SMITHSONIAN INSTITUTION UNITED STATES NATIONAL MUSEUM

PROCEEDINGS

OF THE

UNITED STATES NATIONAL MUSEUM

VOLUME 45

WASHINGTON GOVERNMENT PRINTING OFFICE 1913

ADVERTISEMENT.

The scientific publications of the National Museum consist of two series—Proceedings and Bulletins.

The Proceedings, the first volume of which was issued in 1878, are intended primarily as a medium tor the publication of original papers based on the collections of the National Museum, setting forth newly acquired facts in biology, anthropology, and geology derived there- from, or containing descriptions of new forms and revisions of limited groups. A volume is issued annually or oftener for distribution to libraries and scientific establishments, and, in view of the importance of the more prompt dissemination of new facts, a limited edition of each paper is printed in pamphlet form in advance. The dates at which these separate papers are published are recorded in the table of contents of the volume.

The present volume is the forty-fifth of this series.

The Bulletin, publication of which was begun in 1875, is a series of more elaborate papers, issued separately, and, like the Proceedings, based chiefly on the collections of the National Museum.

A quarto form of the Bulletin, known as the ‘“‘Special Bulletin,” has been adopted in a few instances in which a larger page was deemed indispensable.

Since 1902 the volumes of the series known as “‘Contributions from the National Herbarium,” and containing papers relating to the botan- ical collections of the Museum, have been published as Bulletins.

RicHARD RATHBUN, Assistant Secretary, Smithsonian Institution, in _ charge of the United States National Museum.

DEcEMBER 5, 1913. II

TABLE OF CONTENTS.

Bartscu, Pauy. New land shells from the cia telaide:—— No. 1993, June 21, V9lat et a es

New species: Orba worcesteri, Cochlostyla calusaensis, C. olaniva- nensis.

The giant species of the molluscan genus Lima obtained in Philippine and eae a waters.—No. 1978. petro ae ROL Sie te cies Re a

New species: Lima (Callolima) emithi, L. C. ) Tiina DE: “(C) rathbuni, L. (C.) borneensis, L. eae verdensis, L. (A.) cele- bensis, Ee (A.) butonensis.

The Philippine mollusks of the genus Dimya.— Pm lume ytd OTS, Tas ule ed 3 yaa en ae

New species: Dimya filipina, D. lima.

Berry, Epwarp W. A fossil flower from the Eocene.— Rees weenie OS. Oa Ti oo ae on ye ea es

New genus: Combretanthites. New species: Combretanthites eocenica.

Berry, S. STILLMAN. Some new Hawaiian cephalopods.— een OO Sunie, 4 AOL Ebates me

New genus: Laetmoteuthis.

New species: Laetmoteuthis lugubris, Scaeurgus patagiatus, Eu- prymna scolopes, Teleoteuthis compacta, Abralia trigonura, Pte- rygioteuthis microlampas.

Bruner, LawRENCE. Results of the Yale Peruvian expe- dition of 1911. Orthoptera (Addenda to the Acridiidee— short-horned locusts).—No. 2001. June 11, 1913%____--

New species: Schistocerca maculata.

Cxiarx, AusTIN Hopart. Three interesting butterflies from eastern Massachusetts.—No. 1987. June 13, 1913*_-_--_-

CocKERELL, T. D. A. Two fossil insects from Florissant, Colorado, with a discussion of the venation of the Aesh- nine dragon flies.—No. 2000. June 21, 1913?____-.-_--

New species: Oplonxschna lapidaria.

1 Date of publication.

Page,

549-553

235-240

305-307

261-263

563-566

585-586 363-364

577-583

Il

IV TABLE OF CONTENTS.

CRAWFORD, J. C. Descriptions of new hymenoptera, No. aaee. 6-=No.1979., “May 22, 1913 4) Cag aaa ee

New genera: Bruchocida, Coccidoxenus, Zacalochlora, Trichomalopsis, Aplastomorpha, Cassidocida, Thriposoma.

New species: Protandrena swenki, Augochlora maculiventris, Tele- nomus goniopis, T. latisulceus, Ganaspis hookeri, Podagrion shirakii, Bruchocida vuilleti, B. orientalis, Tanaostigmodes portoricensis, Coccidoxenus portoricensis, Anastatus formosanus, Bruchobius colemani, Zacalochlora milleri, Trichomalopsis shirakii, Aplasto- morpha pratti, Cassidocida aspidomorphx, Pleurotropis anastati, Thriposoma grafi, Zagrammosoma flavolineata, Z. centrolineata, Z. nigrolineata, Sympiesis metacomet, S. massassoit, S. maculipes, S. bimaculata, Anagrus giraulti.

Descriptions of new hymenoptera, No. 7.—1984. Pleas 2 VOLS Be ye iho Se a let CA oe We ee

New genus: Pareniaca.

New species: Aglaotoma texana, Eucoila hunteri, Psilosema pratti, Hontahia magnifica, Pareniaca schwarzi, P. buscki, Polycystus foersteri, Cercocephala atroviolacea, Derostenus agromyze, D. ari- zonensis, D. variipes, Entedon thomsoni, Pleurotropis utahensis, Cirrospilus flavoviridis.

Dati, Witit1am Heatey. Diagnoses of new shells from the Pacific Ocean.—No. 2002. June 11, 19131 ___._..._.... 587-597

New genus: Halicardissa.

New subgenus: Cosmioconcha.

New species: Tritonofusus jordani, Boreothropon gorgon, Amphissa (Cosmioconcha) palmeri, A. (C.) pergracilis, A. (C.) parvula, Liotia lurida, Bolma bartschii, Margarites simblus, Calliostoma nepheloide, Pecten (Pseudamusium) arces, Cuspidaria subglacialis, Psephidia cymata, Lyonsia (Allogramma) amabilis, L. (A.) oahuénsis, L. pugetensis, Lyonsiella magnifica, Poromya (Dermatomya) tenut- concha, Erycina colpoica, Rochefortia compressa, Aligena nucea, Vesicomya (Archivesica?) suavis.

Dyar, Harrison G. Results of the Yale Peruvian expedi- tion of 1911. Lepidoptera.—No. 2006. July 22, 1913'.. 627-649

New genus: Alitmznas.

New species: Phulia altivolans, P. nannophyes, Andina coropune, Actinote binghamex, Phycoides omosis, P. birivula, Lymanopoda shefteli, L. harknessi, L. keithi, Euptychia hotchkissi, E. leguia-limai, Thecla tyleri, T. bennetti, T. brocela, T. ulia, T. muela, T. excisi- costa, Hylephila isonira, Argopteron xicca, Lerodea gracia, Lerema miqua, Hesperia archia, Saturniodes orios, Virbia elisca, V. catama, Altimenas tapina, Mesembreuxoa fasicola, Porosagrotis propriens, Metalepsis cerphiphila, Lycophotia albiorbis, Hyssia elxochroa, Lasionycta comifera, Cobubatha rilla, Plusia monozxyla, Anomis sophistes, Epirrhe diltilla, Triphosa quasiplaga, Spododes unifacta, Polypoetes marginifer, Jocara suiferens.

New subspecies: Papilio madyes montebanus, Heliconius melpomene hyper plea.

1 Date of publication.

TABLE OF CONTENTS.

GILBERT, CHESTER G., and JosrpH E. Poaurn. The Mount Lyell copper district of Tasmania.—No. 2005. July 22, RP eee ere ees et LOS VOL NC Uk Ns Sale OMS AU as eh

GirauLt, A. ArsENE. A systematic monograph of the Chalcidoid hymenoptera of the subfamily Signiphorinze.— LS gE EOMR P22) ee Eo Is oa ACOA Ppa A

New species: Signiphora flava, S. flavella, S. basilica, S. pulchra,

S. maxima, S. melancholica, 8. fasciata, S. hyalinipennis, S. macu- lata, S. nigrella, S. fax, S. funeralis, S. corvina, S. australica.

Houuister, N. Mammals collected by the Smithsonian- Harvard expedition to the Altai Mountains, 1912.—No. [OR GPRS hea a 107 RE RAS oR TAN

Know tron, F. H. Description of a new fossil fern of the genus Gleichenia from the Upper Cretaceous of Wyo- fine NO. 1994.) Sune 21; 1913 *. ede eR Doe

New species: Gleichenia pulchella.

JORDAN, Davip Starr, and JOHN OTTERBEIN SNYDER. Description of the Yachats ‘‘Smelt,” a new species of Atherinoid fish from Oregon.—No. 1999. June 21, 1913 1_

New species: Atherinops oregonia.

Lyon, Marcus Warp, Jr., Treeshrews: An account of the mammalian family Tupaiide.—No. 1976. November 29, UTS SS a Sg UE eid Sem coals reams Poe ag arenas a SRA Me, UAW A, ST

New genera: Anathana, Tana.

New species: Tupaia riabus, T. anambe, Anathana wroughtoni, A. pallida, T. linge, T. paitana.

New subspecies: Tupaia longipes salatana, T. montana baluensis, T. gracilis edarata, Dendrogale melanura baluensis, Tana tana besara, T. t. utara, T. t. tuancus, T. cervicalis mase.

Mattocy, J. R. Three new species of Anthomyide (Dip- tera) in the United States National Museum collection.— ier Ose a Uume ME POTS Ds yk Oe yt ea

New genus: Paralimnophora.

New species: Tetramerinz femorata, Paralimnophora brunnesquama, Anthomyia bidentata.

Pearse, A. S. Notes on a small collection of amphipods from the Pribilof Islands, with descriptions of new species—No. 1998. June 4, 19131 ___________ __2 Le.

New species: Gammarus pribilofensis, Chironesimus multiarticulatus.

Vv

Page.

609-625

189-233

507-532

555-558

575-576

1-188

603-607

571-573

1 Date of publication.

VI TABLE OF CONTENTS.

Pierce, W. Dwicut. Miscellaneous contributions to the knowledge of the weevils of the families Attelabide and Brachyrhinide.—No. 1988. May 238, 19131. __________-

New subfamilies: Entiminx, Dirotognathiny, Psallidiine, Brachy- rhinine.

New iribes: Rhynchitini, Eremnini, Tropiphorini, Pandeleteini, Alceidini, Polydrusini, Blosyrini, Sciaphilini, Thylacitini, Tri- gonoscutini, Calyptillini, Celeuthetini, Trachypheleini, Simoini, Eustylini.

New genera: Amydrogmus, Hadromeropsis, Glaphyrometopus, Brady- rhynchoides.

New subgenera: Panscopidius Neopanscopus.

New species: Hugnamptus punctatus, Amydrogmus variabilis, To- sastes cinerascens, Peritaxia elongata, Melamomphus nigrescens, M. ciliatus, Dyslobus bituberculatus, D. denticulatus, Panscopus (Phymatinus) sulcirostris, P. (Panscopidius) squamosus, P. (P.) dentipes, P. (Nomidus) impressus, P. (N.) ovalis, P. (Neopan- scopus) squamifrons, P. (N.) carinatus, Pandeleteius dentipes, P. depressus, Glaphyrometopus ornithodorus, Bradyrhynchoides con- strictus, E’picerus wickhami, E. benjamini, Pantomorus (Phace- pholis) nebraskensis, P. (P.) metallicus, P. (P.) texanus.

New varieties: Ewgnamptus collaris fuscipes, E. c. ruficeps, E. punc- tatus niger, E. angustatus testaceus, Merhynchites bicolor cockerelli, M. b. ventralis, M.b. piceus, M. b. viridilustrans, Cimboccera pauper sericea.

Poauet, JosEPH EK. See under GILBERT, CHESTER G-- -- --

Ricwarpson, Harriet. The isopod genus Ichthyoxenus Herklots, with description of a new species, from Japan.— Ge LOOKS: Mum i POV GE ral ih Ue es cs EO ep een ee pa

New species: Ichthyoxenus japonensis.

Rirrer, WittiaAmM E. The simple Ascidians from the north- |

eastern Pacific in the collection of the United States National Museum.—No. 1989. June 25, 19131______-_--

New genus: Hartmeyeria.

New species: Molgula oregonia, Eugyriotdes dalli, Halocynthia wash- ingtonia, Hartmeyeria triangularis, Culeolus sluiteri, Styela macren- teron, S. hemicespitosa, S. sabulifera, Corynascidia herdmani, Agnesia beringia, Phallusia vermiformis, P. unalaskensis.

New subspecies: Halocynthia haustor foliacea.

Rouwer, S. A. A synopsis, and descriptions of the Nearctic species of sawflies of the genus Xyela, with descriptions of other new species of sawflies.—No. 1981. ery LOLS tee Na is eek ne LIU. othe bed pce een enone

New genus: Allantopsis.

New species: Xyela salicis, X. pini, X. alni, X. winnemanz, X. errans, X. brunneiceps, X. dissimilis, X. slossonx, X. nevadensis, X. similis, X. californica, X. coloradensis, Pamphilius (Pamphi- lius) greenei, Allantopsis thoracica, Tenthredella carolina, T. fisheri, Proselandria peruviana, Stromboceros (Neostromboceros) assamen- sis, Pteronidea pulchella, P. vanduzeei, Pristophora xanthotrachela.

1 Date of publication.

Page.

365-426

609-625

559-562

427-505

265-281

TABLE OF CONTENTS.

Rouwer, S. A. Descriptions of thirteen new species of parasitic hymenoptera and a table to certain species of the genus Ecphylus.—No. 1991. June 4, 1913*______-

New species: Atanycolidea apicalis, Xylonomus (Xylonomus) lep- ture, Pristaulacus floridana, Triaspis fiskei, Heterospilus leptostyli, Cxnopachys scolytivora, Ecphylus hubbardi, E. lycti, E. schwarzi, E. johnsoni, E. californicus, E. lepturgi, E. bicolor.

New parasitic hymenoptera belonging to the tribe Moridini-—No. 1986. May 22019132. oo Sole ll New species: Xylonomus (Xylonomus) plesius, X. (X.) ruficovis, X. (Maerophora) yukonensis, X.(M.) eastoni, X. (M.) duplicatus,

X.(M.) piceatus, X. (M.) modestus, Odontomerus atripes, O. alas- kensis, O. errans, O. dichrous.

SmitH, Huew M. Description of a new Carcharioid shark from the Sulu Archipelago.—No. 2003. June 21, 1913 *-

New genus: Eridacnis. New species: Eridacnis radcliffer.

The Hemiscyllid sharks of the Philippine Archi- pelago, with description of a new genus from the China eee NO LOO 7. UNS 21, LOTS os oe ae ee Se eae

New genus: Cirrhoscyllium. New species: Cirrhoscyllium expolitum.

SNYDER, JOHN OTTERBEIN. See JoRDAN, DAVID STARR. - - -

STEJNEGER, LEONHARD. Results of the Yale-Peruvian expedition of 1911. Batrachians and reptiles—No. 1992. PIRSHR ENE men Syn UME r eae PNM, ce Lit NL atl all Pk

New species: Bufo inca, Eleutherodactylus binghami, E. footei, Steno- cercus ervingi, Oreosaurus lacertus.

Trur, FrepErRicK W. Description of Mesoplodon mirum, a beaked whale recently discovered on the coast of North Carolina.—No. 2007. November 29, 19131!___________-

Wicxuam, H. F. Fossil coleoptera from Florissant in the United States National Museum.—No. 1982. June 13, RPE antes eo ee Stes Lage cs ep Man RL eR Cilla

New genera: Aleocharopsis, Miolithocharis, Miostenosis.

New species: Agabus florissantensis, Anisotoma sibylla, Aleocharop- sis caseyi, A. secunda, Staphylinus vulcan, Miolithocharis litho- graphica, Lithocoryne arcuata, Cryptophagus bassleri, Tenebroides corrugata, Anthaxia exhumata, Lucanus fossilis, Aphodius grana- rioides, Diplotaxis aurora, Leptura antecurrens, L. ponderosissima, Systena florissantensis, Bruchus dormescens, Miostenosis lacordairet, Blapstinus linellii, Platydema bethunet.

1 Date of publication.

VII

Pag?.

533-540

353-361

599-601

567-569 575-576

5041-547

651-658

283-303

Vill TABLE OF CONTENTS.

Wituiams, Henry Sater. New species of Silurian fossils ESE from the Edmunds and Pembroke formations of Washing- ton County, Maine.—No. 1985. July 22, 19131_____._. 319-352

New genus: Palxopecten.

New species: Whitfieldella edmundsi, Chonetes edmundsi, C’. cobscooki, Brachyprion shaleri, Palzopecten cobscooki, P. transversalis, Chone- tes bastini, Camarotechia leightoni, Lingula scobina, Actinopteria bella, A. fornicata, A. dispar, Grammysia pembrokensis, Leiopteria rubra, Modiolopsis leightoni, Nuculites corrugata.

New variety: Lingula minima var. americana.

1 Date of publication.

SO CON DD Or O

WWWNNHNPNMNWNW NHN MWD EF EB EB BBM Be KE ee NF OM ON OOH WNFOWOOAN OOK WD

33. 34, 30. 36. 37. 38. 39. 40.

bist OF TLLUSTRATIONS.

PLATES.

. Condore Island treeshrew, Twpaia dissimilis from Ellis’ manuscript journal,

HSU aeperees ae Sey ete LCE LK A a Tete RE ets Mb Ae MERA SIME Ue Nena ey San

. Malay Peninsula treeshrew, Twpaia lacernata Wilkinsoni..........-------- Weking o: treeshrews of the genus Tupata 2.0.90. foe ef ee . Indian treeshrew, Anathana ellioti from Waterhouse.............-..------ . Pigmy treeshrew, Dendrogale murina from Schlegel and Miiller...........- . Skins of treeshrews of the genera Tana and Urogale..........-..--.------ . Pentailed treeshrew, Ptilocercus lowti after Gray.........---------------- . skulls of treeshrews of the genus Tupaia......-..--+--- 222. e eee eet Paskulls of treeshrews of the genus Tupaia......--2-----2-5-4-sesse55ee == . Skulls of treeshrews of the genera Tupaia and Ptilocercus....-.-.--.--.--- . Skulls of treeshrews of the genera Tana and Urogale............---------- EE PACOHM OMIA) SIMEDAT (6 i031 28 oe Ue Ale ee Se eee mee MCA ALIA WANTS oi se ec eee cele aeaies FeO hie See eae meen ( Calloling) phil ppinensts...... <x, a)s s's.-.2- ean 29s me see eae ae eens Re COOL) SET BER SIR re oo 8 I AS etal Was Sek I me REID ACE ALONG) PALRDUTE SO 2 A001.) xin adizin, seedy iden Saiemincs Sowa aeee eee BPE eae ea ON tTIUG PLEIN Ser Sd, Te I hs SN Dr Ue Act SL tae ONPORLC COMCDONSE BH i. hoc he Lathe Seella oid oie arain soma es acjanis gen eae eee BePPe UREN LSE CLL COCIIOIS Olen trae oC ctete 2 Lercha/ SW cla ahs at eNeecicie ce aes ME OCA TUNIS A) ie cS ie ke Lo in ee Ra oi Me eS ae ma cox (Lower Bocene) dossil flower: «2.62.20» .!s<iemisea 2) s7osecc anced ee Mie coleoptera irom. Wlorissant., .. cn.) os. no. a Sucaes «eine ene eee ee seas colcoptera from, Mlorissamt 2 .05-\2.0. 52. eae tind os ace asee cea aeons coopers from, Wlorissant. | oi) - a2 «nye «slam dare iin en os one See wage moni colcoptera trom Florissant. 2/322 <inadel- ssicjaec ss hs ese bloom canes meoenl colooptera irom Mlorissam tis. |. 6 ine.cscon ond ls bee Gine vOeulce eaauen RPE GHA LET EE OT TTI BM ERD rom ei aes oh wk ie oe ca ie e/g maow Dinvas from the Philippines. - 20.45.) 2s <i... esecics--- se Saye merew.epoctes Of Sulirign Topstig) 2 3s ee ler 2 ene PROM ADP Ces Of Sllurian FOR. 8 oo sec p/ciase a sande Ganceon thous see ROMA peCION OF SIUTEAT HOSS sie ea ae cline oi Ue Sik upc Gk a ee . Euphydryas phaeton (Drury); underside of a suffused female representing the

WarIOLV SIPCMMGIOtneC KEP: ook ou et yi lel dace Simple Ascidians from the northeastern Pacific...................------- Simple Ascidians from the northeastern Pacific...................-.---- Simple Ascidians from the northeastern Pacific..................-.----- Simple Ascidians from the northeastern Pacific. ....................---- Collecting stations in the Altai Mountains. ..................22...-.---- Natives of the Altai and habitat of Platycranius...................------- Scenes in the Altai Mountains................... ates ata ie era ate Skull of Ovis ammon from Altai Mountains. ..................-----------

Ix

Facing page.

184 184 184 184 184 184 184 184 184 184 184 240 240 240 240 240 240 240 240 240 264 304 304 304 304 304 308 308 352 352 352

364 506 506 506 506 532 532 532 532

x LIST OF ILLUSTRATIONS.

41.\Skull of Procapra altaica from 'Suok..Plaing. 2:2. / S05. Pee ce ee 42. Skull and teeth of Procapra altaica from Suok Plains..................... 43. New land shells from the Philippine Islands. .................-......... “wa new 10s) fern OL the genus Glewhenia..u. 002255 22a seen seu eee ae eee MC TIF ROBCYLLUUN? ELDOLILUT. = sion aus adie mice Cie cic ects om eta ee etc ae EOAthEMnoDs oregonia, RCW species. 6.27.2 US ee ee ae ME OUCIIS TAOCUP OL. » a. oon wis leis ia ale se Merate tg exe sue Walenta) in eae eee PeeeOUsned S6CtIONS Of Orewa. 0s Sacks bis Se er eee 49. Photomicrographs of polished ore sections... ..-.:----5--.22222¢2-.2..200, 50. Protomicrographs of polished ore sections. ........-------...-..--.------ 51. Photomicrographs of polished ore sections......--.-..-..----.-----.----- maryppeaked whale, Mesoplodon minum... -)2225 00. 25. Rs ae Beaevaews of head.of beaked, whales.) uci. j20 cE So a oe ee eae Baouperior view of skull of beaked/ whale. 03.5050 coe 55) Interior view of skull of beaked whale... -.. 2.202.242 bo. a ae ae auateral view of skull of beaked) whale. 20002000 )000c002 uehe e Bago) Views of mandible of beaked whales. 02.0 5))05. 50s ee

TEXT FIGURES.

Diagram showing the form and relationships of the individual bones of the skull of Tupaia as determined by examination of young individuals......... Diagram illustrating the rhinarium (1) in the genera Tana and Urogale (2) in the genera Tupaia, Anathana, and Dendrogale..........-..--.1------------ Palmar surface of right forefoot and plantar surface of right hindfoot of Tupaia glis ferruginea. Hy. thn. hypothenar pad; 7. d.1, 7. d.?, 7. d.3, 7. d.4, first, second, third, and fourth interdigital pads; prz. e, proximal external pad; pra. in, proximal internal’ pad; then, thenar pad 2220). ee ee par gis ferrugimea: 'Tapanul Bay, Sumatra -..000. 2 ee Tupaia glis ferruginea, Upper and lower toothrows X24. Tapanuli Bay, amaranth a) Sel cn Wo 0 La ea ea Map of the Malay region, showing the distribution of the forms of the genus Tupaia, excepting the members of the gracilis, javanica, and minor groups... Map of the Malay region, showing the distribution of the minor, gracilis, and javanica groups of the genus Tupaia. 1. Tupaia javanica; 2. T. minor minor; 3. T. minor malaccana; 4. T. minor sincipis; 5. T. gracilis gracilis; Ga graciiis injlata; 1. ..T. .gractlis edarata 40.2 /2 Rea) ee ee Anathana wroughtoni. British Museum, Mandvi, India.............-...----- Upper and lower toothrows of Anathana wroughtonit. British Museum, Mandvi,

Map showing the distribution of the genus Anathana, contrasted with the dis- tribution of Tupaia on the Asiatic Continent. 1. Anathana ellioti; 2. A. wroughiony, 3. A. palhda; T. genus Tupaia::.. 25. .b3e eee eee

Himuicogale melanura melanura, type X<1F). 2.42 WS. SU eee

Upper and lower toothrows of Dendrogale melanura melanura, X3}.-----------

Map of the Malay region, showing the distribution of the genera Dendrogale and Ptilocercus. A.—Dendrogale frenata; B.—D.murina; M-D. melanura melanura; N—D. melanura baluensis; 2. Ptilocercus lowii lowii; 3. P. lowii PERVETUTIS =o a eNchc a no ele Hale a okie wikia eke CARO as le el I ee

Pee LENG DESOTO, TYPO CL Hoon woke dace ae ane eed i ee ee

Upper and lower toothrows of Tana tana besara, type X2....-.-..------+----+--

32 33

33

75

dil 120

121

125

127 128

LIST OF ILLUSTRATIONS.

Map of the Malay region, showing the distribution of the genera Tana and Urogale. 1. Tanachrysura; 2. T. paitana; 3. T.tanautara; 4. T. tana besara; 5. T. tana tana; 6. T. tana bunox; 7. T. tana sirhassenensis; 8. T. tana tuan- cus; 9. T. cervicalis cervicalis; 10. T. cervicalis masx; 11. T. lingzx; 12. T. BERANE OPE LIPO GILC CUCLOLEL ea coe A's) Sas 21 apa aie ia yah Suse eta alata Ga aN A a PE CUM ELT UV DO 6 baccCese sid iia oe ote aie ciaa Cate ie Rae o ce arene aes eee Upper and lower toothrows of Urogale everetti; type, X2.-....-..----.-------- Piilocercus lowi continentis, X14, Sembrong River, Johore, Malay Peninsula. - Upper and lower toothrows of Ptilocercus lowi continentis, x4, Sembrong River, enlore (Veal Fr Chinese. Yoh ho tees woe See eh el eee Coccidoxenus portoricensis. Outline of antenne of female................--.-. Zagrommosoma flavolineata. Fore wing of female with stipple to show infus- Serer ARCA) uit te Ug arlene ae a SE mele a arto a Nacns Gee OAM ere ea Eesoraon Of Compretanthites cocented.-. 282 oc ss oe eae semana aes Ponetion view OL -Allantopsis UOracied . 22. ae a2 os 5 os oe as we Bee eee SORE Pareniaca schwarzi. Side view of head of female............-...-------------- arnuOnenus JeLlungnausit.. Kemale. XSb. 0.2. .5..-2---+ less 5.2 seems Pa IEC GCLLINGIRIUSTE..s MGIC. OX eas otha x) ences 5 = Sn ee oie vc a elrae Ichthyoxenus jellinghausw. Seventh leg of female. X7}...........--...------ Ichthyoxenus japonensis. Lake Biwa, from <Acheilognathus cyonostigma. Fe- PLYV CHILI epee ntIUz oUNIOUS A ce AR UE GBS) a alice Rl yo ruee leas a Wo een en Ichthyoxenus japonensis, Lake Biwa, from Acheilognathus limbatum. Male.

Ichthyoxenus japonensis, Lake Biwa, from Acheilognathus cyonostigma. Seventh ResrasiOMIa len, Kaa soe 2 cs oaks als eelale's calecels Lites ste gene See a a Dermal denticles from side of Cirrhoscyllium, enlarged.......-.-------------- Front and side views of teeth of Cirrhoscyllium near symphysis, greatly en- larged. The dotted lines indicate the margin of the gums. a,upper jaw; BLO NVAS TN OMe SF a a oe as sy alae RSS ateli a Ga) ie ele eto Se Gammarus pribilofensis. a, first antenna; g', first gnathopod; g?, second gnath- MET e OLAIRGNA Te: UL IE, MEODOUS. 00" 3G ei ee = os ae oe te eee ee Chironesimus multiarticulatus. e, third abdominal epimerite; g', first gnath- opod; g?, second gnathopod; 7, first antenna; ¢, telson; uw, third uropod...-.- Oplonxschna lapidaria. a, stigma; 6, triangle and arculus; c, ends of veins m, nua da: radial sector and supplement: <4. 2). jo 2/22. oes See eee Oplonexschna separata. a, triangles from two specimens collected at Station 14, Florissant; 6, Planzschna multipunctata, triangle of hind wing; c, Planxs- Banayorcupai, triangle of hind, wing .2's ee. soe. eee os See So Nae Paraspiniphora laminarum. a, diagrammatic figure of middle tibia; 6, part of middle tarsus, showing armature; c, part of hind tarsus, showing armature; d, scutellum and adjacent parts, showing bristles; e, end of female abdomen... - PmuarnisTaaciiye.. , Under side-of head. 2.602. ..e le. ee esse be ene ee Eridacnis radclifei. Teeth of upper and lower jaws greatly enlarged. .--....--. Eridacnis radcliffei. a, dermal denticle from side; }, cross section of dermal denticle, through middle, showing attachment.............---------.------ parienman of the Mount Lyell district: ..o.i1.0iten<. ona ane aes cea 1 SPELLS eyo a arora LEAR a CaS ea ee SE yeh ee Amn Sk

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TREESHREWS: AN ACCOUNT OF THE MAMMALIAN FAMILY TUPAIID.

By Marcus Warp Lyov, Jr., Formerly of the Division of Mammals, United States National Museum.

INTRODUCTION.

This-review of the treeshrews, constituting the mammalian family Tupaiide, was originally contemplated in 1904 by Mr. Gerrit S. Miller, jr., curator of mammals, United States National Museum, but owing to pressure of other work he was unable to carry it out. In 1910, shortly after I severed my active connections with the Division of Mammals, United States National Museum, Mr. Miller suggested to me the desirability of making a study of the treeshrews. I tookup his suggestion and the present paper is the result. At that time he turned over to me some preliminary notes on the group he had made during a visit to European museums when he was primarily engaged in other lines of research. The increase of new material, both in the United States National Museum and in other museums, made it imperative that the entire field be gone over again. The collections in Washington were first studied, and during the summer of 1911 I visited most of the museums which Mr. Miller’s previous work showed contained material valuable for this revision.

Specifically, the material examined consists of about 800 speci- mens, all of which are listed in the tables of measurements and dis- tributed as follows:

British Museum, 355 specimens, 27 types.

United States National Museum, 324 specimens, 29 types.

Civic Museum of Natural History, Genoa, 37 specimens, no types.

Royal Zoological Museum, Berlin, 29 specimens, 1 type.

Museum of Natural History, Paris, 20 specimens, 1 type.

American Museum of Natural History, New York, 14 specimens, 1 type.

Natural History Museum of Geneva, 3 specimens, no types.

Natural History Museum of Turin, 1 specimen, no types.

In addition to the specimens mentioned above, in most museums, particularly the older ones, there are a number of specimens of very uncertain or generalized localities, which are unsuitable for systematic work, and they are not included in the above figures. :

PROCEEDINGS U. S. NATIONAL Museum, VoL. 45—No. 1976. 80459°—Proc.N.M.vol.45—13——1

2 PROCEEDINGS OF THE NATIONAL MUSEUM. VoL, 45.

In addition to examining specimens in these museums, I have also had for study specimens sent to Washington from the following institutions:

Museum, Philippine Bureau of Science, 12 specimens, 1 type.

Selangor Museum, Selangor, Straits Settlements, 8 specimens, no types."

Academy of Natural Sciences, Philadelphia, 6 specimens, no types.

I was unfortunate in my time of visiting the Natural History Museum in Leyden. The director, Dr. F. A. Jentink, who has direct charge of the mammals, was on his vacation and I was unable to examine the specimens of Tupaiide in that museum. The material contained there as listed by Jentink ? does not appear vitally impor- tant for a systematic review of the group, yet it contains some very interesting and historical specimens, which I regret not having seen. Among them are the unique type of Dendrogale murivna and the only skeleton of the genus Péilocercus that I know of existing in museums and the cotypes of Tana dorsalis. Dendrogale murina is the only species of treeshrew of which I have not seen examples.

I take pleasure in here expressing my thanks to the directors of the museums which I personally visited for giving me the privilege of studying the available material in their institutions, or from which material was borrowed.

The importance of the explorations of Dr. W. L. Abbott in our knowledge of the treeshrews can not be lost sight of. With the exception of less than a dozen specimens in the United States National Museum the entire series of treeshrews there was collected through his untiring efforts. This means that more than a third of the specimens of treeshrews in all the museums of America and Europe have been personally collected by Doctor Abbott. Among them are 29 types. Indirectly he is also responsible for the tree- shrews collected by Messrs. Kloss and Robinson on the Malay Peninsula, or adjacent islands.

The text figures of the skulls and teeth of the various genera were made by Mr. A. J. Engel Terzi.

Measurements.—All the measurements are in millimeters. With the exception of those of the head and body and of tail of skins, they have all been made by the writer, including those of the hind foot, which includes the claws. In most cases the measurements of the head and body and tail were made by the collector in the flesh. In the tables of measurements where the head and body and tail measure- ments are followed by +, those measurements were made by the writer from the:dried skin or mounted specimene Head and body and tail measurenients of specimens preserved in alcohol were also

1 There are, however, in the Selangor Museum, 4 types, none of which I have seen.

2 Cat. Ostéol. Mamm. Mus. Hist. Nat. Pays-Bas., vol. 9, 1887, and Cat. Syst. Mamm. Mus. Hist. Nat. Pays-Bas, vol. 12, 1888.

no. 1976. TREESHREWS: FAMILY TUPAIIDH—LYON. 3

made by the writer. Measurements of the skull followed by + indicate that the measurement is only approximate owing to damage to the skull. Hindfoot measurements followed by + are also only approximate. In the tables of measurements the specimens in the United States National Museum will be recognized by simple cata- logue numbers, as 104362, without qualifying initials or footnotes; those in the British Museum by the well-known separation of these numbers into sections by means of periods, as 99. 6. 12.3; specimens in other museums will be found designated by appropriate and self- explanatory initial letters or by footnotes.

HISTORICAL ACCOUNT.

The earliest published. account of treeshrews is that of Ellis,! one of the surgeons of Captain Cook’s expedition. On Tuesday or Wednesday, 25th or 26th of January, 1780, Ellis remarks: ‘‘Our sportsmen * * * having seen only a few monkies, squirrels, and a cock and hen, the latter of which they shot. According to Linneus this island is their native place.’ The island referred to is Pulo Condore, off the coast of Cochin China. The squirrels men- tioned in the account are not squirrels, but Tupaias. One of them was evidently shot. A rough but very accurate sketch of the animal was made by Ellis and a Latin diagnosis of it written in his journal. This description of the animal was published by Gray in 1860.? Through the courtesy of the officials of the British Museum a repro- duction of a photograph of Ells’s drawing is here printed. There can be no doubt from Ellis’s picture or description that his squirrels were Tupaias (pl. 1).

Tupaias as such were first brought to the attention of the world by M. Diard, a French naturalist, at one time an assistant of Sir Thomas Stamford Raffles, in November, 1820, under the designation of Sorex glis.$

Six months later, May, 1821, the genus Tupaia was first proposed by Sir Thomas Stamford Raffles, and the species ferruginea and tana described, the latter in the present paper being made the type of a new genus.

Specimens of Tupaias had been seen by Europeans several years earlier, and one even sent to Europe. Geoffroy > remarks:

The discovery of this remarkable group of Insectivores has been attributed to both M. Diard and Sir Raffles. The fact is that it belongs to neither of these celebrated travelers, but to Leschenault de la Tour, who had sent in 1807 to the Museum of Paris an individual of the species which has since been called Tupaiajavanica. Never- theless it is only since 1820 that the attention of naturalists has been called to Tupaias, and that these animals have really entered the domain of science.

1 Voyage by Capt. Cook and Capt. Clerke in ships Resolution and Discovery, 1776, 1777, 1778, 1779, and 1780, vol. 2, 1782, p. 340. « 2 Ann. Mag. Nat. Hist., ser. 3, vol. 5, 1860, p. 71.

3 Asiatic Journ. Month. Reg., vol. 10, p. 478, November, 1820.

‘Trans. Linn. Soc. London, vol. 13, p. 256, May, 1821.

6 Belanger, Voyage aux Indes-Orientales, Zoologie, p. 104, 1835.

4 PROCFEDINGS OF THE NATIONAL MUSEUM. vou. 45,

Geoffroy was naturally quite unaware of the existence of Ellis’s © manuscript notes and drawings. Since Diard’s and Raffles’s time the group has become better and better known and its geographic range widely extended. The most important discoveries in regard to the group since 1821 have been the announcement of the genus Ptilocercus by Gray,! 1848, of the group now called Dendrogale by Schlegel and Miiller,? the discovery of treeshrews in the Philippine Islands by Whitehead, about 1879,’ now forming the genus Urogale, and the discovery of treeshrews in India by W. Ethot, about 1849.

DEFINITION AND RELATIONSHIPS.

The Tupaiide are diurnal insectivorous mammals characterized by a general squirrel-like aspect, more or less arboreal habits, orbits completely encircled by bone, alisphenoid canal present, malar bone with a more or less enlarged perforation, separate radius and ulna, and separate tibia and fibula, dental formula J 3 C4, Pm } M. 3, upper molars with typical W pattern. The family is composed of two very distinct groups for a long time regarded as genera, the typical members of the family, Zupaia and the aberrant Péilocercus. The old genus Tupaia has gradually been seen to be a composite genus, and up to the present time has been divided into three sepa- rate genera: Tupaia, Dendrogale, Urogale. In the present paper two more genera are recognized. These genera are now for the first time grouped to form the subfamily Tupaime. The single genus Ptilocercus is here regarded as forming the subfamily Ptilocercine. Hylomys of the Erinaceide was formerly associated with the tree- shrews, but was removed in 1874 by Anderson.*

TUPAIINS.

Tail bushy or close-haired throughout its entire extent.

Ears small and cartilaginous.

Footpads of moderate development.

Supraorbital foramen well developed.

Foramen rotundum entirely distinct from sphenoidal fissure.

Second upper incisor unicuspid.

Upper molars with well-developed bifur- cated mesostyles.

Upper molar teeth without a distinct cingulum.

Lower molar teeth without a cingulum.

PTILOCERCIN.

Tail with terminal portion distichously tufted, naked, and scaly basally.

Ears large and membranaceous.

Footpads relatively large and soft.

Supraorbital foramen absent.

Foramen rotundum confluent with sphe- noidal fissure.

Second upper incisor with a distinct posterior cusp.

Upper molars without mesostyles.

A distinct cingulum encircles the upper molar teeth.

Lower molar teeth with a cingulum on outer surface.

1 Proce, Zool. Soe. London, 1848, p. 23.

2 Verh, Nat. Gesch. Nederl. Overz. Bezitt., p. 167, 1839-44.

2 Thomas, Ann. Mag. Nat. Hist., ser. 6, vol. 9, p. 250, March, 1892. 4 Waterhouse, Proc. Zool. Soc. London, 1849, p. 107.

6 Trans. Zool. Soc. London, vol. 8, 1874, pp. 453-467.

No. 1976. TREESHREWS: FAMILY TUPAIIDH—LYON. 5 Genera and their types. Genus and its type.

Tupaia Rarrwes, 1821, Tupaia ferruginea. | Ptilocercus Gray, 1848, Pétilocercus lowit. Anathana, new, Tupaia ellioti.

Dendrogale GRAY, 1848, Tupaiamurina.

Tana, new, Tupaia tana.

Urogale MEARNS, 1905, U. cylindrura (=

T. everetti.)

The nearest relatives of the Tupaiide are the Macroscelidide, terrestial Insectivores of Africa. Many authors! place the two families in a superfamily or subordinal group, the Menotyphla or Tupaioidea as distinguished from all the other living Insectivores the Lipotyphla.

This grouping appears to me to be a natural one, and the differences that we now find between the Tupaiide and the Macroscelidide are in large measure due to the very different modes of life of the two families, the Tupaiide being quite arboreal in their habits, and the Macroscelidide, terrestrial! and saltatorial. The geographic distri- bution of the two families taken together show many resemblances to the present day distribution of the Tragulidz, rhinoceroses, elephants, anthropoid apes, Cercopithecide, and Megachiroptera, a circumstance lending some weight to their probable common origin. In spite of their great difference there is scarcely an osteological structure in the Macroscelidide that does not have some counterpart in the Tupaiide, and the opposite, the most conspicuous difference being the absence of the alisphenoid canal in the former and its presence in the latter, and the complete bony orbit of the Tupaiide absent in the African family. The skull of the Macroscslidide bears most general resemblance to that of Ptilocercus, and it is interesting to note that a supraorbital foramen is lacking in both, but is a con- spicuous feature of the Tupaiine. The main differential points between the two families are seen in the following table:

TUPAUDA. MaAcROSCELIDIDA.

Alisphenoid canal present. Alisphenoid canal absent. Supraorbital foramen present (except in | Supraorbital foramen absent. Ptilocercus).

Orbit completely surrounded by bone. Orbit not completely surrounded by bone, even postorbital processes lacking.

Radius and ulna separate bones. Radius and ulna fused.

Tibia and fibula separate bones. Tibia and fibula fused.

Metatarsals not unusually elongated. Metatarsals unusually elongated.

Premolars, 3 above and 3 below. Premolars, 4 above and 4 below.

Molars, 3 above and 3 below. Molars, 2 above and usually 2 below

(sometimes 3 below).?

1 Weber, Die Siugetiere, 1904, p. 377. Gregory, Bull. Amer. Mus. Nat. Hist., vol. 27, 1910, p.268. Gill,” Bull. Geol. Geogr. Surv. Terr., No. 2, ser. 2, May 14, 1875, p. 20. Osborn, Age of Mammals, 1910, p. 522.

2See Gregory, Bull. Amer. Mus. Nat. Hist., vol. 27, 1910, pp. 280-285; also Thomas (Proc. Zool. Soc. London, 1890, pp. 445, 446) who remarks on dentition of Petrodromus and the other genera.

6 PROCEEDINGS OF THE NATIONAL MUSEUM. vou. 45.

Although the general appearance of the molariform teeth of the Macroscelidide is quite different from the typical W-patterned teeth of the Tupaiide, yet it is easy to see how the teeth of the former may have been derived from those of the latter. The teeth of the Macroscelidide show a greater departure from the more typical tritubercular teeth of the Tupaiide, just as the limb bones have shown a greater departure from the normal.

OSTEOLOGY.

The skeleton of the Tupaiidz, as represented by the genera Tupaia and Tana, has been rather carefully studied by Blainville,* Mivart,? Anderson, and Gregory. In the British Museum is most of the skeleton of the type of Urogale evereiti, and in the Leyden Museum is a skeleton of Ptilocercus. I have not seen the latter, but Jentink® has published a few notes on it. It is the only skeleton of that genus that I know of existing in museums. I have not seen skeletons of the genera Anathana or Dendrogale, and know of none in collections. Skeletons of Tupaia are found in most of the larger museums, and in the United States National Museum are the following:

Cat. No. 124317, Tupaia glis ferruginea, Singapore.

Cat. No. 174609, Tupaia demissa, Sumatra.

Cat. No. 49468, Tupaia lacernata wilkinsoni, middle of Malay Peninsula.

Cat. No. 111782, Tupaia nicobarica nicobarica, Great Nicobar Island.

Cat. No. 154593, Tupaia javanica, western Java.

Cat. No. 174611, Tana tana tana, Sumatra.

The observations on the skeleton which follow are based upon an examination of these skeletons of the genera Tupaia and Tana and skulls of the other genera. I have also made free use of the observa- tions of Mivart, Anderson, and Gregory.

Skull—The skull of the genus Tupaia is characterized by its rather generalized structure; it is widest just posterior to the middle, and tapers toward either extremity both laterally and supero-inferiorly, the tapering being much more pronounced anteriorly, especially so in the genera Tana and Urogale; posteriorly the skull is gently rounded off. The brain case is relatively large and inflated and widest at the zygomatic roots. The orbit is completely surrounded by bone, is large, directed mainly laterally but at the same time slightly inclined upward and forward. Posterior to the orbit is a temporal fossa of moderate size. The temporal ridges are rather prominent and dis- tinct except for a short distance in front of the lambdoid crest, where they unite to form a short sagittal crest. In Ptilocereus the temporal

1 Ostéog. Mamm. Insect., 1840, pp. 31-35. 2 Journ. Anat. Physiol., vol. 1, 1867, pp. 292-295, and vol. 2, 1868, pp. 145-146. 3 Zool. Res. West. Yunnan, 1879, pp. 108-123.

4 Orders of Mammals, Bull. Amer. Mus. Nat. Hist., vol. 27, 1910, pp. 269-280. 6 Notes Leyden Museum, vol. 7, 1885, p. 7.

No. 1976, TREESHREWS: FAMILY TUPAIIDAI—LYON. 7

ridges remain separated and in Urogale they unite early to form a much more conspicuous sagittal crest than they do in Tupaia. The lamb- doid crest is well marked and gently arched. It begins faintly on either side near the external auditory meatus and becomes well developed along the upper border of the supraoccipital. The palate is long but neither specially wide nor specially narrow. In front are well marked anterior palatine foramina; posteriorly the palate is slightly concave, and ends in a slightly thickened ridge, and a very small blunt median spine. The most anterior part of the posterior edge is about on a line with the posterior edge of the last molars. In the posterior half of the palate in the genera Tupaia and Tana are usually irregular vacuities. The other genera, Urogale, Anathana, Dendrogale, and Ptilocercus, are usually without defects of ossifica- tion in the palate. The external pterygoid fossx are large, short, and wide, formed by the well marked, pointed, and slightly directed inward pterygoid bones, and the pterygoid plate, rather short and triangular, of the alisphenoid. The choane are rather wide, and narrower between the pterygoids than anteriorly. The bull are of moderate size and formed of the endotympanic. The outer edge of the bulla is produced outward so as to cover up or enclose the tym- panic ring or ectotympanic. The small foramen ovale is almost covered over by the antero-external edge of the bulla. The glenoid fossa is rather wide and shallow and limited in front and behind by short and inconspicuous anterior and posterior glenoid processes. Only the minutest trace of a paroccipital process is present. The alisphenoid is pierced by an alisphenoid canal. The foramen magnum is directed downward and backward.

The external opening of the infraorbital canal is situated above the second premolar. In Ptilocercus the canal is much shorter and its external opening is over the last premolar. The internal opening of the canal lies shortly inside the orbit. The lachrymal canal has its opening in a distinct notch except in Ptilocercus and is rather more outside of the orbit than inside of it. Except in Ptilocercus there is a conspicuous supraorbital foramen at the upper outer angle of the orbit, continuous with a groove under the edge of the roof of the orbit. Except in Ptilocercus the optic foramen is separated from the sphenoid fissure by a narrow spicule of bone, and the foramen rotundum lies at the base of the external pterygoid plate. In Ptilocercus the optic foramen is separated from the sphenoid fissure by a broad bridge of bone and the foramen rotundum is blended with the sphenoid fissure. The foramen ovale is situated almost under the antero-outer edge of the bulla except in Piéilocereus where the opening of the foramen is plainly visible in front of the bulla. In