually possess at least traces or indications of a sharp median beak or ‘“‘tooth” on the upper jaw. But this is absent in some kinosternines, and while many testudinids have a notch here, a few (e.g., Terrapene, Cuora) have a beak quite similar to that of chelydrids. Most testudinids (but not Malayemys) have the temporal bar deeply emarginate from below; the chelydrids have this bar at most shallowly emarginate. Chelydrids have the nostril, orbits, and otic opening somewhat smaller than is frequently the case in testudinids. The supraoccipital crest is higher or more steeply arched, the premaxillary pit is usually deeper, and in staurotypines and kinosternines there is a more marked festooning of the contours of the upper jaw than occurs in testudinids. The frontals are always small and excluded from the orbit in chelydrids; this feature is variable in testudinids.

A combination of most of the characters mentioned defines a chely- drid skull. The absence of all but one or two, most often of all, defines a testudinid as contrasted with a chelydrid skull.

Within the Chelydridae determination of chelydrine skulls from staurotypine and kinosternine skulls is at once possible on one key character which offers no difficulty. All chelydrines have the temporal region more fully covered than do any of the more advanced genera.

Discrimination of staurotypine as against kinosternine skulls is more difficult on the basis of any characters which have the smallest proba- bility of holding good if more genera are discovered. Perhaps the premaxillary beak is always more strongly developed in staurotypines and the temporal bar narrower vertically in the same group.

This difficulty in finding differences in the skulls of these two groups is akin to the difficulty in distinguishing dorsal shells. The shells of both subfamilies differ from those of chelydrines in the loss of one pair

8 BREVIORA NO. 2

of marginal scutes and one pair of peripheral plates. Shape and height of the shells and scute shape are essentially the same. Keeling is variable. Only in the plastron is a key difference at once apparent in the absence of an entoplastron. The mobility of anterior and posterior plastral lobes in kinosternines is another differentiating feature.

In which of these groups does the fragmentary Oligocene skull find its natural place?

It is clearly chelydrid sensu lato. It has the sharp premaxillary beak and deep premaxillary pit of a chelydrid. The temporal bar is broken but there is no suggestion of ventral emargination. The nostrils and the orbits are quite small. The supraoccipital crest is only partially preserved but its root gives evidence of a high arch as in typical chelydrids.

The skull is, however, clearly not chelydrine, since the temporal region is fully exposed by posterior emargination as in the two ad- vanced subfamilies, not as in chelydrines. But the postorbital bar is wider than in any staurotypine or kinosternine; this may be a primitive feature.

Is it staurotypine or kinosternine?

It has resemblances to both groups. The premaxillary beak is very strong as in staurotypines, but the temporal bar is very stout in vertical depth as in kinosternines. The pterygoid waist is moderately broad as in kinosternines, not very broad as in Stawrotypus or very narrow as in Claudius. The nostril is very small as in kinosternines and in contrast to the condition in staurotypines. Foramina incisiva are lacking as in staurotypines.

Some features, however, are extremely specialized. The degree of development of the secondary palate is greater than in any presently recognized chelydrid species, significantly greater than in Stawrotypus. Quite unique (unique in the order) is the extreme obliquity and dorsal arching of the secondary palate.

All in all, the skull seems more specialized than that of any living staurotypine, but at the same time more primitive in at least one respect (the strong premaxillary beak) than any living kinosternine, and perhaps more primitive in the width of the postorbital bar than either modern staurotypines or modern kinosternines.

The skull is Oligocene in age. The only known shell to which it might belong is Xenochelys, of the same age and not distant in locality. But the shell of Xenochelys is quite primitive for its group. Can so specialized a skull be assigned with any probability of correctness to so primitive a shell?

1952 A STAUROTYPINE SKULL g

Such an association is by no means impossible. Indeed, in the Princeton skull a few features like the very strong premaxillary beak and the wide postorbital bar may point to a stage of differentiation not very different from that of the Xenochelys shell: advanced in some respects, primitive in others. For the present it seems expedient to refer the Princeton skull with doubt to Xenochelys formosa Hay.

THE GEOLOGIC RANGE OF THE CHELYDRIDAE

The Princeton skull and the American Museum shell of Xenochelys formosa are the oldest known representatives of the Chelydridae (Chadronian Oligocene). An older fossil from the Eocene of Tunis was indeed referred to the family by De Stefano (1903), but his de- scription was based on a few bone fragments associated with the mold of three pleurals. The generic and species names, Gafsachelys phospha- tica, erected on this very insufficient basis may be disregarded as a nomen vanum and need no longer be considered as part of the fossil history of the Chelydridae.

It is, of course, surprising that the oldest representatives of the family should be staurotypine rather than chelydrine. It is, however, possible that future more complete knowledge may connect some of the other forms called dermatemydine by Hay (1908) with the Chely- dridae. The type species of Hoplochelys Pay was first called Chelydra crassa by Cope, and this genus, though possessed of a full complement of marginals and peripherals was regarded as possibly related to Staurotypus by Hay himself. Both this genus (Paleocene of North America) and Baptemys (Lower and Middle Eocene of North America) have the plastron considerably reduced, the bridge narrowed and the posterior lobe pointed, and an arrangement of plastral scutes like that of Recent Chelydra (pectorals meeting femorals). The shape of the shell in both genera is quite like that of staurotypinesand kinosternines, and in Hoplochelys the shell is tricarinate, as it is also in one species of Baptemys. The humerus of Baptemys tricarinata is very like that of Chelydra. However, the first vertebral is never elongate as in stauro- typines and kinosternines. The other vertebrals are never as wide as in chelydrines, though wider in later (Torrejon) than in earlier (Puerco) Hoplochelys. The costiform process of the nuchal is said to be short in Baptemys (Hay, 1908). The skull of Baptemys wyomingensis is known and is quite un-chelydrid in its major characters: the temporal bar is deeply emarginate from below, and the temporal region widely open above, there being a much greater caudal emargination than in

10 BREVIORA NO. 2

chelydrines. The postorbital bar is in consequence relatively narrow. The orbit also is rather large.

None of the conditions just mentioned in which Baptemys and Hoplochelys differ from chelydrids positively debars them from ancestry. For the present, however, and until they are better known and tran- sitional forms are discovered, it seems convenient to retain them in the Dermatemydidae, merely calling attention to their possible special relation to the Chelydridae.

Unless these forms, perfectly suitable in age, are ancestral chelydrids, there is no record of the family until the early Oligocene, and it is then first represented, as the shell and the referred skull of NXenochelys formosa show, by an advanced subfamily.

The first occurrence of apparent Chelydrinae is in Europe and later in the Oligocene. Fragments of doubtful value from the middle Oligocene of Germany have been assigned to “Chelydra sp.” by Reinach (1900), and in the later Oligocene of Germany rather good remains are found of an undoubted chelydrine, “‘Chelydra” decheni v. Meyer. Reference of the latter form to the Recent genus Chelydra is, as Zangerl (1945) has already pointed out, extremely doubtful: though the shell shape is that of a chelydrine, there are curious resemblances to Staurotypus and to Macroclemys rather than to Chelydra. Indeed, H. v. Meyer himself in 1852 expressly admitted that the inclusion of this form in Chelydra depended upon a very wide generic concept, and his idea of the genus was very definitely much wider than that current today.

In the Miocene of both Europe and North America there is a flowering out real or apparent of chelydrine types. In Europe “Chelydra’”’ murchisoni Bell and four other named species of ‘‘Chelydra”’ and Chelydropsis carinata Peters record the rather widespread oc- currence of the subfamily. The remarks above for ‘‘C.”’ decheni apply also to “C.” murchisoni and the other European forms referred to “Chelydra.”’

The carapace of Chelydropsis carinata has been excellently figured by Peters (1869). Unfortunately it does not seem possible to verify in his figure the features upon which Peters relied in distinguishing this form generically. I am unable to interpret the plate as showing the presence of supramarginals, and while a division of the nuchal bone into two parts is clearly shown, I (as also Boulenger in 1889) doubt that this reflects the normal condition of the animal. Never- theless, I consider it probable, if only on the grounds of zoogeography and age, that the genus will stand, though requiring redefinition. It

1952 A STAUROTYPINE SKULL 11

may at least be pointed out that if the eventual much needed restudy of the European chelydrines should reveal that they all belong to one genus, the name Chelydropsis is available.

From the Miocene of North America three chelydrines have been de- scribed. Two are known from skulls only: Chelydrops stricta Matthew and Macroclemys schmidti Zangerl. Both of these are from Nebraska, M. schmidti from the Middle Miocene, Chelydrops stricta from the Upper Miocene. Both are certainly related to Recent M. temminckit, but they are distinct from that form and from one another. Chelydrops is unique among known chelydrids in having a ridged alveolar surface of the maxilla.!. M. schmidti differs from Chelydrops and from M. tem- minckw by the considerably shorter antorbital portion of the face.

The other Miocene North American form (from the Roslyn Miocene of Washington) is known from the carapace only, no portion of the plastron nor any skeletal parts having been recovered. Hay described this form as a new genus and species, Acherontemys heckmani, because of the close articulation of pleurals and peripherals and because the vertebrals were even broader than in living chelydrines. This shell may belong to either or neither of the forms represented by the skulls before mentioned.

Zangerl (1945) has described a skull fragment from the Pliocene (Clarendonian) of South Dakota, which is indistinguishable from Recent M. temminckvi. Gilmore (1923) has described from the San Pedro Valley of Arizona, either Pliocene or Pleistocene, a Kinosternon which is said to differ from Recent K. flavescens mostly in size.

Two species of Chelydra and one of Macroclemys have been described by Hay from the Pleistocene of Florida. The value of these forms, based on fragmentary material, will be difficult to determine. There are also scattered Pleistocene records for the Recent species Chelydra serpentina and Macroclemys temminckit.

Acknowledgments. I am indebted to Dr. Glenn L. Jepsen for the privilege of studying and describing the Princeton skull. Mr. Arthur Loveridge and Mr. C. M. Bogert have generously made available the comparative Recent material under their care in the Museum of Comparative Zoology and the American Museum of Natural History respectively. Dr. E. H. Colbert permitted examination of the types of Xenochelys formosa and Chelydrops stricta. Dr. A. S. Romer and Mr. L. I. Price have read the manuscript. Mr. Sam McDowell is to be credited with the drawings and a number of useful suggestions.

1 The type and figured adult skull fragment has this ridge. The young specimen referred by Matthew to this form lacks the ridge.

12 BREVIORA NO. 2

TABLE 1, FOSSIL SPECIES OF THE CHELYDRIDAE

OLIGOCENE: Xenochelys formosa Hay Chadronian Oligocene (S. Dakota) North America “Chelydra” decheni v. Meyer Upper Oligocene (Siebengebirge) Europe

MIOCENE: “Chelydra”’ murchisoni Bell Miocene (Oeningen) Europe Chelydropsis carinata Peters Miocene (Eibiswald) Europe “Chelydra”’ meilheuratiae Pomel Miocene (Allier) Europe “Chelydra” lorettana (v. Meyer) Glaessner Miocene (Leithagebirge) Europe “Chelydra” argillarum Laube Miocene (Preschen) Europe “Chelydra” allinghensis E. Fuchs Miocene (Viehhausen) Europe Macroclemys schmidti Zanger] Middle Miocene (Nebraska) North America Chelydrops stricta Matthew Upper Miocene (Nebraska) North America Acherontemys heckmani Hay Miocene (Washington) North America and additional European records for ‘‘Chelydra sp.”” and “Macroclemys sp.”

PLIOCENE: Macroclemys temminckii (Holbrook) Zanger! Early Pliocene (S. Dakota) North America

PLEISTOCENE:

Macroclemys floridana Hay

Pleistocene (Florida) North America Chelydra laticarinata Hay

Pleistocene (Florida) North America Chelydra sculpta Hay

Pleistocene (Florida) North America Kinosternon arizonense Gilmore

Pleistocene (Arizona) North America and additional North American records for M. temminckii and

C. serpentina

1952 A STAUROTYPINE SKULL 13

LITERATURE CITED BouLENGER, G. A. 1889. Catalogue of the chelonians, rhynchocephalians and crocodiles in the British Museum (Natural History). London. 311 pp.

Fucus, ErtKa 1938. Die Schildkrétenreste aus dem oberpfalzer Braunkohlentertiar. Palaeontographica, Abt. A, Vol. 89, pp. 57-104.

GILMoRE, C. W. 1923. A new fossil turtle, Kinosternon arizonense. Proc. U.S. Nat. Mus., Vol. 62, pp. 1-8.

GLAESSNER, M. F. 1933. Die Tertiirschildkréten Niederdsterreichs. Neues Jahrb. Min. Geol. Pal., Abt. B, Vol. 69, pp. 353-387.

Hay, iO; PF: 1908. Fossil turtles of North America. Carnegie Institution of Washing- ton Publication No. 75. 568 pp. 1916. Description of some Floridian fossil vertebrates belonging mostly to the Pleistocene. Rept. Florida Geol. Survey, Vol. 8, pp. 41-76.

Horrman, C. K. 1890. Schildkréten in Bronn’s Klassen und Ordnungen des Tierreichs. Leipzig. 442 pp.

LauBg, G. C. 1900. Neue Schildkréten und Fische aus der bohm Braunkohlenformation: Abhandl. deutsch. naturwiss.-med. Ver. B6hmen ‘‘Lotos’’, Vol. 2, no. 2, pp. 37-56.

Martruew, W. D. 1924. Third contribution to the Snake Creek fauna. Bull. Amer. Mus. Nat. Hist., Vol. 50, pp. 59-210.

Meyer, H. von 1852. Ueber Chelydra Murchisoni und Chelydra Decheni. Palaeonto- graphica, Vol. 2, pp. 237-247.

Peters, K. F. 1855. Schildkrétenreste aus den 6sterreichischen Tertiér-ablagerungen. Denkschr. math-naturwiss. K]. Akad. Wiss. Wien, Abt. 2, Vol. 9, pp. 1-22. 1869. Zur Kenntniss der Wirbelthiere aus den Miociinschichten von Eibiswald in Steiermark. J. Die Schildkrétenreste: Denkschr. math-naturwiss. K]. Akad. Wiss. Wien, Vol. 29, pp. 111-124.

14 BREVIORA NO. 2

PomEL, A. 1854. Catalogue méthodique et descriptif des vertébrés fossiles décou- verts dans le bassin hydrographique supérieur de la Loire et surtout dans la vallée de son affluent principal l’Allier. Paris. 193 pp.

REINACH, A. VON 1900. Schildkrétenreste in Mainzer Tertiirbecken und in benachbarten ungefahr gleichalterigen Ablagerungen. Abhandl. Senckenberg. naturf. Ges., Vol. 24, pp. 3-135.

SIEBENROCK, F.

1897. Das Kopfskelet der Schildkréten. Sitz.-Ber. Akad. Wiss. Wien, Abt. 1, Vol. 106, pp. 245-328.

1907. Die Schildkréten Familie Cinosternidae m. monographisch bearbeitet. Sitz.-Ber. Akad. Wiss. Wien, Abt. 1, Vol. 116, pp. 527-599.

1909. Synopsis der rezenten Schildkréten mit Berucksichtigung der in historischer Zeit ausgestorbenen Arten. Zool. Jahrb., Suppl., Vol. 10, pp. 427-618.

STEFANO, G. DE 1903. Nuovi Rettili degli strati a fosfato della Tunisia. Bol. Soc. Ital. Ecol., Vol. 22, pp. 51-80.

TEppPNER, W. 1915. Ein Chelydra-Rest von Goriach. Mitt. naturwiss. Ver. Steiermark, Vol. 51, pp. 474-475.

WIuuiaMs, E. E. 1950. Variation and selection in the cervical central articulations of living turtles. Bull. Amer. Mus. Nat. Hist., Vol. 94, pp. 511-561.

ZANGERL, R. 1945. Fossil specimens of Macrochelys from the Tertiary of the plains. Fieldiana, Geol. Ser., Vol. 10, pp. 5-12.

ZDANSKY, O. 1924. Ueber die Temporalregion des Schildkrétenschadels. Bull. Geol. Inst. Univ. Upsala, Vol. 19, pp. 89-114.

1952 A STAUROTYPINE SKULL 15

PLATE 1

Staurotypus salvinit M.C.Z. 4989: Dorsal, ventral, anterior and lateral views of skull. X about 114.

16 BREVIORA NO. 2

TES +

Lor*5

PLATE 2

Xenochelys formosa Princeton 13686: Dorsal, ventral, anterior and lateral views of referred skull fragment. About natural size.

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. May 23, 1952 NUMBER 3

NOTES ON SIPHONOPHORES 3. Nectopyramis spinosa n. sp.

By MARY SEARS!

In a number of samples collected by the ‘‘Dana”’ in tows with 1000 to 3000 meters of wire out, there were several nectophores and bracts with rows of peculiar jagged spines (Fig. C). These spines are quite unlike those on other deep-dwelling species, such as V ogtia pentacantha Kolliker, V’. spinosa Keferstein and Ehlers, and Athorybia rosacea Forskal. In fact, the specimens have little in common with either the Hippopodidae or the Anthophysidae. However, one nectophore was described by Moser (1925, p. 425, Pl. 25, figs. 5-7) as Hippopodius (?) cuspitata, although it actually closely resembled the specimens found in the “Dana” collection. The chief difference is that hers had the peculiar jagged spines scattered irregularly over its surface, whereas on the ‘‘Dana” specimens these are arranged in discrete rows. Never- theless, the spines themselves seem to be identical in structure. Like the ‘‘Dana’’ nectophores, most of the characteristics described and figured by Moser (1925) make it appear almost certain that her speci- men was also a Nectopyramis, not a Hippopodius. In all examples, the shape of the nectosac, its relation to the hydroecium, and the fact that the nectophores have all been taken singly and at considerable depths are all peculiarities of Nectopyramis. On the other hand, the soma- tocyst is tubular and not “represented by a series of divergent canals”’ (Bigelow, 1911a, p. 338) as defined in the brief description of the genus.

The original account was based on one species, Nectopyramis thetis. Soon thereafter a second species, Nectopyramis diomedeae, with many of the same characteristics was recorded (Bigelow, 1911b). Since then no additional species have been reported, and little more has been

1 Papers from the ‘‘Dana”’ Collection No. 38, and Contribution No. 607 of the Woods Hole Oceanographic Institution.

2 BREVIORA NO. 3

learned about the genus. Hence, there has been little reason for modifying the original description. Nevertheless, our ideas concerning the relationships of this genus have changed considerably. Thus, the family of the Monophyidae has been broken up and the individual genera transferred to other well-established families (Totton, 1932, p. 328) with which they had obvious affinities. Nectopyramis, there- fore, is now referred to the Prayidae. In this family, the somatocyst and radial canals are structurally quite varied; in some, they are simple, in others branched. At first, these were used as a criterion for separating genera (Bigelow, 1911b), but more recently, a transitional form with branched radial canals and a simple somatocyst has been

GG PASLEY DEL a ¢ \ AAA AaAeng |

r ‘4 Cc = - 1 eae Aaah ae f Sal ate A wel tA) = a 4 A Ir \h .

r fj y \ i U r 2 f fe i] s 4 |

1 Ae} +,

of ¥!

iY ney mr, if y 7 fi 4 4 > eer ° .7¢ Si M4 ay 4

&

as,

discovered (Leloup, 1934, p. 11). This suggests that when more is known about this family, differences in the structure of somatocyst and canals may prove to be of specific rather than generic value.

If, then, we consider that the shape of the somatocyst is in all probability a specific character in Nectopyramis, as may be the case among other prayids, we have two species originally described by Bigelow (1911la; 1911b), N. thetis and N. diomedeae, both of which had branched somatocysts, but simple radial canals and two species, N. cuspitata Moser and N. spinosa n. sp. (described below) with simple tubular somatocysts. The two latter are both spiny, but the two are readily distinguished for Moser’s species is irregularly covered by the spines, but in spinosa these are arranged in regular rows along more or less distinct ridges.

1952 NOTES ON SIPHONOPHORES 3

NECTOPYRAMIS SPINOSA N. sp.

The type specimen is a nectophore about 5 mm. in length taken at “Dana” St. 3933" at 11°18’S., 50°13’E. on 20 December, 1929, in a stramin net 150 cm. in diameter towing with 2000 meters of wire out. The type specimen will be deposited in the University Museum, Copenhagen, Denmark.

Other specimens were taken as follows:

St. 3556". 2°52’N., 87°38’W. 14. IX. 1928. S150. 2000 m. wire. 1 nectophore.

Stabi(.4 15 28's5 160739 B.. 4230 TIT... 1929) S150. 2000 m. wire. 2 bracts.

St. 3920"". 1°12’N., 62°19’E. 9. XII. 1929. S150. 1000 m. wire. 1 bract.

St, 3964")., 25:19'S., 36:13 BE.) 15. 11. 1930. E300. 3000 m. wire. 1 bract.

St. 3998. 7°34’S., 8°48’W. 1. III., 1930. S150. 1000 m. wire. 1 bract.

Nectophore (Fig. A). The unique rows of jagged spines make it obvious that the nectophore is new. Although probably flattened on preservation, in dorsal or ventral view, it is roughly triangular much as are the nectophores of the other known species of Nectopyramis. The nectosac opens into the base of the triangle and like N. thetis “is comparatively small, shallow, and saucer-shaped” (Bigelow, 1911a, p. 338) with four primary radial canals along its sides. It is uncertain whether or not these eventually branch because of preservation. A large deep hydroecium lies just above it with a slit-like opening on the ventral surface. This extends the entire length of the hydroecium. The somatocyst insofar as can be determined appears to be a tubular rod extending from the base of the hydroecium to slightly above its apex. The stem and appendages are entirely missing.

Bract (Fig. B). Although bracts have not been found attached to the nectophore, there seems to be little doubt that the bracts with rows of the same jagged spines belong to this species. These are globular, about 2.5 mm. high and slightly more than 3 mm. wide. As in the nectophore, the hydroecium is so deep that it extends somewhat into the upper half of the bract. Closely associated with the hydroe- cium is a long tubular somatocyst which follows the outline of the bydroecium. On two of the specimens, two threadlike branches are given off dorsally and appear distally to have a globular connection.

4 BREVIORA No. 3

BIBLIOGRAPHY BicELow, H. B.

191la. Biscayan plankton collected during a cruise of H.M.S. ‘“‘Research”’ 1900. Pt. XIII. The Siphonophora. Trans. Linn. Soc., London, Zool., 2nd ser., 10(10): 337-357, pl. 28.

1911b. XXIII. The Siphonophorae. Reports on the Scientific Results of the Expedition to the Eastern Tropical Pacific, 1904-1905... . Mem. Mus. Comp. Zool. 38: 171-402, 32 pls.

1931. Siphonophorae from the Arcturus Oceanographic Expedition. Zoologica. N. Y., 8(1): 525-592, figs. 185-220.

BicELow, H. B., and M. Spars 1937. H. 2. Siphonophorae. Rept. Danish Ocean. Exped., 1908-10, to the Mediterranean and Adj. Seas, Biol. 2: 1-144, 83 text figs.

LELoup, E.

1932. L’eudoxie d’un siphonophore calycophoride rare, le Nectopyramis thetis Bigelow. Bull. Mus. Hist. Nat. Belg. 8(3): 1-8, 5 text figs.

1933. Siphonophores calycophorides provenant des campagnes du Prince Albert I de Monaco. Rés. Camp. Sci., Monaco, 87: 1-64, 1 double plate.

1934. Siphonophores calycophorides de l’Océan Atlantique tropical et austral. Bull. Mus. Hist. Nat. Belg. 10(6): 1-87, 15 figs.

1936. Siphonophores calycophorides (suite) et physophorides provenant des campagnes du Prince Albert ]* de Monaco. Rés. Camp. Sci., Monaco, 93: 1-35, 2 double plates.

Le.oup, E., and E. HENTscHEL 1935. Die Verbreitung der calycophoren Siphonophoren im Siidatlan- tischen Ozean. Wiss. Ergeb. Deutschen Atlantischen Exped... . “Meteor” 1925-1927, 12(2): 1-31, 17 text figs.

Moser, F. 1925. Die Siphonophoren der Deutschen Siidpolar-Expedition. Deut- schen Siidpolar Exped. 17, Zool. 9: 1-541, 33 pls., 61 text figs.

Totton, A. K. 1932. Siphonophora. Great Barrier Reef Exped., 1928-29, Brit. Mus. (N. H.), Sci. Repts. 4(10): 317-374, 36 text figs. 1936. Plankton of the Bermuda Oceanographic Expeditions. VII. Siphonophora taken during the year 1931. Zoologica, N. Y., 21(4): 231-240.

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. JUNE 9, 1952 NuMBER 4

A UNIQUE REMOPLEURIDID TRILOBITE

By H. B. Wuirrincton

INTRODUCTION

The species described below, and made the type of a new genus, has been known for more than 75 years. Previous illustrations are both few and inadequate, and no attempt had been made to reconstruct the exoskeleton. When such an attempt is made (text-fig. 1) this unique remopleuridid is shown to have been a most unusual trilobite, in which the long, forked hypostome reached back to the anterior margin of the pygidium. It could not enroll, as it would seem that many (or most) trilobites could. If the hypostome was rigidly attached to the cephalic doublure, then only very restricted movements of the thorax and pygidium, relative to the cephalon, were possible.

I am indebted to Dr. G. Arthur Cooper, U. S. National Museum, Washington, D. C., for allowing me to study and photograph material in his charge.

SYSTEMATIC DESCRIPTION Family REMOPLEURIDIDAE Hawle and Corda, 1847

Genus HYPODICRANOTUS gen. nov.

Type species. Remopleurides striatulus Walcott, 1875, pp. 347-349, fig. 27, from C. D. Walcott’s locality in the Trenton limestone of Trenton Falls, Oneida Co., New York.

Discussion. The cephalon of Hypodicranotus striatulus differs from that of the type species of Remopleurides (Whittington, 1950, pp. 540-

2 BREVIORA NO. 4

543, Pl. 70, figs. 1, 2, 4, 5) in that: (a) the anterior part of the glabella, the tongue, projects for a greater distance in front of the eyes; (b) three pairs of glabellar furrows are present, and the second is relatively farther back than that of Remopleurides; (ec) the broad spine that curves back beside the thorax is not the genal spine, but a lateral cephalic spine; (d) the hypostome is long and deeply forked, not a

Text-figure 1. Hypodicranotus striatulus (Walcott, 1875). Reconstructions of ventral (left) and right lateral (right) appearance of outline of exoskeleton, approx. X 3, based upon M. C. Z. Nos. 1616, 1617. The positions of the raised lines on the cephalic and thoracic doublure and the hypostome are indicated in the ventral view. m = median suture; p = pit in doublure. In both draw- ings a general outline of the pygidium is indicated by dotted lines.

sub-rectangular plate wider than long. The thorax of Hypodicranotus probably consists of eleven segments, and is like that of Remopleurides, with the wide axis, prominent articulating processes and sockets, diagonal pleural furrows, and backwardly curved and pointed pleurae.

1952 UNIQUE REMOPLEURIDID TRILOBITE 33

The axial and pleural furrows in //ypodicranotus are shallow, there are no axial spines, and no segment has the pleural spines exceptionally elongated. The pygidium of Hypodicranotus is poorly known, but appears to be rectangular in outline, longer than wide, with the axis much shorter than that of Remopleurides. The dorsal exoskeletons of other remopleuridid genera may readily be distinguished from that of Hypodicranotus or Remopleurides.

Hypodicranotus is known from the Trenton Group of New York and Ontario, from rocks of the same age in Wisconsin, the Prosser limestone of Minnesota, the Kimmswick limestone of Missouri and Illinois, the Viola limestone of Oklahoma, and from the Ordovician strata at Silliman’s Fossil Mount, Baffin Island.

HypopIcRANOTUS STRIATULUS (Walcott, 1875) Plate 1, figs. 1-10; text-figure 1.

Lectotype (selected Raymond, 1925, explanation of Plate 3). Mus. Comp. Zool. No. 1616, original of Walcott, 1875, p. 347, fig. 27A, from the dark-gray colored Trenton limestone of Trenton Falls, Oneida County, New York. The large collection from this locality in the Museum of Comparative Zoology includes the trilobites Ceraurus pleurexanthemus Green, 1832; Calliops callicephalus (Hall, 1847); Leonaspis ? trentonensis (Hall, 1847); Diacanthaspis ? parvula (Walcott, 1879); Ilaenus cf. americanus Billings, 1859; [sotelus gigas DeKay, 1824; Flevicalymene senaria (Conrad, 1841), as well as bryozoans, brachiopods, crinoids and asteroids. This locality, from which both W. P. Rust and Walcott collected, is probably in the Denmark member of the Sherman Fall formation of Kay (1937, pp. 267-268).

Paratypes. Mus. Comp. Zool. Nos. 1617, 1618, 1537, same locality and horizon. Additional material, cranidia and hypostomes, is included under Mus. Comp. Zool. Nos. 3267-3269. No. 1617 is original of Walcott, 1875, p. 347, fig. 27a.

Description. Dimensions of lectotype, Mus. Comp. Zool. No. 1616, in millimetres:

Cephalon:

WWemeule Ga Clbtall) ie cape an. Sock. ye cust areas a ee tA fos igicy: 10.4

emma (CXS OTOL ree cena mac. Nec ett teen eye ds eect she ay ee 18.2 Wircthecttsceneall ingles en eee tye teres eo gh 16.4 height atsecond elabellar furrow... 2. ~.2. 6. sete oe oe 6.5 Maximuniswidth across palpebral lobes .......-:...-...5.- Ieee

WEN HME OMEN CU GWC neh. nhs Cos cto noae mae aac ree se P ez

4 BREVIORA NO. 4

Thorax: Widthvaxis first Seamentees ecu eee ae ory eo oe eee 10.2 : ‘Smumith)> “> approxi ee ee eee ee 4.3 “‘ first segment (across tips of pleurae)................13.6 ‘* seventh segment (across tips of pleurae)............ Tet Length (sagittal) complete thorax, approx................. 13.0

Dimensions of hypostome, Mus. Comp. Zool. No. 1617, in milli- inetres:

bength: (Saritbtall ic meh ets doko ose heed het a oe eee OZ

Re (ExSAriblalmn utp hue a1. oo eae eee ae ee 20.2 Maximum width (at about half the length)............... 7.4 Widthsofantadledbodyss... ee. pei ioe ee eee 3.7

The cranidium includes the occipital ring, glabella and tongue, and palpebral lobes. The occipital ring is longitudinally flat, trans- versely moderately convex, bounded laterally by the posterior branches of the facial sutures, which are straight but run diagonally back from the posterior corner of the eye lobe, then curve to run into the posterior margin at right angles. Inside and parallel to the suture is a faint depression, the axial furrow, and just inside the point where the suture cuts the posterior margin is a rounded notch, the articulating socket. The occipital furrow is shallow. Between the eye lobes the glabella and palpebral lobes are gently and evenly convex both longitudinally and transversely. In the mid-line the tongue of the glabella of the lectotype projects forwards 1.8 mm. in front of the eye lobe, and is convex and bent down so that the anterior part overhangs the sutural margin. There are three pairs of glabellar furrows, visible because they are both unornamented and slightly depressed (PI. 1, figs. 2, 4). Each furrow runs in a curve convex forwards, and they are equally spaced from each other, the distance (exs.) between them slightly less than the equal distance (exs.) of the outer end of the first from a point opposite the posterior end of the eye lobe, and the outer end of the third from a point opposite the anterior end of the eye lobe. The furrows commence a short distance inside the palpebral furrow, and do not extend to the mid-line. The first is the most strongly convex, the second commences farthest out, and the third farthest in. The palpebral rim is flat, depressed slightly below the lobe, outwardly sloping, widest posteriorly, narrowing forwards and disappearing at the anterior end of the eye lobe. Posteriorly the palpebral furrow runs into the junction of the extremity of the occipital furrow and the axial furrow. The eye lobe is long, in dorsal aspect curved, most strongly in the posterior part. The eye surface is gently convex

1952 UNIQUE REMOPLEURIDID TRILOBITE 5

transversely, and slopes steeply. The arrangement of the minute, convex facets is shown in Plate 1, figure 10. The outer margin of the eye lobe is defined by a narrow, convex border which commences at the posterior corner and runs forwards to merge anteriorly with the border of the cheek. The anterior branches of the facial suture curve to run at first forwards and inwards from the outer, anterior, corner of the eye lobe, and then run inwards and upwards to the mid-line. Thus in anterior aspect (PI. 1, fig. 5) the margin of the tongue is a curve convex upwards. The free cheek is triangular in outline, outwardly sloping, widest behind the eye lobe, the genal spine short and pointed. The cheek narrows forwards to disappear opposite the anterior part of the eye lobe. From the lateral border opposite the median part of the eye lobe a broad spine curves back, narrowing and reaching to a point opposite the axis of the fifth thoracic segment. The inner margin of the proximal part of this spine is parallel to, and just outside of, the outer margin of the cheek and genal spine. The doublure of the cephalon is broad anteriorly, gently convex, crossed by a median suture. Antero-laterally and laterally the doublure is narrower and bent upwards. A small pit is situated on the exsagittal line passing through the anterior end of the eye lobe, and about mid-way across the doublure (PI. 1, fig. 8; text-fig. 1). The hypostome is long and forked, and its position relative to the rest of the exoskeleton is shown in text-figure 1. It is gently convex, in both longitudinal and transverse directions, the transverse convexity greatest at the curved anterior margin. The convexity (in ventral view) is interrupted only by the shallow depression in the median region just in front of the crotch of the fork. The middle body is circular in outline, faintly elevated, defined postero-laterally by a distinct furrow. It is divided into three sub-equal sections by shallow radial depressions, the strong- est leading back to the crotch of the fork, the other two directed antero-laterally (Pl. 1, fig. 7). On the inside of the exoskeleton of a smaller specimen (PI. 1, fig. 9) these sections appear more pronounced, and on both sides the postero-lateral pair are the more prominent. Anteriorly the middle body merges with the border, and the anterior wings appear to be short, triangular, and upwardly directed. Each postero-lateral border is extended as a prong of the fork, the inner margin of the prong straight, the outer margin curved, so that the maximum width of the hypostome is in front of half the length, and the prong tapers to a sharp point. The doublure of the hypostome (Pl. 1, figs. 4, 9) is narrow along the lateral border, but extends in under the fork to a line almost under the margin of the middle body.

6 BREVIORA NO. 4

The inner edge of the doublure is flexed up sharply. Mus. Comp. Zool. No. 1618 (PI. 1, fig. 4) shows that the doublure of the prong is convex dorsally, with a low, median, longitudinal ridge.

The thorax is moderately convex transversely, the axis broad, narrowing rapidly backwards. It seems to consist of eleven segments. On the left side of the lectotype 10 segments can clearly be seen (Pl. 1, fig. 2). Behind this only what may be part of the left pleura of the eleventh segment, and part of the pygidium, are preserved. Another specimen (PI. 1, fig. 6) shows ten partly disarticulated seg- ments and an incomplete pygidium The last segment has the pleura pointing almost directly posteriorly, whereas that next in front has the pleura directed back and slightly outwards, and the lateral margins are curved. This penultimate segment is very like the tenth of the lectotype. The axial ring is moderately convex transversely, the articulating furrow shallow, the articulating half-ring of length (sag. = sagittal) about half that of the ring (PI. 1, fig. 6). There is no axial furrow as such, but inside the articulating sockets and processes a triangular area (broadest anteriorly) of the outermost part of each ring is slightly depressed. The inner margin of this area is a diagonal curved line sub-parallel to the pleural furrow. The articulating process, on the anterior margin of the segment, is slightly raised as well as forwardly projecting. Postero-laterally the inflation is con- tinued across the pleura as a low, curved ridge, dying out at about two-thirds (exs. = exsagittal) the length. This ridge, and a depression on the inner side, define the pleural furrow. The pleurae are gently convex (tr. and exs.), outwardly sloping, the narrow (tr. = transverse) inner part directed transversely, the outer part curved back and pointed. The shape and inclination of the pleurae are shown in Plate 1, figures 1, 2, 6.

Only a small part of the pleural lobes of the pygidium is preserved in the lectotype. The only other specimen known (PI. 1, fig. 6) is also incomplete. The outline was evidently rectangular, longer than wide, the ill-defined axis convex, short, and wide. Behind and beside the axis the pleural lobes slope down in a curve concave in longitudinal profile. The lateral and posterior margins, and the doublure, are unknown.

Scattered over the cranidium, but largest and most closely spaced on, and adjacent to, the palpebral lobes (though absent from the palpebral rim), are small crescentic depressions, the points of the crescent facing forwards, and the concave, anterior margin raised (Pl. 1, figs. 1, 2, 4). Similar but larger structures are present on the

1952 UNIQUE REMOPLEURIDID TRILOBITE

free cheeks, the genal spine, and the base and outer part of the long lateral spine (Pl. 1, fig. 1). There is a tiny median tubercle on the occipital ring, closer to the furrow than the posterior margin. Faint, small crescentic depressions are scattered on the median part of the axis of the thorax. Strong, well-spaced, raised lines run approximately longitudinally on the inner part of the lateral cephalic spine, the thoracic pleurae, and on the doublure of these areas and the cheeks (Pl. 1, figs. 1-4, 6, 8). The lines tend to run in curves convex inwerds on the dorsal surface of the pleurae, and transversely in curves convex forwards on the dorsal surface of the pygidium. On the inner part of the doublure of the thoracic pleurae, however, they run in curves concave inwards. These lines die out on the median part of the cephalic doublure, and are replaced by faint, anastomosing, transverse grooves. On the lateral borders and fork of the hypostome (PI. 1, fig. 7) the longitudinal lines are strongest, and that which runs just inside the inner margin of the prongs is a prominent ridge. The middle body shows a faint pattern of narrow, anastomosing ridges running longitudinally on the postero-lateral sectors. On the anterior sector of the middle body and the anterior border, fine, well-spaced, anasto- mosing grooves, like those on the median part of the cephalic doublure, run transversely.

Discussion. This species was first described by Walcott (1875, pp. 347-349, fig. 27), and the original material later redescribed by Raymond (1925, pp. 57-58, PI. 3, figs. 4, 5). An incomplete cran- idium from the same general locality as the type (U.S. Nat. Mus. 92528) was also figured by Foerste (1920, p. 222, Pl. 22, figs. 18 A-C). Raymond (1921, p. 31, Pl. 9, fig. 7) described a hypostome from the Middle Trenton of Trenton, Ontario, and referred it to H. striatulus, and stated that a cranidium had also been found at Governor Bay, near Ottawa. Specimens kindly loaned to me by Professor G. Winston Sinclair, from the Middle Trenton of Lakefield, Ontario, contain cranidia and hypostomes like those of the type material. A second species of Hypodicranotus is H. missouriensis (Foerste, 1920, pp. 220-222, Plz 21, fig. 17; Pl. 22, figs. 17A, 17B; Bradley, 1930, pp. 246- 247, Pl. 30, figs. 4-9) from the Kimmswick limestone of Missouri and Illinois. The type material (U. S. Nat. Mus. No. 78438) includes cranidia and a hypostome. The outline of the latter, and the prom- inence of the postero-lateral areas of the middle body, distinguish it from H. striatulus. The characteristic hypostome of Hypodicranotus also occurs in the Trenton of Duck Creek Quarry, near Green Bay, Wisconsin (U.S. Nat. Mus. Nos. 72181, 87687), the Prosser limestone

Ss BREVIORA NO. 4

of St. Paul, Minnesota (U.S. Nat. Mus.), and I have collected one from about 100 ft. above the base of the Viola limestone, in the road cut on U.S. highway 77, in Carter County, 21 miles north of Springer, Oklahoma. The matrix at this locality was a finely-granular, light grey-brown limestone, which yielded abundant graptolites and the trilobites Cryptolithoides ulrichi Whittington, 1941, Trinodus_ sp., Robergia sp., and an asaphid. The specimen figured by Roy (1941, p. 155, fig. 114), as Remopleurides sp., from Silliman’s Fossil Mount, Baffin Island, seems also to be the hypostome of this genus The genus thus occurs in central and eastern North America in rocks of Trenton age, and the Baffin Island beds may be of a similar age.

The exoskeleton of Hypodicranotus striatulus, apart from the hypostome, is like that of other remopleuridids of Middle Ordovician age, e.g. the Irish specimens (Whittington, 1950, pp. 540-548, PI. 69, figs. 5-10, Pl. 70, figs. 1-6) and undescribed species from the Edinburg limestone of Virginia. The latter have a sub-rectangular hypostome, not forked, like that of the type species (Whittington, 1950, Pl. 70, fig. 2). The posterior margin reaches back to a point lying no farther back than the occipital furrow. The long, forked hypostome of Hypo- dicranotus is not only unique, so far as is known, among remopleuridids, but its relatively great length is in excess of that of any other trilobite known to me. It Is evident that here is one genus of trilobites that could not enroll.

The pit in the antero-lateral cephalic doublure (PI. 1, fig. 8, text- fig. 1) is a feature which I have observed in several different species of Middle Ordovician remopleuridids. Silicified specimens show the pit to be the opening of an upwardly-directed tube, which narrows inwards and terminates near the lower, anterior corner of the eye surface. The position and nature of this opening do not suggest that it is homologous with the Panderian opening, which, if present in the cephalon, is situated in the postero-lateral cephalic doublure. Not all remopleuridids appear to show this opening (e. g. those described by Ross, 1951, pp. 84-91, Pl. 20, do not), and I have not observed a similar opening in any other group of trilobites.

It is tempting to speculate on the mode of life of the holaspid Hypodicranotus. It occurs in company with shallow-water marine forms, many of which are indisputably benthonic. The long hypostome prevented enrollment, but did provide some protection for the ventral surface. If no movement was possible at the hypostomal suture, then the amount of possible movement of the thorax and pygidium in the vertical plane, relative to the cephalon, must have been severely

1952 UNIQUE REMOPLEURIDID TRILOBITE Q

limited. The mode of articulation of the thorax precludes any con- siderable movement relative to the cephalon in the horizontal plane. There is ample room for the appendages to project downwards and outwards between the hypostome and the thoracic pleurae. But we know nothing of the type of appendage possessed by remopleuridid trilobites, and without this information have little basis for speculation on the mode of locomotion, manner of feeding, ete. Was Hypodicran- otus a burrowing, crawling, floating, or swimming form? No definite answer is possible but Iam inclined to think of it as either floating or swimming.

REFERENCES Brab.ey, J. H. 1930. Fauna of the Kimmswick Limestone of Missouri and Illinois. Contrib. Walker Mus., vol. 2, no. 6, pp. 219-290, pls. 23-30.

Forrste, A. F. 1920. The Kimmswick and Plattin Limestones of Northeastern Missouri. Denison Univ. Bull., J. Sci. Lab., vol. 19, pp. 175-224, pls. 21-23.

Kay, G. M. 1937. Stratigraphy of the Trenton Group. Bull. Geol. Soc. Am., vol. 48, pp. 233-302, pls. 1-10.

RAYMOND, P. E. 1921. A Contribution to the Description of the Fauna of the Trenton Group. Geol. Surv. Canada, Mus. Bull. 31, Geol. Ser. 38, pp. 1-64, pls. 1-11. 1925. Some Trilobites of the Lower Middle Ordovician of Eastern North America. Bull. Mus. Comp. Zool., vol. 67, no. 1, pp. 1-180, pls. 1-10.

Ross, R. J. 1951. Stratigraphy of the Garden City Formation in Northeastern Utah, and Its Trilobite Faunas. Peabody Mus. Nat. Hist., Yale Univ., Bull. 6, pp. 1-161, pls. 1-36.

Ox 1S. KK 1941. The Upper Ordovician Fauna of Frobisher Bay, Baffin Land. Mem. Field Mus. Nat. Hist., Geol., vol. 2, pp. 1-212, 146 figs. Wa.tcort, C. D. 1875. New Species of Trilobite from the Trenton Limestone at Trenton Falls, N. Y. Cincinnati Quart. J. Sci., vol. 2, pp. 347-349, fig. 27. WHITTINGTON, H. B. 1950. Sixteen Ordovician Genotype Trilobites. J. Paleont., vol. 24, no. 5, pp. 531-565, pls. 68-75.

EXPLANATION OF PLATE I

Hypodicranotus striatulus (Walcott, 1875). Trenton limestone, probably Denmark member of the Sherman Fall formation, Trenton Falls, Oneida County, New York, al] X 3, except figure 10, X 17144, Mus. Comp. Zool. collections.

Figs. 1-3, 5, 8. Lectotype, No. 1616, left lateral, dorsal, right lateral, anterior, and antero-ventral views, presumed original of Walcott, 1875, p. 347, fig. 27A, and original of Raymond, 1925, PI. 3, fig. 4.

Figs. 4, 10. Paratype, No. 1618. 4, dorsal view. The right half of the dorsal exoskeleton of the cephalon and three thoracic segments are preserved; on the left is seen the lateral cephalic spine, and external moulds of the doublure of several pleurae. The left, and tip of the right, prongs of the fork of the hypostome are exposed from the inner side. The hypostome has been slightly displaced. 10, the anterior portion of the right eye surface, showing the arrangement of the facets.

Fig. 6. Paratype, No. 1537, dorsal view.

Fig. 7. Paratype, No. 1617, ventral view, presumed original of Walcott, 1875, p. 347, fig. 27a, and Raymond, 1925, Pl. 3, fig. 5.

Fig. 9. No. 3269, dorsal view of interior of hypostome.

BREVIORA Unique Remopceuripip TRicopite. PLATE 1

Se aaa

*

j |

Cee ae ian

7?

. &

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. AveustT 29, 1952 NuMBER 5

TRICHOLIMNAS CONDITICIUS IS PROBABLY A SYNONYM OF TRICHOLIMNAS SYLVESTRIS

(Aves, Rallidae)

By JAmeEs C. GREENWAY, JR.

Comparison of the type of Tricholimnas conditicius Peters and Griscom (Proce. New England Zool. Club, vol. 10, 1928, pp. 99-103) with a series of T'richolimnas sylvestris (Sclater) makes it appear most probable that the type specimen is an immature female of sylvestris. Furthermore, some recently discovered notes in the handwriting of collector Andrew Garrett cast grave doubt upon the theory that Garrett shot the bird on Apaiang Atoll, between Makin and Tarawa, in the Gilbert Islands, as Thomas Barbour postulated in the original description of the supposed species. Both Mr. Peters and Mr. Griscom agree to this.

Naturalists have wondered since then whether a habitat, such as this low sandy island affords, could have supported a population of these rails. The very distance from congeners on New Caledonia and Lord Howe Island, off the coast of Australia, made this seem unlikely, quite apart from the disparity of habitats. However, the arguments brought forward in the original description were difficult to circumvent.

This type and only known specimen of conditicitus was found pre- served in alcohol in the Museum of Comparative Zoology after a period of long oblivion. With it was a label: “Kingsmill Islands, 1861, Andrew Garrett, Collector’. Since it could be proved without question that Garrett was actually on Apaiang Atoll of the Kingsmill, or what is now called the Gilbert group, in the autumn of 1859, it seemed likely that he did collect the bird there and that the date ‘1861’’ represented the year of acquisition by the museum. However, it now seems quite

2 BREVIORA NO. 5

as probable that the specimen has been confused with a consignment of specimens that came from the Kingsmills and that the label was written in the museum under a misapprehension.

Andrew Garrett was a dealer who had in his stock specimens from all over the world. In a letter to Charles E. Hamlin at the Museum of Comparative Zoology, written on April 10, 1878, from Huahine, Society Ids., he says: “In return I shall be glad to receive any land or fresh water shells from the East Indies and marine shells from any part of the world except Eastern States, California, and Polynesia.’” And again, ‘“You can send me in return for this box everything you can spare from your duplicates no matter from what part of the world ... don’t forget the museum publications for which I will send the ferns in compensation.”

From Hilo, Hawaii, he writes on Oct. 10, 1856, that he plans a voyage on the whaler ‘Lydia’, the captain of which was an acquaintance of his.

In a letter to Jas. M. Barnard dated April 29, 1857, and written at Hilo, he gives a short history of his life. He had been a sailor and had visited in past years the ‘“‘Ladrone, Bonin, Loochoo, some of the Australian and several of the East India Islands ...my taste for Natural History, which I dearly love, was acquired while visiting foreign places.”’

The most important bit of evidence has recently been found. A short note in Andrew Garrett’s handwriting and on the same paper used by him in his correspondence is entitled “Remarks upon the Birds of Apaiang one of the Kingsmills Is.’”’ and is reproduced here in its entirety. It bears no date but was probably written in 1859.

“During my short residence at the above location [Apaiang Atoll, Sept.-Oct. 1859] I collected one or two specimens of every bird I could get. As I had not time to prepare their skins, I packed them in alcohol. There is only one species of land bird, and this I never saw but obtained one of its tail feathers which I send you. I think it is a species of Hawk.” (Italics are mine.) Unfortunately this feather cannot be found. Two species of migrant hawks have been reported from the Palau Island; possibly a stray bird reached Apaiang.

Andrew Garrett was an excellent naturalist; Barbour has called him “an extraordinary genius.”’ Although his interests were centered upon fishes and shells, he knew birds. In a contemporary list of birds, mammals of Hawaii, sent to the Museum of Comparative Zoology, he says, (no.) “7 Phaethon This fine bird builds its nests and roosts among the precipices bordering the sea in Hamakua and Hilo.” He

1952 TRICHOLIMNAS CONDITICIUS 3

then describes the bird accurately. And again, (no.) “8 Fulica common about our fresh water ponds.” It is quite unlikely that he would consider Tricholimnas to be a sea bird. Nor is it likely that he would mistake the short, decomposed tail feather of T'richolimnas for that of a hawk.

Unfortunately there is no record of the actual provenance of the specimen but the weight of evidence points to its having come from Lord Howe Id. It is probable that it found its way into Andrew Garrett’s collection by an exchange, or perhaps through the good offices of a whaling friend.

Further questions occur. Assuming that the type was a member of a population which had for some time inhabited Apaiang Atoll, why was it not found by the contemporaries of Garrett? The answer is that none of them visited the island. However, Kubary, Finsch and others did land upon islands of neighboring groups and no such speci- men was recorded. No bird even faintly resembling this has ever been recorded except from Lord Howe Id. and New Caledonia. If the assumption is correct, we must presuppose that wind born or ship born individuals reached Apaiang and no other neighboring islands and that the population was extirpated a very short time after Garrett visited there. It would be strange if such a population had established itself on this flat, almost desert, island in view of the fact that its un- mistakable relatives prefer forests and hills and that the intervening islands afford more suitable habitat. That it became extinct in a number of island groups during the twenty years 1860-1880 is not at all probable.

Granted that no proof can be offered and that at best we are dealing with probabilities, the best calculated guess would appear to be that the type of Tricholimnas conditicius came from Lord Howe Id.

No specimens of sylvestris were to be found in the United States in 1928 when conditicitus was described. Comparison of the type with a series of twelve in the Rothschild collection, now in the American Museum of Natural History, New York, cause most of the characters ascribed to geographical variation to disappear. The paler crown, throat and underparts, the browner head and throat are doubtless the result of long immersion in alcohol and it is surprising that more color change has not taken place. The only differences are then the length of wing and bill, which are two millimeters shorter than those of the smallest female of sylvestris available. Since G. M. Allen, in the original description indicated that the skeleton was obviously that of

4 BREVIORA NO. 5

an immature bird, perhaps it may reasonably be assumed that this character is due to age. Measurements, in millimeters, are as follows:

“conditicius” sylvestris lafresnayanus CE so WViini oe. | eS ihre ee 132 135-139 133-142 185 Dail. avs. S30 58 58-63 61-68 109 Bill pha: Sarin 45 54-56 47-50 52 Marsus 1K: ahs ae 47 47-50 43-48 54

In the original description the length of the tail of the type of conditicius is said to be 68 mm. Mr. Peters agrees with me that it should read 58 mm.

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. August 29, 1952 NUMBER 6

CONTRIBUTIONS TOWARD A RECLASSIFICATION OF THE FORMICIDAE

I. Tribe Platythyreint (Hymenoptera) By Wiuu1AMmM L. Brown, JR.

Museum of Comparative Zoology Harvard University

The tribe Platythyreini has included the sole genus Platythyrea Roger as treated by former authors. My own investigations show that three additional genera (Probolomyrmex, Escherichia and Eubothro- ponera) must be included. Probolomyrmex Mayr and Escherichia Forel have heretofore been placed among the Proceratiini because of their depigmented condition, atrophied eyes (workers), and especially their vertical, fused and approximated frontal carinae and the closeness of the antennal insertions to the median line and to one another, accompanied by fusion of frontal carinae with the greatly crowded clypeus. Also, these two genera have, according to the describers of the included species, only one tibial spur to each of the two posterior pairs of tibiae. I believe that all of the characters just mentioned are correlated with adaptation of the insects to hypogaeic or other crypto- biotic conditions of life; they appear in widely separated genera of ants and other hymenopters, such as Proceratium, Discothyrea and others in the Formicidae, Psilobethylus, etc. in the Bethylidae, and so on, as rather consistent combinations. The similar modifications of doryline and some other ants may be partly due to hypogaeic or subhypogaeic adaptation, but it would seem that the legionary habit may somehow be more important in accounting for this particular structural modi- fication.

For our present purposes, it will be sufficient merely to recognize two facts: (1) the characters combining to produce the “‘proceratiine

y BREVIORA NO. 6

habitus” can and do act to form similar-appearing groups of genera, even though these genera may have the most diverse ancestry; (2) the number and condition of the spurs of the two posterior pairs of tibiae have been found to possess much less taxonomic value than Emery and Wheeler granted in their widely-used keys; in fact, the spurs are now known to be present as pairs, single units, or even to be absent in different species of the same genera in tribes where once the spurs were considered universally constant and tribally diagnostic.

With these facts in mind, one can proceed at once to a more natural classification of the genera and tribes of Ponerinae. Probolomyrmex and Escherichia have always seemed anomalous as members of the Proceratiini, and they are not, in fact, at all closely related to any other proceratiine genus. Following a slight clue dropped by Mann in 1923, when he mentioned the similarity of the pilosity and sculpture between his Probolomyrmex boliviensis and the species of Platythyrea, I have carefully compared the two last-named genera and find that they agree in an astoundingly complete way. In fact, the point-by- point agreement is so close that I must consider Probolomyrmex to represent a direct derivative of Platythyrea modified for a highly cryptobiotic existence in the same way, as I also believe, that the proceratiines are only modified ectatommines. If Platythyrea and Probolomyrmez are to be associated, then Escherichia should probably go along with them. While I have never seen an example of the latter genus, it appears from all accounts to be very close to Probolomyrmez, and may even be synonymous and representative of an ergatoid or other intermediate female caste of one of the Ethiopian Probolomyrmex species.

A fourth genus that must be included in the Platythyreini is Eubo- throponera, in spite of its unfortunate name. In his original diagnosis of this Australian group, Clark stated, . . . certainly close to Bothro- ponera Mayr;” he called the tarsal claws “simple’’ and gave the palpal formula as maxillary 4, labial 2 segments. I have examined types (Museum of Comparative Zoology) of all the Eubothroponera species described to date except E. brunnipes Clark, and find that all species, at least in the worker caste, possess single, small but distinct median teeth on all tarsal claws. The palpi in this genus are unusually long for a ponerine group, and the maxillary pair may reach the foraminal border behind if fully outstretched. Most of the few specimens available were inconveniently situated or else had the maxillae re- tracted, but in spite of this it is clear that all six species seen have more of both maxillary and labial palpal segments than Clark claims

1952 PLATYTHYREINI (HYMENOPTERA) 3

for them. In E. reticulata Clark and E. tasmaniensis (Forel), a full count was made under good conditions, and in these species the formula was maxillary palpi 6, labial palpi 4 segments. The basal segment, especially in the maxillary palpus, is short, fairly broad, and the segment most likely to be overlooked. Comparing the proportions of the visible segments of the palpi of the less favorably situated speci- mens with those upon which a direct count could be made, I feel quite safe in assuming that the six species I have seen all possess a 6, 4 formula in the worker.

Now this formula is the primitive one in Platythyrea, as I can con- firm (vide infra), and since all other characters save one combine to link the two genera, they must at least be placed in the same tribe. In fact, the characters separating them are relatively minor ones, though constant and consistent in combination in the series before me. The body in Eubothroponera is smaller and generally more compact (less slender and elongate), and the characteristic sculpture of Platy- thyrea is less well developed in Eubothroponera, but still basically the same on head and alitrunk. The pilosity must be used to separate the two genera until a study can be made including all castes of a large number of species belonging to both (see key, below).

In examining the palpi of some representative species of Platythyrea for this work, it was confirmed, as has been long known, that certain _ of them have a palpal formula of 6, 4. What is not so generally known, or perhaps even unknown until now, is that certain species of the Old World tropics possess lesser numbers of segments. The New World species P. angusta Forel and P. strenua Wheeler and Mann have elongate palpi, with the 6, 4 segmentation; the basal segment, par- ticularly of the maxillary palpi, is very short, and the total similarity of the palpi to those of Eubothroponera is very striking. Among the Old World forms, P. cribrinodis (Gerstaecker) very definitely has short palpi, formula 3, 2. In P. schultzei Forel, the maxillary palpi have 2 clearly visible segments each, and there may possibly be a third small basal one, though no third segment could be seen in the single specimen examined; the labial palpi are definitely 2-segmented. Other species, such as P. sagei Forel, were not advantageously placed for a definite palpal count, but had short palpi with formulae almost certainly under those of angusta for both pairs. Also seen in one doubtfully determined species from the Old World were palpi with definite counts of 6 and 4, but very short in overall proportions. It seems that Platythyrea species vary considerably from one to the next in palpal characters, and anyone possessing the material needed for a thorough survey of

4 BREVIORA NO. 6

the group by means of dissection may be able to correlate this vari- ation with other characters, and thereby be able to recognize more than one group or genus among the array of Platythyrea species.

Furthermore, the males of Platythyrea seem, from external exami- nation only, to have very distinctive terminalia worthy of further study.

Key to the genera of the tribe Platythyreini, based on the workers and probably applicable to the females

1. Frontal carinae and clypeus fused and projecting anteriorly over the mandibles; antennae somewhat incrassate apically, inserted close together on the anterior part of the clypeo-carinal process. (Small, usually pale- colored forms, under 4 mm. in length. Ethiopian, Neotropical, Indo- TATISOr TAT) eb te ee ea, hes Re RE ac nell ie cia ee ha oh Probolomyrmex Mayr'

Escherichia Forel Frontal carinae and clypeus not projecting anteriorly over mandibles; antennae inserted distinctly posterior to the clypeus and apart from one another, the insertions covered by the broad lobes of the frontal carinae, funiculus not or scarcely incrassate in most forms. (Larger, usually

pigmented forms, mostly over 4 mm. in length).................... 2 2. Distinct erect pilosity present and widely distributed (widespread in Acasbrallis; siecertet Sei Soak biatch Sen aa oe Eubothroponera Clark

Distinct erect pilosity limited to mouthparts and gastric apex. (Tropico- politan; occurring naturally or as tramp species in some warm temperate L:CPat Otc) | eee dy OMEN: Aa ace Rae mR NE ROSE Berend Platythyrea Roger

Below are listed the platythyreine genera as known to me at present, and also a citation of the genotypes and the principal references to the literature. The references are not intended to be complete, but are designed to give the interested worker a start toward the gathering of pertinent titles.

PLATYTHYREA Roger

1863, Berlin. Ent. Zeitschr., 7: 172. Genotype: Pachycondyla punctata Fred. Smith, 1858, soldier (recte worker), male; designated by Bingham, 1903.

Besides the characters cited in the key (above), it should be men- tioned that this genus, and also Eubothroponera, are further marked by

1 Probolomyrmex and Escherichia are supposed to differ in that the latter has small compound eyes and a discernible postmesonotal groove or line, but it is important to note that Forel himself, in the original description of Escherichia, compared the novelty with several proceratiine genera, but made no mention of Probolomyrmex! Certain Probolomyrmezx (P. boliviensis Mann, P. parvus Weber) are known only from the female caste.

1952 PLATYTHYREINI (HYMENOPTERA) 5 having a complete and apparently functionally flexible suture sepa- rating the pro- and mesonotum, and also by having a large and a smaller spur on each one of the two posterior pairs of tibiae.

Emery’s survey of the world species (1911, Gen. Ins., Fase. 118, pp. 28-30) is comprehensive for its time, but is now far out of date due to the addition of many forms since it was written. Wheeler’s “Ants of the Belgian Congo” (1922, Bull. Amer. Mus. Nat. Hist., 45: 57-60, 758-761, 1007) gives references to African and Malagasy species.

EUBOTHROPONERA Clark

1930, Proc. R. Soc. Victoria, Melbourne, (n.s.) 43: 8-9. Genotype: Eubothro- ponera dentinodis Clark, 1930, worker; original designation, nec E. tasmani- ensis (Forel) designated by Clark in 1934.

With the original generic description, Clark characterized and figured three species (pp. 9-11, fig. 1): FE. dentinodis Clark (p. 9,, fig. 1, nos. 6, 6a); E. micans Clark (p. 10, fig. 1, nos. 7, 7a); FE. bicolor Clark (p. 11, fig. 1, nos. 8, 8a); he also included in the genus Forel’s Pachy- condyla (Bothroponera) tasmaniensis (p. 11), and gave a “‘key”’ (p. 9) to the three Western Australian species newly described.

In 1934 (Mem. Nat. Mus., Melbourne, No. 8, pp. 32-34, pl. 2, figs. 15-17) Clark redescribed the worker of E. tasmaniensis (Forel) (p. 32, fig. 15) from Tasmania and described the new species E. reticu- lata (p. 33, fig. 16) from New South Wales and F. septentrionalis (p. 34, fig. 17) from Queensland.

A seventh and last species was added by Clark when he described E. brunnipes (1938, Proce. R. Soc. Victoria, Melbourne, 50: 361-362, fig. 3) from Reevesby Island, Sir Joseph Banks Group, South Australia (biology, loc. cit., p. 356).

The above, I think, includes all the recognized species of Eubothro- ponera, each reported only from the worker caste. The species are quite uniform in structure, and it is felt that Clark has failed in some instances to properly differentiate them; consequently, a review of the genus is necessary.

6 BREVIORA NO. 6

PROBOLOMYRMEX Mayr

1901, Ann. Naturhist. Hofmus. Wien, 16: 2-3. Genotype: Probolomyrmex filiformis Mayr, 1901, worker; monobasic.

M. R. Smith (1949, Proc. Ent. Soc. Washington, 51: 38-40) has reviewed briefly the five species known to that date. A sixth species was described at about the same time by Weber (P. parvus Weber, 1949, Amer. Mus. Novit., No. 1398, pp. 3-4, fig. 2, female) from Africa. A description with a good habitus drawing of the female type of P. boliviensis is given in the original reference by Mann (1923, Psyche, 30: 16-18, fig. 2). A seventh species, apparently remaining undescribed, has been taken in the Canberra region of eastern Australia by Mr. Tom Greaves.

EscHERICHIA Forel

1910, Zool. Jahrb. Syst., 29: 245-246. Genotype: Escherichia brevirostris Forel, 1910, op. cit., pp. 246-247, worker; monobasic.

Although there are entries in the catalogs of Emery and Wheeler, this genus and species seems not to have been reported a second time. It should be noted, however, that Weber’s description of Probolomyrmex parvus, cited above, may just possibly apply to the female of Esche- richia brevirostris when the latter becomes known in association with workers, as the characters and type localities given for the two species do not, to my mind, exclude the possibility of conspecificity.

; | 3 i ks : i i A - ® j = hi P N j q bin ah Ave ‘ea 1 bas jj ts | i ; Vey Hi f ha) Test hae fa pe ; rr.

me A hi ; ee = Oe i pO AN ae PN awit ty 6 ER j 7 . Pa oy) Vg i " y ‘Ms ; a . = * y . ty # v

be T :

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. SEPTEMBER 29, 1952 NUMBER 7

ON THE NOMENCLATURE OF THE PACIFIC GRAY WHALE

By WILLIAM E.. SCHEVILL!

The gray whale of the Pacific has until recently been called Rhachianectes glaucus (Cope). In 1937 van Deinse and Junge published an important paper of which the main part is a thorough comparative study of subfossil Eschrichtius robustus bones from the Netherlands, comprising one adult cranium and two juvenile crania, one of which was associated with the mandibles, scapulae, and other parts of the skeleton. These they compared with an adult Rhachianectes glaucus skeleton in the British Museum, as well as with the published material on both forms. Although the subfossil bones are naturally worn and fragmentary, as well as being for the most part juvenile, they present ample characters to indicate that they and the gray whale are conge- neric. At this point I would like to emphasize that the following criticisms of van Deinse and Junge’s conclusions are directed solely at their taxonomy; this involves no disparagement of their important and valuable osteological work, which has demonstrated the generic identity of Eschrichtius Gray 1864 and Rhachianectes Cope 1869 (= Agaphelus Cope 1868). But they also argue for specific identity, and thus unite Rhachianectes glaucus (Cope) 1868 with Eschrichtius robustus (Lilljeborg) 1861. Moreover, they argue that Dudley’s scrag whale (1725), named Balaena gibbosa by Erxleben (1777, p. 610), is the same form, and therefore say (1937, p. 181), ‘After this it is clear that the name of this whale must be changed into Eschrichtius gibbosus (Erxleben). We must emphasize that with the bones we have now at hand we cannot give any argument that the Pacific and Atlantic representatives of Eschrichtius should be specifically different.” But by the same token these bones cannot prove that these whales are specifically identical.

1 Contribution No. 626 of the Woods Hole Oceanographic Institution.

2 BREVIORA NO. 7

Additional and better material of VY’. robustus may present more specific characters, but until such material turns up, it seems unfortu- nate to take a well-grounded name of a well-characterized living form, known from complete specimens, and replace it with that of a form so imperfectly known from such incomplete material that specific characters cannot be entirely established; instead it seems better not to change the name of the well-known form.

It is even less desirable to use the name (gibbosa Erxleben) of a form which from the beginning has been an unrecognizable literary curiosity. Considering the available information, which is scanty though oft repeated, the most that can be said for Balaena gibbosa Erxleben is that it may be an Eschrichtius, but since it is not specifically well characterized and since nothing sufficiently like it has ever again been reported from anywhere near the type locality (‘the Coast of New England’’) or elsewhere in the Atlantic, it seems to be stretching probability to submerge in it known forms from other regions. De- tailed consideration of the uncertainties in attempting to identify Dudley’s secrag whale (Balaena gibbosa Erxleben) are omitted here. From time to time cetologists have tried to identify this beast, but have always been frustrated by various irreconcilable data, the chief of which is perhaps the utter lack of specimens. Cope’s attempt to embody gibbosa Erxleben was undermined by his confusion over a dilapidated specimen of Balaenoptera acuto-rostrata, so that he formally withdrew in 1884 (p. 1124). Although he did not publish details, it is probable that Cope used much the same reasoning as van Deinse and Junge in associating Dudley’s scrag and the Pacific gray whale in the same genus. Thus it appears best to continue to use robusta Lilljeborg for the subfossil European form.

As to the nomenclature of the Pacific gray whale, it seems reasonable to retain the trivial name g/aucus until better material of E'schrichtius robustus may demonstrate specific identity (although Cederlund (1939, p. 282) believes the holotype sufficient for this). It is therefore sug- gested that the living Pacific gray whale be called Eschrichtius glaucus (Cope).

This status may be summarized thus:

Genus Eschrichtius Gray 1864 (p. 350). Type species (by subsequent designation by Lilljeborg 1865, p. 494): Balaenoptera robusta Lilljeborg 1861. Species Eschrichtius robustus (Lilljeborg) Balaenoptera robusta Lilljeborg 1861, p. 602. The Grias6 whale. Subfossil in northwestern Europe (English Channel to Baltic Sea).

1952 PACIFIC GRAY WHALE 3

Uschrichtius glaucus (Cope) Agaphelus glaucus Cope 1868a, p. 160, 1868b, p. 225. Rhachianectes glaucus Cope 1869, p. 15. The California or Pacific gray whale; devilfish. Recent in North Pacific Ocean (Baja California to Korea).

REFERENCES

Cederlund, B. A. 1939. A subfossil gray whale discovered in Sweden in 1859. Zoologiska Bidrag fraan Uppsala, 18, pp. 269 - 286, 5 pls. Cops, E. D. 1868a. [Remarks on Cetacea at meeting of 23 June.] Proc. Acad. Nat. Sci. Phila., 1868, pp. 159-160. 1868b. On Agaphelus, a genus of toothless Cetacea. Proc. Acad. Nat. Sci. Phila., 1868, pp. 221-227. 1869. Systematic synopsis of the species of the cetaceans of the west coast of North America. Proc. Acad. Nat. Sci. Phila., 1869, pp. 14-32. 1884. Catalogue of aquatic mammals of the United States, by F. W. True. Review. American Naturalist 18, no. 11 (Nov.), pp. 1123-1124. vAN DeinsE, A. B., and G. C. A. JUNGE 1937. Recent and older finds of the California gray whale in the Atlantic. Temminckia, 2, pp. 161-188, pls. 4 - 11. Dup.tey, Pau 1725. An essay upon the natural history of whales with a particular account of the ambergris found in the sperma ceti whale. Philos. Trans. Roy. Soc. London, 33, no. 387, pp. 256-269. ERXLEBEN, J. C. P. 1777. Systema regni animalis ... Classis I. Lipsiae, pp. xlvili + 636 + [66]. Gray, J. E. 1864. Notes on the whalebone-whales; with a synopsis of the species. Ann. Mag. Nat. Hist. (8) 14, no. 83 (Nov.), pp. 345-353. LILLJEBORG, W. 1861. Hvalben, funnaijorden paa Grason i Roslagen i Sverige. Forhandl. Skandinav. Naturf., 8de Mgde, 1860, Kigbenhavn, pp. 599-616. 1865. [Note quoted in] Gray, J. E., Notice of a new whalebone whale from the coast of Devonshire, proposed to be called Hschrich- tius robustus. Ann. Mag. Nat. Hist. (8) 15, no. 90 (June) pp. 492 - 495.

eo pennant: peal Opt otis sais ie ia ripe YE OMe vba: mee

pee aah pont: » agin

eae iss a 1a) abpy _ Ny

: m y ie is en ntie Py es onal

: Pa Pee Ctrl ki de ‘eueieiag of a tea a Alot Let al

, ey

: } ae! A 4 A i ; devrabtered ad caveat & ya Cont er fs, + uf vie yg

an My Havel Aq BORE aru Yids

ae eri aE, Sealy) Helilyg™ ter kira’ 2 Lanka sO ale spe oe = ir} wae aaa! ai Na 1

ann lhs Mieke. a “Hpadindey! aie hye nthe alti

aae- Ta Pee aay ia Rhee: SUA died iii

ie a) as rs Lia OF a ,

Fi Sh) Aint hatin? aly Ww Spent nay) eisopnAi Pipes

pet. of aA Jt, Lee AL poll seni’ tim rates muah

ay : a & ry : 7 = ? aT | j TO te t a P| a aed a Asset siti gine i os uy ay coe Aart a

‘fice ea #3 She ue r

al *

ie

ear PD Ateer eles 4 sh a) eh Salt Hiity Meet at hiybPakgpalig ay Us

a CGA Bs Te Aten natine ert more ; ra a ws ty Hy uly +e tele © a —. ie aa a

a

oa Lisa nite i sil fi Big, cess atl? peelg Ws? ign) ey Lar UY aa “e ME in or on tien ane i) ey gens (icone Pett ack, wit , yuiieg RAS RAG os A OORT oa a. hater Boer: | gta ai ink Agri siniy go Wout 4 Y eer tat feo henge 2! lisa A” WAlhe ga: fat Wpegn fc? 12h adie vol Kany § Prose meas ai eet itt oh a

7 pa oN J tad > - ae | :

ae ia : i .

—— y al 9 4 Doig S .

ee ae, Vieni Py act = an ¥- eee

—_ »

BREVIORA

Museum of Comparative Zoology

(AMBRIDGE, Mass. OcToBER 21, 1952 NUMBER 8

A NEW SPECIES OF THE CYCLOSTOME GENUS PARAMYXINE FROM THE GULF OF MEXICO

By Henry B. BiagELow and WILLIAM ©. SCHROEDER!

Up to the present time the myxinids, represented in the North Atlantic by the unibranchiate genus Myzine alone, have been unknown in the Gulf of Mexico. During 1951, however, the experimental trawlings of the U. S. Fish and Wildlife Service vessel “Oregon’’, in the northern part of the Gulf, yielded three specimens of a myxinid with six pairs of external gill openings that agree closely with the genus Paramyxine Dean 1904, reported previously from Japan only.’ Paramyzxine, originally described from a single specimen, as the new species P. atami Dean 1904, is characterized, by its author, by having the outer “branchial ducts of distinctly unequal length, the most anterior several times the length of the posterior” (Dean, p. 22), so that the most anterior pair of external gill openings is opposite the 4th or 5th pair of internal gill sacs; and by having the successive pairs of gill openings close together. In other known myxinids with more than one pair of gill openings the successive pairs of external branchial ducts are of approximately equal lengths, or the Ist pair is only a little longer than the rearmost pair, and the successive pairs of gill openings are more widely spaced. On the type specimen the gill openings them- selves were oval, with their long axes transverse (except for the 6th opening on the left-hand side, which receiving the oesophageal duct is larger than the others), not round as they are in other polybranchiate species. But the gill openings were of various shapes in other specimens described subsequently by Matsubara (1937), showing that their shape is not a generic character as Dean supposed,

Contribution No. 610, from the Woods Hole Oceanographic Institution. 2 We owe to Mr. Stewart Springer the opportunity of studying these interesting specimens.

2 BREVIORA NO. 8

or even a specific character, but is, likely, dependent to a considerable extent on the degree of muscular contraction. This leaves only the relative lengths of the external branchial ducts and perhaps the close spacing of the gill openings, as compared with other myxinids in which they are few in number, as bases for separating the genus Paramyxine from other polybranchiate myxinids. We accept the genus provision- ally, awaiting the opportunity of examining other myxinids with corresponding numbers of gill openings.

Characters that Dean (1904, p. 22) included as specific for P. atami were the precise number (6) of pairs of gill openings; Ist pair of ex- ternal branchial ducts about 3 times as long as the 6th pair; 12 and 13 lingual teeth in the outer rows with 13 in each of the inner rows; anterior 3 lingual teeth of each row fused with one another at their base; white gill openings and conspicuously dark mucous pores. His illustration also shows a rather pointed snout, resembling that of Myzxine, but a somewhat broader and more paddle-shaped caudal fin fold.

The gill openings of the type specimen of atamz occupied a space about 9 per cent as long as the distance from the tip of the snout to the most posterior pair of gill openings,! with those on each side arranged close together in a regular row, and they are arranged simi- larly in an excellent specimen of atami, from Suruga Gulf, Japan, received recently by the U. S. National Museum (U. S. Nat. Mus. No. 161442). But Matsubara’s (1937) observations have shown that the serial arrangement of the gill openings, like their shapes, may vary widely from individual to individual, and even between the two sides of the body of a given specimen, for they were irregularly arranged in 13 of his 14 specimens. Muscular contraction may play a role here. Matsubara also found that the dental formula is not a precise one, for his specimens had only 10 or 11 lingual teeth in each row, outer and inner, with only the first two fused at the base instead of three as was true of the type specimen.

The Gulf of Mexico specimens agree in general with the accounts and illustrations of the Japanese atam7 and with the National Museum specimen, in body form, with rather pointed snout and a caudal fin fold somewhat wider than that of Myaine. Their barbels are similar to those of atami in length and in arrangement; they have the same number (6 pairs) of external gill openings; the external branchial ducts (as measured by probing) are about as long relative to the 6th pair;

1 As scaled from Dean’s (1904, pl. 1, fig. 3) illustration.

1952 NEW SPECIES OF CYCLOSTOME 33

the foremost pair is situated about as far rearward while the gill openings cover about as long a space relatively; the distribution of mucous pores is about the same; also the linear dimensions in general, as appears from the following tabulations:

Table 1 Distances, in Per Cent of Total Length Gulf of Mexico atami, atami, U.S. specimens, 338, Japan type Nat. Mus. 055 and 590 mm. 550 mm. specimen 500 mm. Snout to foremost mucous pore 7.5- 8.7 6.8 9.0 Snout to foremost gill openings 22.5-23.5 26.0 27.0 Ist gill opening to 6th gill OVEDIN Pesag 7 Sins susie Shae a 2.4— 5.7 2.5 2.4 6th gill opening to tip of caudal (TA S10) fo ae ae eee ee 72.0-74.3 (alee: 73.0 Snout to origin of anal fin fold 37.1-50.0 41.4 43.4 Cloacal pocket (center) to tip of caudal fin fold:.....:..- 13.4-17.5 12.8 1322, Table 2

Numbers of Mucous Pores on Each Side Gulf of Mexico atami, type! atami, U.S.

specimens Nat. Mus. specimen Snout to foremost pair of gill openings................. 15-15 to 19-19 19-20 16-16 6th pair of gill openings to cloacalkpocketinus. + 4244¢-- 44-45 to 57-57 58-59 47-47 Center of cloacal pocket to tip of caudal fin fold.......... 11-11 to 13-14 ? 10-10

The number of lingual teeth, too, is about the same in a Gulf of Mexico specimen of 505 mm. (14 and 13 in the outer rows, 11 and 12 in the inner rows) as it was in the type specimen of atami, with the same number (3) of anterior teeth fused basally in each of the outer two rows. And while two only, of the anterior teeth in each of the inner rows are fused in the Gulf of Mexico specimen, three were fused in the type, and Matsubara (1937, p. 15, text fig. 3) has already reported and figured Japanese specimens that agreed with the Gulf of

1From Dean 1904, p. 11, text fig. 2, D.

4 BREVIORA NO. 8

Mexico one in this respect. The series of mucous pores on each side is interrupted near the anterior end of the cloacal pocket on the U. 8. National Museum specimen of atami (Fig. 6), just as it is in the Gulf of Mexico specimens (Fig. 5) in Myzine too, for that matter to continue rearward at a higher level. Representation of it in Dean’s (1904,pl. 1, fig. 3) illustration as continuing unbroken past the cloacal pocket seems to have been an error by the artist. But the rearward pores are situated higher above the ventral edge of the caudal fin fold in the Japanese atami (Fig. 6) than in the Gulf of Mexico specimens (Fig. 5).

The depth, too, of the body relative to the distance from tip of snout to Ist pair of external gill openings is greater on all three of the Gulf of Mexico specimens (34 per cent to 37 per cent) than is pictured for atami either by Dean (30 per cent) or by Matsubara (22 per cent), or than it is on the U. S. National Museum specimen of atami (29 per cent). And while the hagfishes are so soft of body that measure- ments of different specimens cannot be accepted as precisely com- parative unless taken under equal tension, the difference in this respect between the two geographic populations may be large enough to be significant. The shape of the caudal fin fold also differs rather notice- ably, its lower margin being nearly straight in all three of the Gulf of Mexico specimens while it is pictured as moderately convex in the two Japanese specimens of atami which have been illustrated (Dean 1904, pl. 1, fig. 3; Matsubara 1937, pl. 1, fig. A), likewise in the National Museum specimen of atami (compare Fig. 5 with Fig. 6).

Furthermore, neither of the two larger Gulf of Mexico specimens shows any trace of a dorsal fin fold anterior to the level of the cloacal pocket, whereas Matsubara’s (1937, pl. 1, fig. A) illustration of atami clearly shows such a fold, extending forward for one third to one half the distance from the level of the cloacal pocket toward the level of the 6th pair of gill openings while the dorsal fin fold on the National Museum specimen of atami is continued forward as a definite though low dermal ridge to abreast of the 6th pair of gill openings.!

These differences between the Gulf of Mexican and the Japanese representatives of the genus seem sufficient, in combination, for recog- nition in nomenclature. We therefore propose for the former the new specific name springeri, as defined below. What is most interesting, however, in this case, is not that the Gulf of Mexican population

1The anal fin fold also, of the National Museum specimen of atam7 is continued forward

nearly to the 6th gill opening as a prominent dermal ridge. But the appearance of the latter, in alcohol, suggests that it was the result of aulmuscr contraction, not a normal] characteristic,

1952 NEW SPECIES OF CYCLOSTOME 5

differs in minor ways from the Japanese it would be astonishing if such were not the case but that two populations so widely separated should resemble each other so very closely, and that myxinids with five to seven pairs of external gill openings should have been found at localities as widely separated as the Gulf of Mexico, Japan and China, New Zealand and South Africa, but nowhere else in the oceans.! This instance does not stand alone as it concerns the Gulf, for the chimaeroid Hydrolagus alberti Bigelow and Schroeder 1951a, recently discovered in deep water there, seems more closely allied to the Japa- nese H. mitsukurti (Dean) 1904, than it is to either of the two species of its genus that are known from the North Atlantic, i.e., affinis (Brito Capello) 1868, and mirabilis (Collett) 1904, or than it is to H. collet (Lay and Bennett) 1839, of the Pacific coast of North America. Equally unexpected was the recent discovery, in the Gulf, in moder- ately deep water, of a representative (Springeria folirostris Bigelow and Schroeder 1951), of the rajoid family Anacanthobatidae, for the latter had been known before only off the Natal coast of southeast Africa; also of two representatives of the skate genus Cruriraja in Cuban-Floridan waters (C. atlantis and C. poeyi Bigelow and Schroeder 1948), for this genus, too, had been known only off Natal, southeast Africa, previously.

Altogether, the bottom zone in the Gulf and around Cuba at 75 to 600 fathoms has yielded three new species of sharks, eleven new batoids, and one new chimaeroid since 1938, a foretaste of the rich harvest of unknown fishes to be expected from a more thorough ex- ploration of the deep slopes and of the sea floors of the Gulf and of the Caribbean.

PARAMYXINE SPRINGERI, new species”

Type. A specimen, 590 mm. long, Oregon Station No. 489, Lat. 27°44’N., Long. 85°09’W., 254 fathoms, Sept. 29, 1951 (U. S. Nat. Mus. No. 161512).

Additional material. Another specimen 505 mm. long from the same trawl haul, and a third, 338 mm. long, from Oregon Station No. 321, Lat. 27°27’N., Long. 87°19’W., 220 fathoms, April 28, 1951. The sex has not been determined for any of these specimens.

Description. Dimensions, in per cent of total length, of type (590 mm.), 505 mm. specimen, and 338 mm. specimen.

Depth of body, at Ist gill openings, 9, 7.8, 8.9.

1 For a synopsis of the species of myxinids, see Holly, 1933. 2 Named after Stewart Springer who discovered this interesting cyclostome.

6 BREVIORA NO. 8

Distance, snout to Ist gill openings, 23.5, 23.2, 22.5.

Distance, Ist gill openings to 6th gill openings, 2.4, 2.6, 5.7.

Distance, snout to origin of anal fin fold, 50.0, 37.1, 39.8.

Distance, center of cloacal pocket to tip of tail, 14.5, 13.4, 17.5.

Depth, in per cent of distance, snout to Ist gill openings, 38, 34, 37.

Number of mucous pores on each side, type (590 mm.), 505 mm. specimen and 338 mm. specimen:

Snout to Ist gill openings, 19-19, 18-18, 15-15.

1st gill openings to 6th gill openings, 4-3, 3-3, 6-6.

6th gill openings to anterior end of cloacal pocket, 55-56, 57-57, 44-45,

Anterior end of cloacal pocket to tip of caudal fin fold, 13-14, 11-11, 12-12.

Snout obtuse-conical, much as in Myzine; body cylindrical an- teriorly, about 80 to 100 per cent as thick (transversely) at region of gill openings as deep there, but increasingly compressed posteriorly, until only about 38 to 54 per cent as thick as deep at level of cloacal pocket, and flat sided thence rearward to paper-thin margin of caudal fin fold. Depth of body about 5.4 to 6 per cent as great as total length at level of most anterior mucous pores, about 6.3 per cent at anterior end of cloacal pocket on type, and 6.6 to 8.6 per cent on smaller specimens.

Fleshy rostrum, anterior to nasal opening, broadly rounded an- teriorly and wider than long,! the nasal opening conspicuous below it. Barbels in the typical myxinid arrangement, the Ist pair about two- thirds as long as the 2nd pair, the 3rd _ pair a little the longest. Mouth, when closed, with the usual puckered or irregularly stellate appearance, the conical projection that flanks it on either side about as long as the 2nd pair of barbels. Thirteen and fourteen lingual teeth in the outer rows, eleven and twelve in the inner rows of 505 mm. specimen’, individual teeth conical with slightly blunted tips, curving a little rearward, the most posterior of each inner row strongly so; the 5th and 6th tooth longest in each row; the 7th and the following teeth successively shorter along each row; the rearmost only about one-third as long as the 5th and 6th. The first three teeth in each outer row, and the first two in each inner row fused at their bases; the outer row in each side about 1.2 times as long as the inner row; the longest teeth in the outer rows about 1.3 times as long as the

longest in the inner rows.

1 Longer than wide in Myzine. : 2 The teeth of the type specimen cannot be counted without dissection.

=~

1952 NEW SPECIES OF CYCLOSTOME

The six pairs of gill openings are low down on the sides, their fore-aft position as given in Table | (p.3), occupying a shorter space on larger specimens (2.4 per cent of total length on the type) than on smaller, the series on the two sides converging rearward, with the 6th pair only about three-fourths as far apart as the Ist pair on the type, about four-fifths as far apart on the smallest specimen; the 6th opening on the left-hand side (receiving the oesophageal duct) larger than the others as is the rule among myxinids, and irregular in shape, the other gill openings ranging in shape from round to oval with the long axis transverse or slightly oblique, the precise shape seemingly dependent on muscular contraction; the largest (apart from the 6th left-hand) a little less than one-half as long as the 3rd (longest) pair of barbels. The gill openings are either in regular serial arrangement on each side as on the type (Fig. 2) and on the smallest specimen, or more or less irregular at least on one side, as on the 505 mm. specimen (Fig. 3), a difference perhaps associated with muscular contraction. The Ist pair of external branchial ducts (as sounded by probing, on one side of the body) are about three times as long as the 6th pair, both on the type and on the 505 mm. specimen.

The mucous pores range in size from easily visible to so minute as to be discernible only on close examination. Their location on the body and in serial arrangement is as in the other myxinids; 19 pores on each side from snout to Ist gill openings on the type, 4 on one side and 3 on the other abreast the gill openings, 55 and 56 between gill openings and cloacal pocket and 13 and 14 rearward from the anterior end of the cloacal pocket on the type specimen; their numbers on the other specimens as given in Table 2 (p.3). Totals of 77 pores on the 338 mm. specimen, of 88 on the 505 mm. specimen, and 92 on the 590 mm. specimen (type) show that the pores increase in number with growth. The pores along the section occupied by the gill openings are ventral from the latter, which it seems is characteristic of the poly- branchiate myxinids in general.

The pores rearward from the gill openings are nearly in line with the latter, are low down on the sides nearly to the anterior end of the slit-like cloacal pocket, where the series is interrupted, with the nine posterior pores situated at a definitely higher level, a distinctive character of this species as contrasted with the Japanese atami (p. 4). On the type, those rearward from the cloacal pocket are also noticeably larger (hence more conspicuous) than those farther forward, and they are somewhat more conspicuous than the more anterior pores on the

8 BREVIORA No.8

505 mm. specimen as well. But this regional difference in size of mucous pores is not a specific character; rather it is likely a phase of muscular contraction, for the pores of the caudal series on the smallest specimen are no larger than those anterior to the cloacal pocket.

The point of origin of the anal fin fold, about midway of the body on the type specimen, is considerably farther forward on the other two specimens (see Table 1, p. 3) showing that its precise situation is variable within rather wide limits. On the type, the fold itself is so narrow that its width is hardly measurable, and it is traceable rearward only about half the way from its point of origin toward the cloacal pocket. But it is wider on the other specimens (8 to 13 per cent as wide as the body is deep), and traceable rearward nearly or quite to the cloaca. To what degree these differences are associated with different stages in growth, or in the sexual cycle, is a question for the future.

Neither the type, nor the 505 mm. specimen shows any trace of a dorsal fin fold anterior to the level of the cloacal pocket, their backs being evenly rounded, in which respect they contrast with P. atamz as pictured by Matsubara (1937, pl. 1, fig. A). And while the smallest (338 mm.) specimen has a low fleshy ridge all along the back nearly to the level of the mouth, the fact that it is flanked on either side by a shallow and indefinite furrow suggests that it is the result of muscular contraction, rather than a definite and persisting fin fold. The caudal fin fold fringes the rear section of the trunk, from about opposite the anterior end of the cloacal pocket on the dorsal side, around to the posterior end of the cloaca on the ventral side. Its lower outline is nearly straight as remarked above (p.4). The irregular fluting of its broadly rounded, posterior-dorsal margin on the smallest specimen (Fig. 5) seems to represent its normal outline better than its margin on the type, (Fig. 1) where it seems to have been injured.

One of the most interesting features of P. springeri, reported to us by its discoverer, is that no one of the specimens discharged any mucus at all when brought on board, or so little that none was noticed.

Color. All three specimens are greyish brown, faintly tinged with reddish, below as well as above, the edges of the caudal fin fold and of the anal fin paler; the gill openings encircled with white, rendering them conspicuous against the dark background; the mucous pores appearing as dark brown dots, each of them at the bottom of a shallow depression of the skin.

Size. The maximum size is not known.

1952 NEW SPECIES OF CYCLOSTOME 9

Range and habits. So far known only from the northern side of the Gulf of Mexico, at the localities listed on page 5. The depths of capture, added to the fact that fishermen have never reported it, show that it is a deep-water species. Nothing else is known of its habits.

REFERENCES BriceLtow, Henry B., and WILLIAM C. SCHROEDER 1948. New genera and species of batoid fishes. Journ. Marine Research, vol. 7, no. 3, pp. 548-566. 1951. A new genus and species of anacanthobatid skate from the Gulf of Mexico. Journ. Washington Acad. Sci., vol. 41, no. 3, pp. 110-113. 195la. Three new skates and a new chimaerid fish from the Gulf of Mexico. Journ. Washington Acad. Sci., vol. 41, no. 12, pp. 8383-392. Brito CapELLO, FELIX DE 1868. Catalogo dos peixes de Portugal. Jorn. Sci. Math. Phys. Nat. Acad. Lisboa, vol. 1, no. 3, pl. 3, fig. 1A. 1868a. Descripcaao de dois peixes novos de Portugal. Jorn. Sci. Math. Phys. Nat. Acad. Lisboa, vol. 1, no. 4, pp. 314-317. Co..etTt, R. 1904. Diagnoses of four hitherto undescribed fishes. . . . Selsk. For., 1904, no. 9, 7 pp. DrAN, BASHFORD 1904. Notes on Japanese myxinoids. Journ. Coll. Sci. Imp. Univ. Tokyo, vol. 19, art. 2, 23 pp., 1 pl. Ho.uiy, MAxIMILiAN 1933. Cyclostomata, in Schultze and Kukenthal, Das Tierreich, lief 59, 62 pp. Lay, G. T., and E. T. BENNETT 1839. Fishes, in Richardson and others, Zoology of Captain Beechey’s voyage, pp. 43-75. MatsuBaRa, K1YoMATsu 1937. Studies on deep sea fishes of Japan. III. On some remarkable variations found in Paramyzxine atami Dean. Journ. Imper. Fisheries Inst. Tokyo, vol. 22, no. 1, pp. 18-15, 1 pl.

10 BREVIORA NO. 8

Fig. 1. Type specimen, x 14; the mucous pores somewhat exaggerated.

Fig. 2. Ventral view of anterior part of type specimen, x 14; the mucous pores somewhat exaggerated.

Fig. 3. Ventral view of gill region of specimen 505 mm. long; natural size; the mucous pores somewhat exaggerated.

Fig. 4. Lingual teeth of specimen 505 mm. long, from the right-hand side; outer row above, inner row below, anterior ends to the left, x 2.

Fig. 5. Side view of caudal region of specimen 338 mm. long, x 14; the mucous pores slightly emphasized.

Fig. 6. Side view of caudal region of P. atami, 500 mm. long, from Suruga Gulf, Japan, U.S. Nat. Mus.,No. 161442;the mucous pores slightly emphasized.

7 ai? [ 1% ; : i iu \ Rie ania iA bd Dia cae is OH eh | Td TA eR Na

ma - iy yk J f er

BREVIORA

Museum of Comparative Zoology CAMBRIDGE, Mass. OcToBER 21, 1952 NuMBER 9

NEW SPECIES OF EARTHWORMS FROM THE ARNOLD ARBORETUM, BOSTON

By G. E. GAtTEs if

From Allolobophora caliginosa (Savigny) 1826 there have recently been split off: A. nocturna Evans 1946, and A. zowana Evans 1948. The new species were thought to be distinguished by differences in location of first dorsal pore, extent of male porophores, etc., and es- pecially by number of segments. These characteristics had not previ- ously been given such weight in lumbricid taxonomy, may be subject to variation and, as in the case of segment number, may be tedious to use when dealing with large numbers of worms.

While working on collections from the Arnold Arboretum and other localities near Boston, it was found that material with characteristic caliginosa tubercula pubertates could be rapidly sorted into two groups according to the presence or absence of genital tumescences on segment xxxlii. Further study showed that those specimens with tumescences on xxxili were referable to caliginosa as apparently restricted by Evans after breeding the worms in the laboratory. The worms with no tumescences on xxxill were found to be consistently distinguishable from caliginosa by the same sort of characteristics as in the case of nocturna and iowana and, as well as those species, worthy of specific status.

ALLOLOBOPHORA ARNOLDI N. sp.

Type. Museum of Comparative Zoology, cat. no. 4441.

Length, 55-100 mm., diameter, 4-6 mm. Segments, 152-194 (normal specimens). Unpigmented and almost white, or of greyish appearance or, especially in older worms of second season, with a brownish coloration. First dorsal pore, on 10/11, 11/12, or 12/13. Clitellum, on xxvii, 46xxvii, or xxviii to xxxiv or Yoxxxv. Tubercula

3, BREVIORA No. 9

pubertates of double origin, on xxxi-xxxill. Genital tumescences, in- cluding a and b setae, on 1x—Xi, XXX, XXxll, xxxiv, and occasionally also On XXVI.

Spermathecal pores, on 9/10-10/11 on ¢ lines, setae, male poro- phores, calciferous sacs, calciferous glands, typhlosole and last hearts, etc., as in caliginosa.

Type locality. Arnold Arboretum, Boston, Mass.

Distribution. Known at present only from Boston and vicinity.

Remarks. A. arnold? is distinguished from caliginosa by the constant absence of genital tumescences on xxxili, occasional presence of tumes- cences on xxvi (instead of xxvii), slightly greater anterior extent of the clitellum (xxvii or xxviii instead of xxviii or xxix), greater number of segments, and perhaps by a slightly more anterior first dorsal pore.

From iowana, arnold: is distinguished by the constant presence of genital tumescences on xxxiv, constant absence of those tumescences on xxxill, by the two-part tuberculum about as in calzginosa, possibly also by a slightly more posterior first dorsal pore, greater number of segments and absence or slighter development of pigmentation (not dark reddish brown).

From nocturna, arnoldi is distinguished by the constant absence of tumescences on xxxiii and xii, smaller size, fewer segments, less re- stricted male porophores, and possibly by a more posterior first dorsal pore.

Data as to variation in both caliginosa and arnold: in the Boston area will be presented, if possible, in another contribution.

He

In lumbricid taxonomy considerable weight has been given in the past to rather small differences in the tubercula pubertates. Thus, for instance, the species in the following pairs have been distinguished from each other by a one-segment homoeosis of the tubercula: Octo- lasium cyaneum (Savigny) 1826 and O. lactewm (Orley) 1881, xxx-xxxili and xxxi-xxxiv; Lumbricus rubellus Hoffmeister 1848 and L. castaneus (Savigny) 1826, xxvili-xxxi and xxix—xxxiil. Further, Dendrobaena rubida (Savigny) 1826 has been distinguished from D. subrubicunda (Eisen) 1874, by a one segment difference in length of the tubercula, XXIX-xxx and xxvili-xxx. Genital tumescences, as indicated above, may also be of considerable value in lumbricid taxonomy.

Accordingly, it seems advisable to treat a worm distinguishable at present from the caliginosa-complex only by differences in tubercula

1952 NEW SPECIES OF EARTHWORMS 3

and tumescences as specifically distinct. ALLOLOBOPHORA MOLITA Nn. sp?

Type. Museum of Comparative Zoology, cat. no. 4442.

Length, 81 (+ ?) mm., diameter, 5 mm. Segments, 140 (+ ? pos- terior amputee?). Pigmentation lacking. First dorsal pore in 712/13. Clitellum on xxviii-xxxiv, possibly reaching slightly onto xxvii. Tubercula pubertates, longitudinally placed bands in bc, extending across entire lengths of xxx—xxxiil; with lateral margins nearly smooth, but median margins (quite definitely lateral to b lines) slightly indented by 30/31, 31/32, 32/33. Genital tumescences, including a and b setae, present on ix—xi, xxxli and xxxili (feebly developed).

Spermathecal pores, male porophores, setae, calciferous sacs and glands, typhlosole, last hearts, as in caliginosa.

In the clitellum intersegmental furrows are obliterated and dorsal pores are occluded but the epidermal thickening is slight. Sperma- thecae (in x and xi) are iridescent and presumably contain spermatozoa obtained from a copulatory partner. Male funnels are iridescent, also indicating sexual maturity in spite of the feeble development of the clitellum.

Remarks. A. molita is distinguished from all of the caliginosa- complex: caliginosa, nocturna, iowana and arnoldi, by the four-segment tubercula pubertates. Further distinction may be provided by an apparent restriction of genital tumescences in clitellar region to xxxii- xxxill. From all of the complex, except possibly iowana, molita is also distinguished by the quadripartite (instead of double) origin of the tubercula. In zowana, tubercula are band-like but nothing was said as to origin.

Type locality. Arnold Arboretum, Boston, Mass.

Distribution. At present known only from the type locality.

REFERENCES Evans, A. C. 1946. A new species of earthworm of the genus Allolobophora. Ann. Mag. Nat. Hist. (11) 138: 98-101. 1948. On some earthworms from Iowa, including a description of a new species. Ann. Mag. Nat. Hist. (11) 14: 514-516.

bene he bata “#

“i ‘a a stink > a

ae

aed a ths cai sigh oe) ley nen

iit Wee eae tela Ae RR - Mele, ae

4. ‘y a ) py pa ua? O71) + ad sgpiiay jm | eed uN

iy uncer ah. (ap! Pica! ill | a ae nee ren

ny hire teas, Ot dvd Peart, Dad a poe wide 2 a lS SY Seal AA spihaeh metey 1s hi) Theil ae

7 ty reals ral lations: pres A TY On dead Ce “seth iaoh WL 2 hierar ny ? ent Say ah it evennie igh lap iutas) a en

oe (aii, COO PL wed. Tal ®.71e gt Mean

ivy aa ee "poheabigy grit ithe i? ee ous 4 toe a ep

aos Ne Me Aras Legh. Tab oyd yp ee

yi f=

ran

eee: ica bi) We MSE) Selene acta ann load + i ee ABABA. 08 aieashal uo teuenalue> qt hte tele

f . ctrl Ramictad iets) agysity ny Nog her yhiagt gga pete i iy <

ere me eye eo ee Anshan att) Wa, Lene atdeaty ol cheat i 3h ona ae Vleet FONG Lae

es . ¥ rd

7 ; f : és Feta y My \ ook

| 5 AV, obs west) fn Be “he 1% Ps ett) tia Bk Bikede ot

1S,

Re roy) cane ee i il i) Ris Ree Os AY mi, eee era iy ai vulr hyi hire ngitrut’ beat)

wg’ i!

Ng ky gies ap teed j U ee Bee hae lie is . ie * ade A 3) ) LE REA Ng . kl a at a .

wd te vo toltiot he } OTS at! 0 "ads ve ‘hartet ' i bated tippi a d id rary pared Ai ifthe hs ee ee ; : 7 ¥ ) ee > 7 i a i th ll 1) ; blot J sy ¢ De ¥ 4 toa sera) “At eS : 1 Ow rae,

r it ety Me . Jon Copa ae ye: P u Oy fa enue by an A ar Dt h wt) ; ut dil, 2 on ahh Peet ae HA’ , ry OT a ae i : 49 ie |e fay = # an : _ oi : xt “0 F » i , ay ig or 7 gq ay iy a 7

sp ars

ee eae

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. DECEMBER 29, 1952 NuMBER 1(0)

ON THE EARTHWORMS OF NEW HAMPSHIRE

By G. E. GATES

So far only one record of earthworms in New Hampshire has been encountered in the literature. The worms were found “under the bark of trees” at elevations of 2,000-3,000 feet in the White Moun- tains by a German delegate to the International Zoological Congress at Boston in 1907. The specimens went to an Irish Museum where they were identified as Dendrobaena rubida (Savigny) 1826, by South- ern (1910). The species had not previously been reported from North America and since 1910 there has been no further record from this continent. The record was given in a short article on another kind of worm and, though the article was published in Philadelphia, the record was not included in Smith’s review (1917) of the Lumbricidae of North America.

Through the kindness of Dr. P. J. Darlington, Jr. who donated the bait remaining from a fishing trip, it is now possible to list a second species for New Hampshire.

Southern (1910, p. 18) quoted his collector who stated that “earth- worms were very rarely met with in the forests at elevations of 2,000- 3,000 feet.’ Similar statements have been made in the past with ref- erence to forests of northern New England as well as of northern New York. Accordingly, Dr. Darlington’s observations appear to be worthy of record:

‘““My experience has been that it is always possible to find earth- worms in fair numbers even in the heavy woods in the more remote parts of New England, or at least in the White Mts. and the Connecti- cut lakes districts of New Hampshire. I have looked for them many times when I wanted them for bait for trout and I have always been able to find them under cover along the banks of brooks. I am fairly

2 BREVIORA No. 10

sure that there are at least two species. One,... (see below) .. . al- though it occurs on the stream banks, is usually well above the water level under stones or logs or in the roots of vegetation in sandy places that are damp but not wet. The other species is darker and much smaller, in fact hardly large enough to use for bait and it seems to oc- cur in much wetter places, under stones on gravel bars or sandy banks very close to the water level... . Practically every brook in New England is fished these days, and many of the fishermen bring worms with them and throw out what may be left at the end of their fishing trips, so that different species must have been introduced many times, even in the deep woods.” (Darlington, 7n Jit.)

ALLOLOBOPHORA ARNOLDI Gates 1952

In steep bank of East Inlet Brook, about 2,000 ft. above sea level, near spruce, in heavy but not virgin forest, above Second Connecticut Lake, May 30, 1952. Four clitellate specimens. Dr. P. J. Darling- ton, Jr. collector.

Length, 95-105 mm. Diameter, 5.5 mm. Segments, 91 (amputee), 106 (amputee), 156, 158. First dorsal pore, 712/13 (1), ??13/14 (2). (@ litellum, $xxvil-xxxiv. Tubercula pubertates bipartite and.as in

A. caliginosa (Savigny) 1826. Genital tumescences, on ix-xi (4), XXX, Xxxil and xxxiv (4), xxix (1);

Remarks. Alive, these worms appeared to be quité without pigment and they looked in that respect much as Octolasiwm cyanenm it Savigny) 1826 usually does.

The epidermis of segments xxvi, anterior half of xxvii, and of XXXV, is somewhat thickened and of a translucent appearance quite distinct from that of normal epidermis as well as from the yellow or white opacity of the clitellar epidermis, on three specimens. The demarca- tion of opacity and translucence on xxvii is indistinct, one gradually passing into the other, on the fourth worm.

On the last segment of the amputees, rudiments of setal follicles and of nephropores are still recognizable. In one of those specimens the new anal region is small and scarcely distinguishable from the ninety- first segment but probably is demarcated by a quite fine, greyish trans- lucent furrow (presumably a rudimentary interségmental furrow). In the other amputee no special anal region is recognizably demarcated.

A. arnoldi was split off from A. caliginosa when it was found in Boston collections that all specimens having genital tumescences on

1952 EARTHWORMS OF NEW HAMPSHIRE 3

XXX, XXxll and xxxiv (with none on xxxill) could also be distinguished from caliginosa by greater clitellar length and by larger segment number.

The presence of worms with arno/di characteristics in northern New Hampshire shows that those characteristics by which the species was recognized and defined are not merely those of a local variant in the Boston area. The occurrence of arnold? in many other states, as well as in Canada, may be anticipated.

REFERENCES Gatss, G. E. 1952. New species of earthworms from the Arnold Arboretum, Boston. Breviora, No. 9: 1-3. Smitu, F. 1917. North American earthworms of the family Lumbricidae in the col- lections of the United States National Museum. Proc. U. 8. Nat. Mus. 52: 157-182. SouTrHERN, R. 1910. A new species of enchytraeid worm from the White Mountains. Proc. Acad. Nat. Sei. Philadelphia, 62: 18-20.

i. @ 7 e. y o/

“i

oy yee s.

oe * ommaieds Ver aes Veni Van

; . ¢€ huagl’ Fupitre =i) Mii: Cee ites Greyaha ew Nasemile 1 5 ee Gaby WN goat WIA} ie aia ae et Tr

. ; ) tag o a a Poy i ceils f ire! Hp 4 ye » PPM Agee ty ‘ion reve ay de Aw i ea inti aa: tile Ay ae RM pe ea Ogee Nee aie a eee ihe ti.6 Sire. i a ae at “4 for unre ate eth

'

Al a ies) Pe Dhoni ; aca ne ag a 0 1 _ } is i et Cod Lee nas Vines) s* <3 OP: Sag js eat in] Ls 7 a ai iff

Res - ae ae. rd Lae wel pan hale 0s ar. si Aine tie ah Gt Koh an

eid. = Ban Sr * ek E iit 32 iw (cz aye

7 ae p ; : hale ~ ii are wep aeghh WER Allie Denn, feo te™ Tesi agit

. ‘ie Bs my ah aie May ik a

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. MarcH 20, 1953 NuMBER 11

CHARACTERS AND SYNONYMIES AMONG THE GENERA OF ANTS

Part I

By Wiuiiam L. Brown, JR.

Museum of Comparative Zoology Harvard University

Work now under way is aimed at a revision of the basic classification of the Formicidae down to generic and subgeneric levels. The present classifications are considered both unnatural and impractical, con- cealing as they do large numbers of generic and specific synonyms. I propose to differentiate genera only on the basis of discontinuities of a largely morphological kind, and it is hoped that by this method a useful and accurate key to the genera will become possible. Syn- onymies given here require discussion that is best set forth separately, so as not to clutter more nearly final, synoptic classifications and keys now being prepared.

Myrmecta Fabricius

Myrmecia Fabricius, 1804, Syst. Piez.: 423. Genotype: Myrmecia gulosa Fabricius, by designation of Emery, 1911.

Myrmecia (Promyrmecia) Emery, 1911, Gen. Ins., Fasc. 118: 18-19. Sub- genotype: Myrmecia aberrans Forel, by original designation. NEW SYNONYMY.

Myrmecia Clark, 1952, Formic. Australia, Melbourne, 1: 21.

Promyrmecia Clark, 1952, [bid., p. 119.

The references of Clark should be consulted for further synonymy. Clark has stoutly defended the separation of Myrmecia and Pro- myrmecia as distinct genera, first on the basis of ability or inability to jump of the species concerned, and later, when saltation proved to have numerous exceptions in bothdirections, on morphological grounds.

2 BREVIORA no. 11

Years of study of the myrmeciines by this author have produced in the 1952 reference (p. 20) a couplet summarizing the characters supposed to separate Myrmecia and Promyrmecia. The three charac- ters contrasted are: (1) fraction of length by which the antennal scape surpasses the occipital border, (2) distinctness of metanotum, and (3) size, as mirrored in total length. In the first character, it may be observed from the descriptions of species of Myrmecia given by Clark himself in the pages following the couplet, that no less than one-third of the 59 forms recognized disagree with the couplet, and disagree in the direction of Promyrmecia. In at least some cases, the scape pro- portions are exactly as stated in the couplet for the upper limit of Promyrmecia. The distinctness of the metanotum is subject to varying interpretation, but from any reasonable point of view, the generality of the indistinctness of the metanotum in Promyrmecia cannot be defended. In fact, Clark’s figures (op. cit.) in rapid survey are the best contradiction of his couplet that I can offer anyone without a large representation of material at hand for direct study. The “‘size” figures, even as cited in the couplet, are broadly overlapping and useless for the purpose to which they are put. Study of a good repre- sentative series of the species is convincing proof for me that size, as based on any measurable dimension or proportion of the worker or female so far utilized, is graded from the largest Myrmecia to the smallest Promyrmecia without a break.

It is entirely possible that study of the male genitalia, when a sufficient number of males becomes available, will demonstrate a set of cleavages that along with other characters will serve as disconti- nuities for the proper splitting of Myrmecia. Clark’s and other di- visions are here rejected for lack of evidence. It should be pointed out, in order to save possible future confusion, that Clark has syno- nymized Halmamyrmecia Wheeler with Promyrmecia, although he wrongly maintains Myrmecia nigrocincta Fr. Smith, its type, in the genus Myrmecia as he has defined it.

RHYTIDOPONERA Mayr

Ectatomma (Rhytidoponera) Mayr, 1862, Verh. zool.-bot. Ges. Wien 12: 731. Genotype: Ponera araneoides Le Guillou, by designation of Emery, 1911.

Rhytidoponera (Chalcoponera) Emery, 1897, Ann. Mus. Stor. Nat. Genova 38: 548. [Sub] genotype: Ponera metallica Fr. Smith, by designation of Emery, 1911. NEW SYNONYMY.

Rhytidoponera and Chalcoponera treated as distinct genera, Wheeler, 1922,

1953 ANT CHARACTERS AND SYNONYMIES 3

Bull. Amer. Mus. Nat. Hist. 45: 643-644. Clark, 1936, Mem. Nat. Mus., Melbourne, 9: 14-15.

The large genus Rhytidoponera in the present broad sense includes species of typical ectatommine characteristics, distinguished in having the inferior pronotal margins just in front of the fore coxae armed on each side with an acute tooth; the hind coxae are completely unarmed above. Distribution is primarily Australian and Papuasian, with outliers in the ‘“‘Wallacia”’ region and in the southern Philippines to the west, and in New Caledonia in the east.

The separation of Rhytidoponera from Chalcoponera, either generic or subgeneric, has been accepted from the time of Emery’s first di- vision of the few species then known. Rapid accretion of species in both groups has frequently led to puzzlement of authors trying to place new species in one group or the other, and this puzzlement has resulted in several published expressions of doubt accompanying specific descriptions.

The separation has been based on (a) proportions of certain antennal segments in worker and male, (b) development and pectination of the spurs of the middle and hind tibiae in the worker, (c) development of notaulices in the male, and (d) presence or absence of a normal winged female caste (Wheeler, loc. cit. 1922). If one applies these Emery- Wheeler key characters rigorously to a wide variety of species in Rhytidoponera and Chalcoponera, discrepancies are not long in ap- pearing.

The tibial spurs of the middle pair of legs are reduced and narrow (rarely absent) in all species of both genera examined for this work, and may be safely disregarded. The extremes of development of breadth and pectination of the posterior spurs are largely in accord with the conventional separation of the two groups, but exceptions are glaring, and gradual transition from one type to the other is evident among a restricted selection of eastern Australian species. In the species tenuis Forel, which has Chalcoponera antennal funiculi and which has been placed in Chalcoponera by all authors, the posterior tibial spurs are minute and show the strongly reduced pectination supposed to be characteristic of Rhytidoponera s. str. Conversely, certain Rhytidoponera s. str. species, such as R. malandensis Forel, have the posterior spurs large, broad, and strongly pectinate; in fact, R. malandensis has the spurs considerably more broadly pectinate than in any Chalcoponera species I have seen, including the genotype.

In the worker funiculi, the proportions of the ‘‘critical’’ segments

4 BREVIORA no. Il

are intergradient and fully ambiguous in a number of species, among them R. reticulata Forel, C. lamellinodis Santschi (paratypes), C. dubia Crawley, and all the species of the C. impressa Mayr complex. More ambiguous examples or outright contradictory species could be cited, but the above cases should serve to prove the point so far as the worker is concerned.

Concerning the male characters, a case similar to that for the workers can be drawn, but it will serve merely to focus on the New Caledonian fauna, consisting of a handful of forms in which the workers have been placed in Chalcoponera on the usual characters. The males of these species are, unfortunately for the generic division, endowed with the characters of Rhytidoponera s. str. Emery (1914, in Sarasin and Roux: Nova Caledonia, Zool. 1: 397) covers this situation adequately when he states: “The males of the New Caledonian species of Chalcoponera (Rhytid. fulgens, numeensis, atropurpurea and acupuncta) make an exception to the character that I have attributed to the subgenus concerning the structure of the antennae (Genera Insectorum, fase. 118, p. 39). For the present, I am incapable of distinguishing the males of the two subgenera of Rhytidoponera.” Apparently Wheeler overlooked this paragraph when he raised Chalco- ponera to generic rank in 1922, citing in his key the same discredited male characters. I have dissected the male genitalia of several species representing Chalcoponera and Rhytidoponera s. sir., and the prepa- rations fail to show differences except minor ones among the various species, without regard to the old division. The volsellae are uniformly much like those of Myrmica and the Dacetini in the subfamily Myrmi- cinae, and the other parts are on the usual formicid pattern.

It seems fairly obvious that most describers have pretty consistently ignored the formal characters in assigning new species to one group or the other; rather, assignment seems to have been made chiefly on the basis of size and habitus. On such a basis, I can make out not two, but several, species-groups of fairly distinctive relationships, but broadly intergradient one to the next. Of these, the impressa complex seems to have a combination of characters approaching nearest the hypothetical generalized Rhytidoponera. Disregarding certain aberrant species and complexes, such as the turneri group and the New Cale- donian stock, the development of the majority of species seems to have followed two lines: one toward large species of the type of mayri Emery, araneoides Le Guillou, and punctata Fr. Smith, and one toward the type of metallica Fr. Smith and victoriae André. In the farflung

1953 ANT CHARACTERS AND SYNONYMIES 5

regions penetrated by Rhytidoponera s. lat. on the Australian continent, these two broad adaptive types have met with relative success, while the more generalized connecting types have been restricted to favorable forested areas, largely in the mountains of eastern Australia. Thus, while the Rhytidoponera species inhabiting the deserts, scrubs, heaths and eucalypt woodlands of the great, arid, open regions of Australia may seem to be amenable to assortment into two distinct groups, it must be borne in mind that the more generalized forest-loving species, while less familiar to the observer, form an effective bridge joining these two groups.

Before leaving Rhytidoponera, a word is required on the females. It has been assumed that Chalcoponera species always produce normal winged females, and that Rhytidoponera s. str. species do not. In 1950, Clark (in litt.) indicated to me that he had found females of some sort, presumably differentiable from the workers, in the latter group. To date, he has published nothing concerning this find, so that it is fitting that the old assumption be continued until definite information appears in print. My own field investigations in Australia (1950-51) indicate that the R. impressa group (¢mpressa, chalybaea, splendida) normally produces a regular yearly crop of males and females in most mature nests during the first part of the dry season in each of the regions inhabited. Both sexes are winged before the nuptial flight, and ferti- lization and nest-founding are presumed to follow conditions more or less normal among the Ponerinae, as isolated females have been found dealated and inhabiting small cells in various situations in normal habitats. Outside the season for production of winged forms, only dealate females have been found in established nests in addition to the workers and such brood as occurs. When winged forms are found in a nest, these are always of both sexes so far as my observations go.

In species of the metallica group, a situation of a quite different sort is apparent. Among the common and widespread members of this group, such as metallica, tasmanicnsis, and their very close allies, winged or obviously dealate females are very rare or unknown. The victoriae group is similar in this respect, although I have found a dealate female in a mature nest of the species common at Kuranda, in North Queensland, and have also found a lone winged female of victoriae floundering in the damp sand at the surf edge at Seaford, on Port Phillip Bay, in Victoria. Of several thousand nests of victoriae, metallica, tasmaniensis and very close allies of these species that I have opened, not one (with the single noted exception above) has yielded

6 BREVIORA no. ll

a female of the normal type, either alate or dealate. Yet there are a few isolated specimens of normal females in these groups in the Mu- seum of Comparative Zoology collection, most of which seem to have been taken singly on the wing or otherwise.

While it is conceivable that the rarity of females in the nests of such species may reflect the methods of investigation, and not true absence of this caste, another circumstance makes this seem unlikely. I refer to the very widespread occurrence of winged males in nests of the same species at all seasons of the year, a phenomenon that stands out among my Australian observations even though I failed to take quantitative notes that would have made it much more convincing. These males are usually present in small numbers, and may be found in the upper as well as the lower chambers of a given nest. They are very active, and either run to hide quickly or else take to ready flight. A more limited number of observations on the large deserticolous members of Rhytidoponera s. str. indicates that males are to be found in their nests on a somewhat similar basis, though observations through several seasons will be needed to confirm this.

Among some other, more rare and distributionally restricted species, R. croesus Emery appears to have normal females most or all of the time, while R. aspera (Roger) is recorded as having this caste at least some of the time. For most of the species of the “inter-subgeneric”’ and aberrant groups, workers only are known at present, but these forms are so rare and so little-investigated that presence or absence of winged females cannot be assumed on any reasonable grounds. Under the circumstances as outlined, utilization of the presence or absence of a true female caste as a taxonomic character would seem entirely premature.

The situation in the female-less or female-rare species that have been fairly extensively investigated is, however, of considerable interest apart from taxonomic considerations. In two such species, I have seen a male seeking to enter an alien nest, apparently of the same form. In the spring, in southeastern Australia and the dry Northern Flinders Ranges of South Australia, at least, males of these forms are often seen hawking in rapid flight low over the ground in the manner of certain mutillid and thynnine males during the sexual search. Observations in 1951:

On the 12th of September, in a strip of open eucalypt woodland at Burwood, near Melbourne, Victoria, I noticed such a male in flight. on a cool, sunny afternoon. While following him, I suddenly saw him.

a

1953 ANT CHARACTERS AND SYNONYMIES

alight and disappear without hesitation into a small hole in the ground. This hole, when attacked with a trowel, proved to be the entrance to a nest of R. tasmaniensis, a nest containing, so far as I could dig on this occasion, workers and two males, the latter indistinguishable from males taken in nests of this species on other occasions. One of these males, found just below the entrance, must have been the male I saw entering just before I started to dig.

At Wilpena Pound, a sort of natural oasis in the arid Flinders Ranges of South Australia, the evening of November 25 brought an abundant flight of a large brown Rhytidoponera species, entirely males, to our camp pressure lanterns. Later that night, some rain fell and there were electrical storms on the adjacent peaks. On the next day, in an area of T'riodia grass within the Pound, I found a low, gravel-studded mound with wide, slit-like entrance, of the type made by certain arid- land species of Rhytidoponera and Camponotus. The nest was seen in the path taken on the way out, and was not excavated until returning toward camp, late in the afternoon when the sun had fallen very low. While I was bending over to deal the hard clay mound a first blow, a large male of Rhytidoponera came, flying in low over the ground from a distance, and landed directly on the lip of the entrance slit, which it immediately entered. This was surprising, as no sign of life had been detected around the mound at either time of inspection (the species concerned, probably R. mayri Emery, is, with its close relatives, a primarily crepuscular and nocturnal forager). The nest was immedi- ately attacked, and the male recovered quickly several inches down along the main entrance passage. Further digging secured only a few workers of R. mayri Emery.

While these two incidents, observed by chance, are scarcely to be considered definite proof of an established behavior pattern, the obser- vations made to date on various Rhytidoponera species suggest the following hypothesis, now being tested by Haskins (in litt.).

Rhytidoponera (s. lat.) varies by species and species-groups in presence, absence or rarity of a normal female caste, and varies corre- spondingly in behavior of and toward the males. In the species in which normal females are rare or absent, it may be assumed that deposition of fertile eggs is wholly or largely taken over by workers

1 The species described by Clark as R. stridulator and R. dixoni, along with some other forms‘ are supposedly distinguished from R. mayri by details of petiolar structure and body sculpture. Single nests of what I take to be mayri, however, show a wide range of variation in these same characters, and Clark has not made the differences sufficiently clear to convince me that his species are really distinct.

8 BREVIORA No. ll

or highly modified ergatoids. It is possible that apparatus for the retention of sperm isstrongly impaired in such hypothetical individuals; if so, then the need for constant refertilization would seem to call for the constant presence of consort males in the nest. Meager obser- vations seem to indicate that males sometimes, perhaps normally, leave the parent nest and enter another nest of the same species, where they may remain as guests for long periods. If observation and controlled experiment corroborates the above hypothesis, a new and extremely interesting kind of behavior will be added to the multiplicity of patterns known among the ants, and the origin of certain similar male habits among the Dorylinae may be easier to understand.

CENTROMYRMEX Mayr

Centromyrmex Mayr, 1866, Verh. Zool.-bot. Ges. Wien 16: 894. Genotype: Centromyrmex bohemant Mayr, monobasic.

Typhioteras Karawajew, 1925, Konowia 4: 128. Genotype: Typhloteras hamu- latum Karawajew, monobasic. NEW SYNONYMY.

Karawajew based his genus on a specimen with only a single, large pectinate spur on the posterior tibiae, and thereby carried it out to Emery’s Group III of the tribe Ponerini. This placement is only another example of the great faith formerly held by many authors in the constancy and taxonomic importance of the number and condition of the spurs of the middle and hind tibiae. In fact, the character in question is a very poor one upon which to base a classification, es- pecially in the Ponerini. If one reviews the situation in various Centromyrmex species, including Centromyrmexr hamulatus NEW COMBINATION, it is at once apparent that the number and con- dition of the spurs in question differ by species, and possibly even within species. Thus, either the middle, or the posterior, or even both pairs of tibiae may possess the extra lateral spur in a more or less rudimentary condition, while the degree of development of the medial spurs and their pectination is also variable from one species to the next.

In other characters, including the striking general habitus and the probably general termite-eating proclivities, the species here included in Centromyrmex form a very homogeneous and natural-seeming group. To break up this combination on the basis of spur characters, one must, as did Karawajew, go to extraordinary lengths in invoking ‘““convergence”’ as a possible explanation of the common resemblances; to be thorough in applying the same logic, one would have to distribute the few species involved among three or four genera. In any case, the

1953 ANT CHARACTERS AND SYNONYMIES 9

tibial apices of several of the species bear such a dense growth of heavy, spine-like setae that the identification of a spur rudiment of similar size becomes an academic exercise. In addition to the synonymy of Typhloteras with Centromyrmex, it becomes necessary to point out that the subtribe Centromyrmicini of Emery is a NEW SYNONYM of tribe Ponerini if the spur characters will not hold.

PRISTOMYRMEX Mayr

Pristomyrmex Mayr, 1866, Verh. Zool.-bot. Ges. Wien 16: 903. Genotype: Pristomyrmex pungens Mayr, monobasic.

Odontomyrmex André, 1905, Rev. Ent. Caen 24: 207.

Hylidris Weber, 1941, Ann. Ent. Soc. Amer. 34: 184, 190. Genotype: Hylidris myerst Weber, monobasic. NEW SYNONYMY.

Hylidris defended, Weber, 1952, Amer. Mus. Novit. 1584: 15-22.

When, in 1941, Weber first described Hylidris, he did so without realizing that it might be closely related to another genus like Pristo- myrmex. By 1952, he has realized this relationship and is at some pains to mark out its boundaries. His new approach to the situation lies in segregating the African species of Pristomyrmex from the Indo- Australian ones; the former group, rallied about H. myers? as genotype, is given the name Hylidris. He is vague about what genus the Indo- Australian species are to belong to, but these are presumably to remain in Pristomyrmex. The characters of Hylidris are cited briefly, but are not contrasted with those of Pristomyrmea s. str. in any direct state- ment. Instead, Weber gives briefly his views on generic limits, which he feels should be arbitrarily drawn in some cases. (For my contrasting opinion, see the introduction to this paper.)

Summing up, it may be said that Weber’s division of Pristomyrmex is based upon characterization of only one of the resultant groups, and no assurance is given that the characters are exclusive to that group. In point of fact, these characters are not exclusive to the African group. Since Weber claims to have examined the Indo-Australian species, at least in part, it may be wondered that he did not note this fact for himself; perhaps he did note it, and offered the generic-limit discussion as a defense against protests he felt would be lodged against Hylidris.

I cannot follow Weber’s generic split, which I regard as wholly arbitrary. Significantly, Weber fails to mention the old synonym Odontomyrmex, a name put forward by André for a species that appears to have the chief definitive characters of Hylidris. Mann (1919, Bull. Mus. Comp. Zool. 63: 341) delivered the coup de grace to Odonto-

10 BREVIORA no. ll

myrmex when he noted two specimens of his Pristomyrmex obesus melanoticus having a prominent tooth on one side of the pronotum, but none on the other side. We do not need to appeal to abnormal specimens in this case, however, as the known, normal specimens of the Indo-Australian and African regions form a tightly intergradient series with respect to the development of the posterior propodeal and anterior alitruncal pairs of teeth and the smooth to foveate-reticulate sculpture. This series, in my opinion, does not even split into re- spectable species-groups on the basis of the known characters, let alone genera or subgenera. Series of several undescribed species in the Museum of Comparative Zoology and J. W. Chapman collections serve only to fortify this opinion.

GAUROMYRMEX Menozzi

Gauromyrmex Menozzi, 1933, Natuurhist. Maandblad 22: 146. Genotype:

Gauromyrmex bengkalist Menozzi, monobasic. Solenomyrma Karawajew, 1935, Treubia 15: 103. Genotype: Solenomyrma

acanthina Karawajew, monobasic. NEW SYNONYMY. Acalama M. R. Smith, 1948, Jour. N. Y. Ent. Soc. 56: 205-207. Genotype:

Acalama donisthorpet M. R. Smith, monobasic. NEW SYNONYMY.

This genus is very doubtfully distinct from Vollenhovia Mayr, from which it may at present be distinguished by means of the 11-segmented antennae and the bidentate propodeum, vs. 12-segmented antennae and unarmed propodeum in Vollenhovia. The characters are very weak ones, and may be compromised in species in this complex that are presently not available to me. Vollenhovia emeryi Wheeler, with 12-segmented antennae, has minute propodeal teeth, and the median funicular segments are so reduced in length as to be virtually obsolete; the step to Gauromyrmex from this species is a very short one. The amber species Vollenhovia beyrichi (Mayr), if properly placed by Wheeler, would be intermediate in the critical characters. Types of Gauromyrmex bengkalisi and Acalama donisthorpet were

examined, and are considered specifically distinct, but not generically so. Specific synonymy is formalized as follows:

GAUROMYRMEX ACANTHINUS (Karawajew) new combination Solenomyrma acanthina Karawajew, 1935, Treubia 15: 103-104, fig. 23, worker. Acalama donisthorpei M. R. Smith, 1948, Journ. N. Y. Ent. Soc. 56: 207-208, figs. 1, 2, worker. NEW SYNONYMY. Types of Smith’s species and Gauromyrmex bengkalist have been examined through the kindness of Dr. Smith. The former species has

1953 ANT CHARACTERS AND SYNONYMIES 1

been compared with Karawajew’s description and figure, and also with the West Chinese series taken by myself, and mentioned by Smith (loc. cit., p. 206). The comparison gives the distinct impression that one is dealing with a single variable species. Variation is chiefly in size (slight), distinctness and acuteness of propodeal teeth, and depth of pigmentation, but the present evidence does not warrant subspecific distinction in my opinion. This ant appears to be rather common in India and China, and I believe that I have seen it on several occasions in Bengal Province nesting under loose bark, though the actual specimens were lost in a wartime shipment. In life, the distinctive point of recognition lies in the extremely depressed ap- pearance of the ant, even when moving about. It appears to the naked eye somewhat like a flattened Leptothorax, and its attachment to tree trunks and plant cavities may be correlated with the habitus.

At the time of Dr. Smith’s investigation of this insect, he sent specimens to me, and I failed to recognize it as any described species. The published descriptions of Menozzi and Karawajew came to light later. The entire case of these synonymous genera should prove my frequently-held point that the classification of the Myrmicinae is long overdue for complete revision.

Tribe AMBLYOPONINI

Onychomyrmicini Ashmead, 1905, Canad. Ent. 37: 382.

Examblyoponini Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15: 401.

Reneini Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 183. NEW SYNONYMY.

PRIONOPELTA Mayr

Prionopelta Mayr, 1866, Sitzb. Akad. Wiss. Wien 53: 503. Genotype: Pri- onopelia punctulata Mayr, monobasic.

Ponera Fr. Smith (partim), 1860, Journ. Proc. Linn. Soc. London, Zool. 4 (suppl.): 105, nec Latreille.

Examblyopone Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15: 401. Geno- type: Examblyopone churchilli Donisthorpe, monobasic: vide infra.

Renea Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 183. Genotype: Renea testacea Donisthorpe, monobasic. (Nec Renea Nevill, 1880, in Mollusca.) NEW SYNONYMY. Vide infra.

Incredible as it may-seem, Donisthorpe based his two new genera and two new tribes on two new species, both of which are synonymous with Prionopelta majuscula Emery. The same author had already proposed a new name for a synonym of the same species, so he has

2 BREVIORA No. ll

achieved the unequalled feat of proposing for the same species three new specific, two new generic and two new tribal names. The syn- onymy of Prionopelta majuscula follows:

PRIONOPELTA MAJUSCULA Emery

Ponera simillima Fr. Smith, 1860, Journ. Proc. Linn. Soc. London, Zool. 4 (suppl.): 105, “worker,” 2, nec Fr. Smith, 1860, op. cit., p. 104. NEW SYNONYMY.

?Rhopalopone simillima, Emery, 1900, Term. Fiizetek 23: 311. 1911, Gen. Ins. 118: 35.

Prionopelta majuscula Emery, 1897, Term. Fiizetek 20: 595-596, worker, 9. Brown, 1951, Bull. Brooklyn Ent. Soc. 46: 102, Examblyopone churchilla synonymized.

Prionopelta poultoni Donisthorpe, 1952, Ann. Mag. Nat. Hist. (10) 10: 462, nom. pro Ponera simillima Fr. Smith, I. NEW SYNONYMY.

Examblyopone churchilli Donisthorpe, 1949, Ann. Mag. Nat. Hist. (11) 15: 401-402, 2.

Renea testacea Donisthorpe, 1947, Ann. Mag. Nat. Hist. (11) 14: 183-186, fig., worker, o’; pp. 590-591, 9. NEW SYNONYMY.

In his original description of Ponera simillima I, Frederick Smith describes a ‘“‘worker”’ and also gives characters purporting to be those of a winged female. In his treatment of 1932, Donisthorpe mentions as Smith’s type “1 dealated @” from “Dor.” [Dory, A. R. Wallace]. It is by now fairly well known that some of the Dory insect material, at least among the coleopterous collections in the British Museum, is suspected to be from other East Indian localities. Smith’s description fits Prionopelta mayuscula better than it does either of the other two Prionopelta species known from the Indo-Papuasian area (P. kraepelina Forel and P. opaca Emery), so there is no reason to question this particular record for a widespread New Guinea species. The as- sumption must be made, of course, that Donisthorpe’s 1932 assignment to Prionopelta was correct; there is every reason to believe that it was so.

In 1951, I determined that a paratype female of Examblyopone churchilli was synonymous with P. majuscula and returned the type to Dr. E. S. Ross at the California Academy of Sciences. Dr. Ross was the original collector (at Maffin Bay, Dutch New Guinea) of both E. churchilli and Renea testacea, and he has recently sent me workers and males from the type series of the latter species. The workers fit Emery’s diagnosis of P. majuscula very neatly, except for the usual small size difference resulting from Emery’s habitual under-

1953 ANT CHARACTERS AND SYNONYMIES 13

measurement. This being the case, I asked Dr. Ross to compare the female specimens assigned by Donisthorpe to R. testacea with the female holotype (defective) of FE. churchill. He has replied (in litt.) that the correspondence of form, etc. is as good as can be expected, allowing for the missing parts of the EL. churchilli type. The R. testacea workers differ as expected from authentic workers of Prionopelta opaca and P. kraepelini in the Museum of Comparative Zoology and match the differences from P. opaca listed by Emery in the original de- scription of P. majuscula.

P. majuscula is larger than the other two Indo-Papuasian species, and its worker has the dorsum of the head and alitrunk very definitely shining, with minute, spaced punctulation. The female is considerably larger and darker than the worker, and has stronger punctulation; the head is darker than the rest of the body. Both P. kraepelini and P. opaca are not only smaller, but also there is only a slight difference between the worker and female stature in these species; the dorsum of the head is densely and more coarsely punctulate and opaque, especially in opaca. The worker and female of kraepelini and the worker of majuscula are pale to bright yellow in color, while the females of majuscula and opaca and the worker of opaca are darker, ranging from ferrugineous brown to blackish-brown. P.opaca and P. majuscula appear to be widespread on New Guinea and neighboring islands, while P. kraepelini is a more westerly, Indomalayan species that has spread into the Pacific as a tramp and has reached the Philippines in the north at Dumaguete, Negros Oriental: several series (J. W. Chapman and D. Empeso).

af we eee Wits (i

. e . an? « nya { rg if 4 vs ¥! , : “4 7 Mies \ Pe ua hth { a ' we ay @

{ nth ; : , sf 5 7 + i : aie Ag. i " oe 7a) s/s z fb; 13 ey ps fa re

| eed [ uaigh i \e: us ‘ie adi p “ly a ma cog WP yar £0 ha aa Dis. fue tis) r+ Ge ieenber!pid of eel i yet Port ff Tv (re a aia Piet are 4 caer 4 ° rer a8 HD ae ee!

=

oye yh - hia Cu? thi 3

hoi bh) Phen havi) 4 Pr j Leip fd EN ar

«i A Luria JM Sk ee

} : on ial’ s eg i peal 8 i’ ‘ey ee al bije Bible ' oe

oe ee,

a es +4 = Ai Lee a > A pias Gide ib seme, © “UT PT @ 7 air ny. b rn resin ery Lond Da | ui : Fg i , i = | a ce oy : Le

BREVIORA

Museum of Comparative Zoology

CaMBRIDGE, Mass. APRIL 23, 1953 NuMBER 12

A CAVE FAUNULE FROM WESTERN PUERTO RICO WITH A DISCUSSION OF THE GENUS ISOLOBODON

By Tuomas E. ReyNoLps Zoology Department, Massachusetts College of Optometry

Karu F. KoopMAN Biology Department, Queens College

ERNEST E. WILLIAMS

Harvard University

In spite of the diligent work of H. E. Anthony nearly 40 years ago, the vertebrate paleontology of Puerto Rico must still be described as incompletely known. Anthony’s labors have not been followed up in any systematic fashion by other investigators, and much therefore remains to be done.

The present paper places on record a collection from an area of the island relatively neglected by Anthony the extreme western portion of Puerto Rico. The collection, though small and apparently repre- senting a fauna of no high antiquity, permits new locality records for certain forms and includes a very interesting specimen of the genus Isolobodon which has compelled us to inquire into the variability of that genus and into the propriety of taxonomic subdivisions of it.

The materials here recorded were obtained by the senior author during the fall and winter of 1948-49 at the mouth of the Cueva Monte Grande between Mayagiiez and Cabo Rojo (Distrito de Mayagiiez, Pueblo de San German, Barrio de Monte Grande). The specimens have been deposited in the Museum of Comparative Zoology.

There were several levels in the deposit, and part of the deposit was kitchen midden, but no record was kept of the level at which specific material was found.

2 BREVIORA No. 12

We cite below only the mammalian component of the faunule. There are, however, also fish, bird, turtle, snake and lizard bones. The lizard jaws have been identified by Rodolfo Ruibal and Max K. Hecht as all belonging to Anolis cuvieri, the living giant anole of Puerto Rico. No attempt was made to identify limb bones in any group. In the case of the mammals we take this opportunity to cite also localities not previously published but represented in the col- lections of the Museum of Comparative Zoology (MCZ) or the American Museum of Natural History (AMNH).

The Monte Grande Faunule

INSECTIVORA NESOPHONTES EDITHAE

Two mandibular halves.

This large extinct insectivore of Puerto Rico has previously been reported from Hacienda Jobo, Morovis and Utuado. The MCZ col- lections show it also from Ciales, Manati and Rio Piedras.

CHIROPTERA NOCcTILIO LEPORINUS MASTIVUS

One complete skull. Two mandibles.

Though recorded by Anthony (1926) only from Old Loiza, this form was collected by Anthony and Goodwin at Vega Baja (AMNH speci- men) during a 1926 West Indian expedition which has never been fully reported in the literature. The present is therefore the third record for this large fish-eating bat in Puerto Rico.

CHILONYCTERIS PARNELLII PORTORICENSIS

One skull.

The specimen so referred seems somewhat small for the species, but it is much too large to be the small Puerto Rican species of the genus (C. fuliginosa inflata). It is probably a young specimen.

This subspecies seems to be relatively rare in collections, only 11 specimens having been previously obtained, representing four locali- ties: Cayey, Morovis, Pueblo Viejo and Trujillo Alto.

MoNOPHYLLUS PORTORICENSIS

One skull.

1953 A PUERTO RICAN CAVE FAUNULE 3

This species was recorded by Anthony (1926) from five localities: Bayamon, Cayey, Morovis, Pueblo Viejo and Trujillo Alto.

BRACHYPHYLLA CAVERNARUM Four more or less complete rostra. Six complete mandibles. Seven half-mandibles. Recorded by Anthony from six localities: Cayey, Comerio, Corozal, Morovis, Pueblo Viejo and Trujillo Alto.

ARTRBEUS JAMAICENSIS JAMAICENSIS

One skull. Six mandibles. This very common form was obtained or reported by Anthony from 14 localities.

EPTESICUS FUSCUS WETMOREI

One mandible.

Anthony (1926) reported this species from four localities (Maricao, Morovis, Pueblo Viejo and San German). He had collected it also at Trujillo Alto (AMNH specimen) during the 1916 expedition but somehow overlooked this record in his publication.

RODENTIA RaTTUS sp.

Two mandibles. One half-skull. Rattus, as always in the Western Hemisphere, indicates that at least part of the Monte Grande faunule was post-Columbian in age.

ELASMODONTOMYS OBLI QUUS

A fragment of an upper incisor and a third right upper molar appear to represent this species.

This large extinct rodent has previously been reported only from Ciales, Morovis and Utuado. The MCZ collections add it also from Rio Piedras.

ISOLOBODON PORTORICENSIS

Nineteen skull fragments. Fifty-five half- or partial mandibles.

The bulk of the material from Cueva Monte Grande is of a medium sized hypsodont rodent. The enamel folds of the molars are nowhere completely separated to form laminae. The lateral surfaces of these

4 BREVIORA No. 12

teeth have the pattern of striations characteristic of the genera or subgenera [solobodon and Aphaetreus. Even a cursory inspection of the molar pattern, however, rules out A phaetreus (Miller 1929a, pl. 2). The specimens then seem referable to [solobodon. The only species of that genus which has previously been recognized on Puerto Rico is Isolobodon portoricensis, and almost all of the specimens agree well with this form in both size and pattern and may immediately be placed as representatives of this species

Specimens of [solobodon portoricensis from Monte Grande afford a new record of locality but less clearly than the other forms recorded above, since Anthony has already reported this species from nearby Cabo Rojo as well as from Ciales, Manati, Salina, San German and Utuado. (The MCZ adds Saliche and Aguirri.)

Four of the Monte Grande mandibles, however, are not immediately classifiable as I. portoricensis and require special attention. All are much smaller in size than fully adult J. portoricensis. Two are obvi- ously immature. One of these is edentulous; the other has completely unworn crowns to the teeth, which are, therefore, not readily compared with the worn adult molar pattern. However, by breaking the ventral surface of the mandible, it has been possible to discover from under- neath the pattern of the enamel ridges which will be eventually re- vealed by growth. The enamel pattern so revealed is exactly that of typical I. portoricensis.

The third small mandible is larger than the other two and although considerably smaller than adult J. portoricensis shows no morphological evidences of immaturity (last molar not fully erupted, unworn molar pattern, absence of bony shelf behind the last molar). The fourth mandible is similar to the third in size but is edentulous.

Subadult mandibles equivalent in size to these last two Monte Grande specimens are known from many localities elsewhere in Puerto Rico and in Mona and the Virgin Islands and are always indis- tinguishable from the adults except in size. It is, therefore, clear that small size is in itself no bar to considering the four questionable Monte Grande mandibles as I. portoricensis, and we refer the two obviously immature specimens to that species without further discussion. The fourth mandible, lacking teeth, manifests no distinctive characters and may tentatively be referred to the same species.

The third questionable specimen differs, however, in molar pattern from all the numerous specimens of Jsolobodon with which it has been

1953 A PUERTO RICAN CAVE FAUNULE 5

compared. In any [solobodon three important enamel folds are evident, a single main fold from the labial side and two counterfolds from the lingual side. In the Monte Grande specimen in question the anterior counterfold of the first molar shows a distinct constriction about midway along its length and, correlated with this, there is a marked broadening of the lingual lobe anterior to the counterfold. No similar constriction is present on the second and third molars; the anteriormost lingual fold may have been broadened on both these teeth, but fractures in the critical regions make it impossible to confirm this. No con- striction comparable to that on the first molar, nor any broadening of the anterior lingual lobe at all similar to that seen on the first molar and probable on the second and third molars of the Monte Grande specimen, have been found in the more than 250 Isolobodon portori- censis mandibles examined. This third, small, Monte Grande mandible may, therefore, represent an exceedingly rare species, possibly reaching a smaller adult size, but closely related to Isolobodon portoricensis. On the other hand, it may represent a rare mutant condition in a subadult individual of J. portoricensis. Of these two alternatives we consider the second the more economical hypothesis, since we have been unable to find any other consistent differences between the third, small, Monte Grande mandible and the mass of compared J. porto- ricensis. We feel that the counterfold constriction might have been caused by a single mutant gene. It is interesting that this apparently aberrant individual probably never reached full adulthood.

In the course of this investigation, and while small size as a possible species character was still in question, the Monte Grande mandible and other Jsolobodon material from Puerto Rico was carefully com- pared with material identified as [solobodon levir from Hispaniola. It is worthwhile to consider here the status of the latter species, but before doing so a brief resume of its taxonomic history seems in order.

I. levir was first described by Miller (1922) as a distinct genus and species, [thyodontia levir, from two molars found fossil at St. Michel. He did this believing that these isolated teeth were lower molars. Twelve mandibles from the same locality were identified as [solobodon portoricensis without comment. In 1929, after studying much more material from the same locality, he realized that the two molars of “Tthyodontia’’ were actually upper molars of Jsolobodon. In an attempt to save I. levir from complete synonymy, he noted that all St. Michel specimens were smaller than J/solobodon from other Hispaniolan

6 BREVIORA No. 12

(kitchen midden) localities. The latter, in turn, were indistinguishable from Puerto Rican J. portoricensis. Although he could find no morpho- logical characters to distinguish them, he nevertheless separated the St. Michel specimens as a distinct species on size alone. He pointed out, however, that Hispaniolan I. portoricensis came from kitchen middens, whereas J. /evir came from ow] pellet deposits. As he indi- cated, smaller species and individuals are usually found more fre- quently in owl deposits than in kitchen middens, but he believed that the presence in St. Michel caves of relatively large mandibles of the related genus or subgenus A phaetreus ruled out the suggestion that in the two types of Hispaniolan Jsolobodon he was dealing with large- and small-sized samples from the same population. In later papers (Miller 1929b, 1930) he records [. levir from several other Hispaniolan locali- ties (San Gabriel, Monte Cristi, Constanza, Trujin), in Monte Cristi actually in association with I. portoricensis.

After extensive comparisons of I. portoricensis from Puerto Rico and surrounding islands (AMNH material) with J. levir from Hispa- niola (MCZ material from Fort Liberté as well as material from the United States National Museum, especially mandibles from St. Michel, Monte Cristi and Anadel, but also skulls from these and other localities), we agree with Miller that no morphological characters are to be found to separate the two species. We fail, however, to see any clearcut size difference, since the smaller species ‘“‘levir’’ appears to us to grade into the larger species portoricensis. “‘I. levir’? mandibles from Hispaniola can be matched by equally small mandibles from Puerto Rico. We believe also that in some cases Miller introduced a false dichotomy into his measurements by comparing the largest speci- mens identified as J. levir with the largest I. portoricensis: it is unlikely that a bimodal distribution would have resulted in the one case of actual association of the two forms if measurements of all measurable specimens had been used. For these reasons we consider [thyodontia levir Miller a complete synonym of [solobodon portoricensis J. A. Allen. Isolobodon thus emerges as a monotypic genus or subgenus, undiffer- entiated from Hispaniola through Mona and Puerto Rico to the Virgin Islands. As has already been pointed out (Miller, 1918) this wide uniform distribution may well have been brought about by human transport. If that be true just what was the original range of the genus before the coming of man to the Antilles. This would seem impossible to determine now.

1953 A PUERTO RICAN CAVE FAUNULE of

Acknowledgments. We are indebted to the Departments of Mammals of the Museum of Comparative Zoology, the American Museum of Natural History, and the United States National Museum for the privilege of examining and comparing specimens.

REFERENCES

ALLEN, J. A. 1916. An extinct octodont from the island of Porto Rico, West Indies. Ann. N. Y. Acad. Sci., vol. 27, pp. 17-22.

AnTuony, H. E. 1926. Mammals of Porto Rico, living and extinct. N. Y. Acad. Sci., Scientific Survey of Porto Rico and the Virgin Islands, vol. 9, pp. 1-238.

MILER, G. 8. 1918. Mammals and reptiles collected by Theodoor de Booy in the Virgin Islands. Proc. U. 8. Nat. Mus., vol. 54, pp. 507-508.

1922. Remains of mammals from caves in the Republic of Haiti. Smithsonian Misc. Coll., vol. 74, no. 3, pp. 3-5.

1929a. A second collection of mammals from caves near St. Michel, Haiti. Smithsonian Misc. Coll., vol. 81, no. 9, pp. 14-18.

1929b. Mammals eaten by Indians, Owls, and Spaniards in the coast region of the Dominican Republic. Smithsonian Misc. Coll., vol. 82, no. 5, pp. 6-8.

1930. Three small collections of mammals from Hispaniola. Smithsonian Misc. Coll., vol. 82, no. 15, pp. 4-8.

8 BREVIORA NO. 12

PLATE

Crown views of mandibles of Isolobodon. A. “‘Isolobodon levir’’. U.S.N.M. No. 255874 from Monte Cristi, Dominican Republic. B. Isolobodon portori- censis. A.M.N.H. ‘0.1’? Utuado, Puerto Rico. C. Aberrant Monte Grande mandible. M.C.Z. Cueva Monte Grande, Puerto Rico. (These specimens, all of about the same size, have been selected to show the extremes of variability of molar pattern.) 5 x natural size.

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. APRIL 23, 1953 NuMBER 13

FOSSILS AND THE DISTRIBUTION OF CEE EMID Wiens

1. ““Hydraspis’’ leithii (Carter) in the Eocene of India is a Pelomedusid

By Ernest WILLIAMS

For SO years a fossil turtle from the Eocene of India has been referred to the Recent South American chelyid genus [/ydraspis (not of Bell 1828, correctly called Phrynops according to Stejneger 1909, and Lindholm, 1929). In spite of the zoogeographical interest of this record the evidence for this assignment has not previously been re- examined critically.

In view of the prevalent idea that the family reached Australia from Asia, the occurrence of a chelvid in the Eocene of India would be neither surprising nor unwelcome. Reference, however, of an Eocene Indian fossil to a modern South American genus is more suspect and obviously calls for re-investigation. Examination of the original de- scription leads to a verdict unfavorable to both the generic and the family assignment.

The fossil in question was brought to scientific attention just over 100 years ago. In 1852 H. J. Carter in a study of the geology of the is- land of Bombay described as 7T’estudo leithii* the remains of a small turtle from the Intertrappean beds. Carter was not deceived as to the affinities of the form; he was using the generic name T'estudo in a Linnaean sense and explicitly stated that he regarded his fossil as close to Sternothaerus” (= Pelusios). He published two good plates giving a reconstruction of the fossil made from nine partial specimens. These

Testudo leithii Carter 1852 preoccupies Testudo leithii Giinther 1869 for the tortoise of

Egypt. The available and correct name of the latter then appears to be Testudo kleinmanmni Lortet 1883

2 BREVIORA NOwlS

plates and his text description are the basis of all subsequent discussion. The type material cannot now be located in India and has never been restudied.

On the basis of the 1852 description, Gray in 1871 reassigned the Indian fossil. He remarked: ‘‘The description and figure of the cara- pace induce me to believe that the fossil is nearly allied to some of our existing South American species of the restricted genus Hydraspis; and the remains of the head, which are unfortunately imperfect, lead to the same conclusion....’’ This determination has been very generally copied, in spite of the zoogeographical anomaly upon which Gray himself commented.

I find it necessary to disagree with Gray on the basis of the figured morphology of both shell and skull. Plates 1 and 2 are reproductions of Carter’s plates X and XI with a few inessential modifications for clarity.

According to Carter’s plate X (though the area is given in dotted lines only) and according to his express statement in the text there is no nuchal seute in the Indian fossil. Lydekker (1S89b, p. 170) stated: “The omission of a nuchal shield in the restoration of the anterior border of the carapace is probably incorrect.”” Perhaps, however, Lydekker made this statement only on the ground that if the form were []ydraspis it should possess a nuchal scute. Absence of a nuchal scute would rule out all Recent genera of Chelvidae except Chelodina and Emydura (in both of which the scute may be present or absent) and Elseya (in which it is regularly absent). The latter genera are all natives of the Australian region.

In Carter’s fossil the first vertebral is much smaller than the second vertebral. This precise condition is not met with in the living C helyi- dae. In most Recent South American forms including most cf the species of Hydraspis (= Phrynops) the first vertebral] is on the contrary much larger than the second. There is, it is true, an appreach to the condition of the fossil in the Australian genera Ek mydura, Pscudemiydura and Elseya and in the South American Hydraspis tubcrosa (specimers in the British Museum) and perhaps in some specimers of the Scuth American genus Hydromdusa, but in the latter only if the anterior median scute is interpreted as a nuchal withdrawn from the margin rather than as a transversely divided first vertebral. In none of these, however, is the first vertebral as much smaller than the second as it is in the fossil.

In the Indian fossil the gulars are small, and the very bread inter-

1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES i)

gular extends from the anterior margin to the humero-pectora! sulcus, separating the humerals in the midline. This pattern of scutes on the anterior plastral lobe is different from any which is known in the modern Chelyidae. In all the genera except Chelodina, Pscudemydura, and rarely in Emydura (Siebenrock, 1907) the humerals meet in the midline for a significant distance behind the intergular. The intergular also is rarely as broad as in the fossil. In Chelodina the intergular is very large and separates the humerals, indeed extending deep into the area of the pectorals, and, except in C. intergularis Fry, it does not reach the anterior plastral margin, the gulars meeting in front of it. In Pseudemydura (Siebenrock, 1907) the intergular is like that of the Indian fossil in its breadth but as in Chelodina dips deeply between the pectorals. The gular-intergular pattern in the exceptional Emydura subglobosa in which Siebenrock found the intergular separating the humerals is also quite unlike that of the Indian fossil, the intergular being narrow and of quite different shape.

The feeble xiphiplastral notch is another feature in which the fossil differs from Hydraspis and other chelyids except the forms of the Australian region and Batrachemys.

The shell, therefore, is not a good match for that of any known genus of chelyid. It is perhaps most like those of the Australian genera but differs from all of these in significant details, for example, in the presence of neurals, which are lacking in all the Australian genera.

The skull and mandible in their turn provide conclusive evidence against chelyid affinities. The mandible, although incomplete, is stouter and broader than in any known chelyid. The symphysis must have been long, in strong contrast to the condition in chelyids. The skull, shown by Carter only in dorsal view, is radically different from that of any chelyid. The skull roof has undergone emargination from behind as in the Pelomedusidae or most Cryptodira. As a result, the parieto-squamosal arch is absent, but a jugal-quadratojugal bar is present. In the Chelyidae and in no other turtles the skull is emargi- nated from the ventral margin only, and a parieto-squamosal con- nection is (except in Chelodina) always preserved. In chelyids the quadratojugal and the bar of which it was a part are always absent. These are as crucial and clearcut differences as it is possible to obtain between skulls of turtles. Carter’s fossil cannot be a chelyid.

It is most probable that it is a pelomedusid. Reference to this family wou'd, it will be recalled, be a return to the opinion of the original dese-iber, wh» thought the fossil was closest to the African

4 BREVIORA NOn Ie

pelomedusid genus Pelusios. The characters of the shell fit such a reference extremely well. The nuchal scute is almost always absent in pelomedusids. The first vertebral is almost always smaller than the second (exception in Palaeaspis Gray, and sometimes in Pelusios). An intergular separating the humerals is found in Stereogenys podocne- motdes and in Elochelys perfecta as well as in occasional individuals of Podocnemis expansa. The xiphiplastral notch is very variable in pelomedusids. Vertebral shields 2 to 4 of the Indian fossil have strikingly convex anterolateral borders, conspicuously concave postero- lateral borders, as in some Recent Podocnemis and a number of fossil pelomedusids.

A final feature which, as described and figured by Carter, is anoma- lous, may be clinching proof of the pelomedusid affinity of this form. Gray mentioned that Carter’s form was “peculiar also for the under- side of the marginal opposite the (pectoral-humeral) suture being rather broader than the rest and angular on the inner edge, which I have not seen in any of the Recent species.” If we have to do here with sulci between scutes, the situation is indeed peculiar and unique, but it is noteworthy that the lateral marginal scute boundaries are represented by dotted lines in Carter’s original reconstruction (solid lines in plate 2 here) except for the anomalous ‘‘marginal”’ in question. It is noteworthy also that the ventral view of the lateral marginals in Carter’s plates does not match the dorsal view of the same marginals. It is evident that Carter was not certain of the exact scute boundaries here, and it is possible that he has figured as the anomalous “‘mar- ginal” the sutures between bones rather than the sulci between scutes. The lines drawn solidly by Carter in this region are in nearly the right position and have the right aspect to represent in their lateral portions the sutures bounding small mesoplastra and medially the hyo-hypoplastral suture. This interpretation is the more probable because Carter’s figure is a reconstruction from nine specimens, one of which may have shown the sutures in this critical region and not the sulci. If small laterally placed mesoplastra were present, this fact would definitely place the Indian form in the Pelomedusinae of Zan- gerl (1948) with which, on the basis of other resemblances in its shell, it is most plausibly linked.

Carter’s turtle is, therefore, most probably a new genus of pelome- dusine. I cannot distinguish it from all previously proposed genera of pelomedusines because not all of these are themselves well-delimited. Thus I cannot distinguish it from Rosasia (Carrington da Costa 1940)

1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES 5

because at present that genus does not seem to be definable. (On its known characters Rosasia might be a synonym of any one of several genera. The carapacial shield, which alone is known, is not sufficiently diagnostic.) I cannot distinguish the Indian fossil from Dacochelys (Lydekker, 1889a) because there are no comparable parts, that genus having been founded on a mandibular symphysis, a part that is missing in Carter’s fossil. There is also no evidence that Dacochelys is a pelome- dusid.* I distinguish Carter’s form, with some hesitation, from Elochelys Nopcsa 1931 because the gular-intergular pattern is not quite that of E. perfecta, the type of the genus, and I am not persuaded that the other species referred by Nopcsa to that genus (/. major) belongs there. The critical feature of Elochelys also, the absence of a suprapygal, is not determinable in Carter’s form. A possibly trivial feature, the feeble xiphiplastral notch, distinguishes the Indian fossil from Stereogeni's podocnemoides (Reinach, 1903), but with Schmidt (1940) I do not believe that podoenemoides belongs to the genus Stereogenys. Podocnemoides and leithii may indeed belong to the same genus, but that genus is then unnamed.

From the better defined genera of the Pelomedusidae the Indian form is distinguished by the following combination of characters:

(CARTEREMYS, new genus

Type. Testudo leithii Carter 1852.

Diagnosis. Skull roof much emarginate from behind; opisthotics prolonged backwards in sharp crests; mandible with a moderately long symphysis; nuchal absent; first vertebral not divided transversely ; intergular large and very broad, separating humerals; xiphiplastral notch narrow and shallow; pubic and ischial sears distinctive in shape and position.

* An ingenious device by which Lydekker avoided the possibility of a change in the trivial name of Dacochelys has been the source of confusion in regard to this point. The type of the genus is Dacochelys delabechei Lydekker 1889, but Lydekker considered his form probably synonymous (largely on size alone) with Emys conybeariti Owen and therefore (according to Lydekker and Boulenger 1887) with Emys delabechei Bell. No name change is necessary 7f this synonymy is correct and, since the shell of Emys conybearii shows small lateral mesoplastra (Lydekker and Boulenger 1887), Dacochelys delabechei is then a pelomedusine. But the only valid physical type of Dacochelys (the type of the species upon which the genus is based) is the very peculiar mandibular symphysis, which I regard as quite impossible to assign to family. Lydekker’s device has most unfortunately brought the name Dacochelys into the literature (for example in Zangerl, 1948) as a pelomedusine which it may be but which it certainly cannot at present be proved to be.

It should be mentioned that if Dacochelys delabechei were in fact a synonym of EF. delabechei Bell as Lydekker assumed, and if E. conybearii Owen were a synonym of E. delabechei Bell as Lydekker also assumed, Dacochelys Lydekker 1889 would be a straight synonym of Palaeaspis Gray 1870, type Emys conybearzi Owen

6 BREVIORA NO. 13

Horizon. Intertrappean Eocene of Bombay.

Comment may now be made on other records and alleged records of this form.

In 1890 Lydekker reported an entoplastron showing half of an intergular scute from the Intertrappean beds in the Nagpur district as a second occurrence of ‘“Hydraspis leithii.”’ The new fragment was much larger than typical Carteremys leithii and differed also in the much narrower intergular. Lydekker passed over the differences casually with a mention of variability in Hydraspis (= Phrynops) hilar. It is unlikely that he had in hand any member of the genus Hydraspis or Carteremys leith. It is probable that he had some other pelomedusid, and he had himself previously (1887) described a Podoc- nemis indica in the Eocene of India.

Sukheswala (1947, an abstract only) has reported a find of a shell of Testudo (= Carteremys) leithii in the Intertrappean of Worli Fill, Bombay. Here, as in the case of the specimens described by Carter, remains of frogs (Indobatrachus pusillus) were associated. In this instance there is no doubt of the identification.

Dr. Sukheswala has kindly sent me a photograph of this specimen (plate 3). The outline of the shell (somewhat different from that figured by Carter) is clearly shown, as is also the characteristic sculpture of the surface (mentioned by Carter), which while somewhat like that of some chelvids also resembles that of, for example, the American pelomedusine genus Taphrosphys. More important is the evident presence of several neurals, the first vertebral clearly much smaller than the second, and (less certainly) the absence of a nuchal shield. The plastron and the skull are unfortunately missing. As with Carter’s specimens the new shell is small, eight inches long by six wide. The specimen is now in the possession of the Geological Survey of India.

Two other specimens have since been found by Dr. Sukheswala and have been sent by him to the Geological Survey of India. These are recorded in the general report of the Survey for 1948 (West, 1950). These specimens, while recognizable, afford no additional information.

Also recorded by the Survey in the same report was a possible young shell of Carteremys leithii from the carbonaceous shales of an Inter- trappean band near Raibasa in the Chhindwara district, Central Provinces, India. The Survey has generously sent me a photograph of this specimen. The identification must be pronounced doubtful as, indeed, the Survey has regarded it. While this new locality may be

“J

1953 FOSSILS AND DISTRIBUTION OF CHELYID TURTLES

thought of hopefully, it would appear that Carteremys leith is at present known with certainty only from the Intertrappean of the island of Bombay.

Acknowledgments. Dr. A. S. Romer, Dr. P. J. Darlington, Dr. P. E. Vanzolini, Dr. Karl F. Koopman, Dr. W. E. Swinton and Mr. L. I. Price have read the manuscript and given critical advice. The allo- cation of the Carter fossil has been discussed with Dr. Rainer Zangerl. Mr. Arthur Loveridge, Mr. C. M. Bogert and Dr. H. W. Parker have permitted the examination of Recent skeletal material in their de- partments. I wish to thank also the authorities of the Bombay Natural History Society for endeavoring to locate Carter’s types, the Geological Survey of India for furnishing me all the information in their possession in regard to the new specimens of Carteremys and for photographs of certain of them, and especially Dr. R. N. Sukheswala of St. Xavier’s College, Bombay, for his prompt and courteous replies to iny insistent questions and for the gift of the photograph reproduced as plate 3.

REFERENCES

CARRINGTON DA Costa, J. 1940. Um novo quelonio fossil. Portugal Com. Serv. Geol., vol. 21, pp. 107-123. Carter, H. J. 1852. Geology of the island of Bombay. Jour. Bombay Branch Roy. Asiatic Soc., vol. 21, pp. 161-215. Gray, J. E. 1870. Supplement to the Catalogue of Shield Reptiles in the Collection of the British Museum. Part I. Testudinata. London. 120 pp. 1871. Notice of a fossil hydraspide (Testudo leithii Carter) from Bombay. Ann. Mag. Nat. Hist., ser. 4, vol. 8, pp. 339-340. LINDHOLM, W. A. 1929. Revidiertes Verzeichnis der Gattungen der rezenten Schildkréten nebst Notizen zur Nomenklatur der Arten. Zool. Anz., vol. 81, pp. 275-295. Lortet, L. 1883. Poissons et reptiles du lac de Tiberiade et de quelques autres parties de la Syrie. Arch. Mus. Hist. Nat. Lyon, vol. 3, pp. 99-194. LYDEKKER, R. 1887. Eocene chelonians from the Salt Range. Paleontographica Indica, ser. 10, vol. 4, pp. 59-65.

8 BREVIORA No. 13

1889a. On the remains of Eocene and Mesozoic Chelonia and a tooth of 2(Ornithopsis). Quart. Jour. Geol. Soe. London, vol. 45, pp. 227-246. 1889b. Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natural History). Part III. London. 239 pp. 1890. Notes on certain vertebrate remains from the Nagpur district. II. Part of a chelonian plastron from Phisdura. Rec. Geol. Surv. India, vol. 23, pp. 22-23. LypEKKEmR, R. and G. A. BOULENGER 1887. Notes on Chelonia from the Purbeck, Wealden and London Clay. Geol. Mag., ser. 3, vol. 4, pp. 270-275. Nopcsa, F. 1931. Sur des nouveaux restes de tortues du Danien du Midi de la France. Bull. Soc. Geol. France, ser. 5, vol. 1, pp. 223-234. 1934. The influence of geological and climatological factors on the distri- bution of non-marine fossil reptiles and Stegocephalia. Quart. Jour. Geol. Soc. London, vol. 90, pp. 76-140. REINACH, A. VON 1903. Schildkrétenreste aus dem aegyptischen Tertiir. Abhandl. Senckenberg. naturf. Ges., vol. 29, pp. 1-64. Scumipt, K. P. 1940. A new turtle of the genus Podocnemis from the Cretaceous of Arkansas. Geol. Ser. Field Mus. Nat. Hist., vol. 8, pp. 1-12. SIEBENROCK, F. 1907. Beschreibung und Abbildung von Pseudemydura wmbrina Siebenr. und iiber ihre systematische Stellung in der Familie Chelydidae. Sitz-Ber. Akad. Wiss. Wien. Math-naturw. KI. vol. 116, pp. 1205-1211. STEJNEGER, L. 1909. Generic names of some chelyid turtles. Proc. Biol. Soc. Wash- ington, vol. 22, pp. 125-127. SuKHESWALA, R. N. 1947. A fossil tortoise (Testudo leithii) from the Intertrappean of the Worli Hill, Bombay. (Abstract.) Proc. Indian Sci. Congress, 33rd Session, vol. 3, p. 97. West; W. D: 1950. General report of the Geological Survey of India for the year 1948. Rec. Geol. Surv. India, vol. 82, pp. 1-253. ZANGERL, R. 1948. The vertebrate fauna of the Selma Formation of Alabama. Part II. The pleurodiran turtles. Fieldiana, Geology Memoirs, vol, 3, pp. 23-56.

Plate 1. Carteremys leithii, dorsal view of shell and skull, slightly modified for clarity from the original reconstruction (Plate X of Carter, 1852). Skull restored from the more perfect half.

Plate 2. Carteremys leithii, ventral view of shell and mandible, slightly modified for clarity from the original reconstruction (Plate XI of Carter, 1852).

Plate 3. Carteremys leithii, photograph of dorsal surface of shell. (Courtesy of Dr. R. N. Sukheswala.)

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. May 28, 1953 NUMBER | 4

RECORD OR A HERMAPHRODITIC HORSESHOE CRAB, ON CTEL UGS VOY EY Gels NUE ES Ge

By JoHn P. Baptist

lishery Research Biologist U.S. Fish & Wildlife Service

Newburyport, Mass.

A hermaphroditic horseshoe crab, Limulus polyphemus L., was picked up by a small scallop dredge from the channel of Plum Island Sound, Massachusetts on May 16, 1952. As far as can be determined, this is the first such Limu/us to be recorded.

The general appearance was that of a mature male. Its width (125 mm.) was about average for males in this area. (The widths of 606 mature males measured this summer ranged from SS mm. to 140 mm., with an average of 117 mm., while 489 mature females ranged from 130 mm. to 199 mm. in width, with an average of 155 mm. ) It had mating claspers, and the anterior edge of the prosoma was curved upward in the middle, which also is a male characteristic. The left genital aperture was typically male, round and located on a definite papilla. The right genital aperture was typically female, a horizontal slit and not on a papilla (Lochhead, 1950). The genital operculum, showing both male and female genital apertures, may be seen in the upper illustration of Plate 1. On the dorsal surface of the opisthosoma near the telson were darkened areas normally present on mated females. These may be barely discernible in Plate 2. They are pro- duced by the abrasive action of the anterior edge of the male as he clings tenaciously to the female during the mating period. Females which have mated for prolonged periods exhibit deep scars on the last pair of immobile spines, caused by the powerful claspers of the male.

BREVIORA no. 14

bo

Such scars were not evident on the hermaphrodite, suggesting that it had been mated as a female for a comparatively short time.

Gross dissection of the prosoma and histological sections revealed both male and female gonads. The right and anterior portions were full of eggs in various stages of development (Plate 2), comparable to to those of mature females. Histological sections of the tissue on the left side demonstrated sperm sacs full of tail-less sperm, diverticula of the hepatopancreas and connective tissue (Plate 1, lower illustration). The same structures were visible in sections of gonads from norma! males. According to Benham (1885) the tails are apparently produced as the sperm approach the aperture.

The presence of both eggs and sperm confirms the external evidence

that this specimen is a true hermaphrodite.

REFERENCES

BENHAM, W. B.S. 1885. On the testis of Limulus. Trans. Linn. Soc. Lond. (Zool.), (2)

2: 363-366. LocHHEAD, J. H.

1950. NXiphosura polyphemus. In Selected Invertebrate Types, edited by F. A. Brown, Jr. pp. 360-381. John Wiley & Sons, Inc., N. Y.

1953 A HERMAPHRODITIC HORSESHOE CRAB 3

Upper Ventral view of posterior surface of genital operculum (enlarged), showing right genital aperture (Q ) and left genital aperture (<7). Lower Photomicrograph of section through testis (X 120). S, sperm sacs

containing tail-less sperm. H, hepatopancreas. C, connective tissue. (Photographs by Alden P. Stickney)

BREVIORA No. 14

PAG 2

Dorsal view of Limulus polyphemus with carapace of the prosoma removed, showing mass of eggs on the right side, but only a few scattered eggs on the

left side. (Telson has been cut off). (Photograph by Alden P. Stickney.)

BREVIORA

Museum of Comparative Zoology

CAMBRIDGE, Mass. Niay 1, 1953 NuMBER 15

FURTHER NOTES ON THE EARTHWORMS OF THE ARNOLD ARBORETUM, BOSTON

By G. E. Gatss

An opportunity to search once more for mature specimens of the Pheretima sp., represented in collections of the previous spring by a single, specifically unidentifiable juvenile, was unexpectedly provided on September 25, 1952. After securing the desired material, as many of the previous sites as time permitted were re-examined, to permit comparison of fall with spring populations.

In the natural woods, during the summer, the leaf pile of previous collections had been grown over and had become rather dry. Some fifty feet away, and at about the same distance from the site where the juvenile of Pheretima sp. had been found, another large leaf pile, possibly older and with more gravel, had remained damp and without any plant overgrowth. In the peat bog, where vegetation had been mowed and removed, water was no longer running in the brook though still standing in small, scattered puddles, but previously disturbed soil still seemed moist. In the ponds, water level had fallen several inches exposing a marginal strip of bottom muck that was still saturated. The Newton garden had been planted and could not conveniently be re-examined.

The species obtained in the September collections are listed in the ~ following table which also shows sexual stages and number of specimens secured.

2 BREVIORA no. 15

EARTHWORMS OBTAINED IN THE ARBORETUM IN SEPTEMBER, 1952

Natural Woods

Species Original Second Leaf Pile Peat Bog Leaf Pile Near Side Far Side Allolobophora T = arnoldi 0-2-2 0-1-0 0-0-1 0-0-1 caliginosa 0-0-2 0-0-1 limicola 0-2-0 sp. (1) 4-0-0-0 Dendrobaena subrubicunda 3-5-32 1-0-1 Eisenia foetida 12-5-6 7-1-7 2-0-1 Lumbricus castaneus 0-0-2 rubellus 3-1-7 2-2-3-16 6-11-17 terrestris 4-1-0-4 1-0-1 Octolasium lacteum 0-0-1 Pheretima agrestis 0-0-35 0-0-21

Papas indicate juvenile-aclitellate-clitellate or small juvenile-late juvenile-aclitellate- clitellate.

+ Postsexual aclitellate. * Specimen in poor condition. (1) Probably of arnoldi and /or caliginosa.

A. arnoldi. Two specimens from original leaf pile listed as clitellate had spermatophores externally but the clitellum had almost completely re-<