Full text of "Proceedings of the United States National Museum"

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SMlTHSOiNIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM

PROCEEDINGS

OF niE

UNITED STATES NATIONAL MUSEUM

VOLUME 96

UNITKD STA'n-.S

GOVERNMENT PRINTING Of KICK

WASHINGTON : lO^S

The scientific publications of the National Museum include two se-
ries, known, respectively, as Proceedings and Bulletin.

The Proceedings, begun in 1878, are intended primarily as a medium
for the publication of original papers, based on the collections of the
National Museum, that set forth newly acquired facts in biology, an-
thropology, and geology, with descriptions of new forms and revisions
of limited groups. Copies of each paper, in pamphlet form, are dis-
tribui-ed as published to libraries and scientific organizations and to
specialists and others interested in the different subjects.

The dates at which these separate papers are published are recorded
in the table of contents of each of the volumes.

The present volume is the ninety-sixth of this series.

The Bulletin, the first of which was issued in 1875, consists of a
series of separate publications comprising monographs of large zoologi-
cal groups and other general systematic treatises (occasionally in
several volumes), faunal Avorks, reports of expeditions, catalogs of
type specimens, special collections, and other material of similar nature.
The majority of the volumes are octavo in size, but a quarto size has
been adopted in a few instances in which large plates were regarded
as indispensable. In the Bulletin series appear volumes under the
heading Contributions from the United States National Herharivmi,
in octavo form, published by the National Museum since 1902, which
contain papers relating to the botanical collections of the Museum.

Alexander Wetmore,
Secretary, Smithsoniam. Institution.

^iPis

CONTENTS

Page

BoHAHT, G. E. The phorid flies of Guam. No. 3205. Febru-
ary 17, 11)47^ 397-416

New genus: Parafannia.

New si)ecies : Mcgasclia {.Ucgusdia) Hctifimur, J/, suis, M.
stuntzi, M. parabasiscta, Chonoccphalus hirmitus, C. subglaber,
PuUciphora wj/inani, P. nigriventru, Parafannia molluscovora.

C'aldweix, John S. Neotropical lanternflies of the genus
Phinctus in the United States National Museum, with de-
scriptions of four new species. No. 3194. May IG, 1945 ^__ 177-184
New siiecies : Phrirtus aordidufi, P. minulacanihis, P. punctatus,
P. regalis.

( "lakk, Austin H.. and Zetek, Jaaies. The onychophores of
Panama and the Canal Zone. No. 3197. February 21,
194G ^ 205-213

CusHMAX, R. A. The ichneumon-flies of the genus Gryp-
tanura Brulle. mainly tropical American. No. 3193. May
23, 1945^ 139-170

New genus: CremnocryptuH.

New species: Cryptunura lubcrculdta, V. dicostuta, C. quadri-
mncuUita, C. mrdiosirigosa, C. bicarinata, C. piccothoraoB,
C. planiscutcllata, C. polUigaster, C. conica, C. ruficeps, C.
scptctitrionulin, C. apophysis, C. biliiicata, C. rufa, V. lineati-
fi'tiiiir, C. coxata, C. boUvicnsis, C. isthmus, V. cxcalibur, C.
aviridccs, C grucilipcs, C. gracilis, C. gcnulis, (J. parancnsis, C.
trnuiterebrata, V. mnnilifroiis.

New combinations: Cryplnnuni mcxicana (Cres.son), C. uriza-
bcnsis (Cameron), V. propinqua (Cresson), C. prciiosa (Vier-
eck), V. vurirgnla (Hrulle), V. inccrta (('resson), Vrcmnocryp-
tus spiiiifcrus (Cameron), C. cingulatus (Tosquiuet).

. A generic revision of the ichneumon-flies of the

tribe Ophionini. No. 3200. July 17, 1917' 417-482

New gciK'ra : AUiji-h'iphiftu, Potophion, Sintoithinn, 'I'roplitipfiion,
Clistorapha, Jforthonrura, Chilophion, Aulophion.

New species: Auslralophinn iit/latus, Potophion cnudatus, Simo-
phion aicnrinatuH, Trophophion tcniiicrps, liocthonvura arida,
Htauropoctonus chczanus, Aulophion bicarinatus, A. vxcarinu-

tU8.
.New CDmliiiiatlDiis : l{!i i/iirlKijihion /I'luniiipt ntiis ( AsIiiik'JkI ) , J\'.
ligulifrr (Morlcy), Aglnophion purpurnsn uh (Smilli), A.
su)i\'ilriinum (Endt-rleln ), CliHtoruplin suhfuliginosa (Asli-
nn'iMl). Ginophion costalis (<;r(;sson), Vliihtjihion iihnoruiis
(Felt), Sliiuroii'>rt(,nii.H raricgaltts (Ucbldii).

I)at<- of iiubllrntluii.

IV PROCEEDINGS OF THE NATIONAL MUSEUM vol.86

Page

Davis, A. C. Review of the weevils of the tribe Ophryastini

of America, north of Mexico. No. 3207. July 8, 1947 ^ 483-551

New species: Eupagoderes griseus, E. aeneus, E. robustus, E.

pilosus.
New variety : Eupagoderes morUvallis approximatus.

Fennah, R. G. New lanternflies (Fulgoroidea) from South

America. No. 3189. May 9, 1945 ^ 95-104

New genus : Iquitosa.

New species : Pintalia quadrimaculata, P. marmorata, P. oMiqui-
vitta, P. falcata, P. curvivitta, P. daedala, P. vomerifera, Iqui-
tosa shannoni, Ateson seniiluteum, A. luteospersum.

Fisher, Walter Kenrick. Echiuroid worms of the North
Pacific Ocean. No. 3198. April 11, 1946 ^ 215-292

New orders: Xenopneusta, Heteromyota.

New genera : Bonclliopsis, Euhonellia, Nellobia, Lissomyema.

New si)ecies: Thalassema steinhecki, Listriolobus pelodes,
OcJietostoina octomyotutn, O. edax, Arhynchltc inamoenus,
Bonelliopsis alaskana, Eubonellia valida, Nellobia eusoma,
AcantJiohamingia parudola.

New subspecies : Echiurus echiurus alaskanus.

Fisher, W. S. New beetles of the family Eucnemididae from
Central America and the West Indies. No. 3188. May 8,

1945 1 79-93

New genera : Neodiapodius, Hylotastella.

New siiecies : Temnillus aspericollis, Neodiapodius buscki, Dro-
maeolus pulcher, D. panamensis, Fornax poeyl, F. valerio,
Plesiofornax nigrinus, Farsus convexus, ArrMpis cubanus, A.
insularis, Dirhagus albofasciatus, Hylotastella schwarzi, Nema-
todes exiguus.

■ — . New cerambycid beetles belonging to the tribe

Disteniini from Central and South America. No. 3201.

November 26, 1946^ 329-333

New species : Distenia lateralis, D. spinipennis, Cometes etnar-
ginata, C. bicolor.

Gahan, a. B. Eight new species of chalcid-flies of the genus
Pseudaphycus Clausen, with a key to the species. No. 3200.

November 22, 1946 ^ 311-327

New species: Pseudaphycus mcritorius, P. abstrusus, P. malinus,
P. anyustifrons, P. nmndus, P. Umatulus, P. meracus, P. alveo-
latifrons.

. Review of some chalcidoid genera related to

Cerocephala Westwood. No. 3203. December 31, 1946 i__ 349-376
New subfamily: Cerocephaliuae.
New genus: Acerocephala.

* Date of publication.

CONTENTS V

ruge

Gahan, a. B. — Continued

New species: Choctos;)U(i tabida, Accroccphala acnif/ma.

New combinations: CIkx toKitila f rater (Girault), Th<ocoIax
bakeri (Crawford), 'J heocolaxia viridinotnm (Dodd), T. insu-
laris (Dodd), T. insuluris var. grandis (Dodd), T. perpuhhra
(Dodd), 2'. pcrpulchra var. metaUica (Dodd), T. scoUjtivora
(Ashiuead), T. pitijophthori (Aslnnead), Accroccphala atrovio-
lacea (Crawford), Eupclinella canadouHis (Provancher).

Gazin, C. Lewis. Machacroidcft cothen Matthew, the suber-
tooth creodont of the Bi-idger Eocene. No. 3202. December
16, 1946^ 335-347

GujMore, Ciiarijjs W. Notes on recently mounted reptile
fossil skeletons in the United States National Museum. No.
3106. Fi'bruary 5, 1046^ 195-203

. The osteoloory of the fossil turtle Testudo

'pracextans Lambe, Avitli notes on other species of Testudo
from the Oligocene of Wyoming. No. 3199. March 28,
1946^ - 293-310

GoijjMAN, E. A. (See under Kellogg, Remington.)

Heinricii, Carl. The genus i^wTu/eZZaZeller: A contribution
toward a revision of the American pyralidoid moths of the

family Phycitidae. No. 3190. May 18, 1945^ 105-114

New species : Funddln iiinohUia.

Kellogg, Remixgtox, and Goldman, E. A. Review of the

sjjider monkeys. No. 3186. November 2, 1944 ^ 1-45

New .sui)si)ecies : Afcks gcoffroyi yucatanensis, A. g. panamensis.

MiiJ.ER, Robert R. Hy porhamphiut patris, a new species of
hemiramjihid fish from Sinaloa, Mexico, with an analysis of
the generic characters of IhjporJiamphus and Ileniirani-

phus. No. 3195. July 27, 1945^ 185-193

New species: llyporluiinyltus jxitria.

SciiULTZ, Leonard P. A revision of the American clingfishes,
family Gobie.socidae, with descriptions of new geiioi-a and
forms. No. 3187. December 30, 1944 ^ 47-77

New gen«'ra : Acyrtun. InfratridniH, Afcon.

New spe(!ies : Sicj/ascn hildchrandi.

New sub.species : Cotylis niyri])ittnin woodxi.

. A new genus and two new s[)ecies of i)erc()id

fishes from New Guineji. family Centropomidae. No. 3191.

March 6, 1945 ' 115-121

New Bt'nu.s : Xcnamhu.iHia.

New HfMi'cle.M : Xtndtnhassis honcHni, X. Himnni.

' Date of puljllcatloii.

VI PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Page

ScHULTz, Leonard P. Three new sciaenid fishes of the genus
Ophioscion from the Atlantic coasts of Central and South
America. No. 3192. April 25, 1945 ^ 123-137

New species: Ophiosoion brasiUensis, 0. venezuelae, 0. pana-
mensis.

. A revision of the genera of mullets, fishes of the

=■5

family Mugilidae, with descriptions of three new genera.

No. 3204. December 5, 1946 ^ 377-395

New geuera : Xenomugil, Crenimugil, Heteroitiugil.

Zetek, James. (See under Ci.ark, Austin H.)

^ Date of publication

ILLUSTRATIONS

PLATES

Follow I iig
page

1. Lower side of head of Votylis microi^pUus (ri.wlcr) and of (\

nigripinnis niyripinnifi I'oters 64

2. New South Aiuoiicau FiilKoroidea 104

3. South American Fulgoroidea 104

4-6. The pMius FundcUa 114

7. Cephalic process in Phrictus 180

8. Valvula and geuitalia in Phrivttis 180

0,10. Species of Phrictus: Dorsal aspect 180

IL Hyporhamphua patris, new species 193

12. Mounteil skeleton of Crocodiliis clavis Cope 203

l.S. Skull and k)wer jaw of Crorodilus clavis Cope 203

1 1, ir». Skull of Crocodiliis clavis Cope 203

16. I»isea.sed dorsal vertebrae of Crocodilns sp 203

17. Tail, pelvis, hind linihs, and feet of Corythnsaurus casuarius Brown_ 203

18. Skin impre.ssions of Conjthosnunis casuarius Brown 203

19. Restoration of Cori/thosaurus casuarius Brown 203

20. IJchiurus cchiurus alaskanus, new subspecies 292

21. Listriolobus pelodes, new species, and Ochetostoma octomyotum,

new species 292

22. Listriolobus pelodes, new species 292

23, 24. Ochetostoma octomyotum, new species 292

2r». Arhynchite iuamoetius, new species 292

26, 27. IloneUiopsis aUiskatia, new genus and species 292

2S. Euhonellia ralida, new frenus and species 292

20, ;}0. Xellohid cusoma, new genus and species 292

;{1,32. Acunttiohaminf/ia paradola, new species 292

^'.i-?,~). Frechis enupo Fislier and MacClinitie 292

1^6. Urechis caupo, F. chilcnsis, and U. unicinctua 292

37. Frcchis caupo and comniensals 292

38-41. Tcstudo praeextans Lanibe 310

42,43. Testudo laticunea Cope 310

44. Testudo f/uadrala Cope; Testudo praeextans Lambe 310

45-46. Mnrhnrroidrs cothen Matthew 347

47. Si>f'cies of Choetospila, Thcocolax, Cerocephala, Acerocephala, and

Theocolaxia 376

48. Species of Thcocolax, Cerocephala, and Acerocephala 376

40-.''>6. Ichrienmon-flios of the tribe Ophionlnl 482

Ti:XT FICTIRES

PnKo

1. Distribution of the forms of spidernionkeys (Atcles) In South Amer-

ica (map) 12

2. Distribution of the forms of spider monkeys (Ateles) in Middle

America (map) 14

3. Holotype of Xmnmhassis honcHsi, new species 119

4. IIolotyi)e of Xctuimbassis simoni, new speci»'H 121

f). Diagranunatic sketches of snout and lower Jnw of Bairdiella chrys-

ura fLacep<>de), Stellifer rastrifer (Jordan), and Ophiosdon

typicus Cill 12.'»

6. ITolotyi»e f»f Ophi'iscion hrasilicnsis, new species 129

7. Iliilotyjx' of (tjiliioscitni rcnczurhic, new species 132

8. JIololyp<' of Ojihi'iscioti paunmcnsis, new species 13.'>

9. Sketch of hearl regions of Ifyporhamphus and Ifcmiramphwt 180

VII

VIII PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Page

10. Lissornyema mellita (Couu) 223

11. Thalassema steinbecki, new species 231

12. Ldstrioloius pelodes, new species 235

13. Listriolohus pelodes, new species 237

14. Ochetostoma edax, new species 244

15. BonelUa viridis : 251

16. BonelUopsis alaskana, new species 252

17. Urechis caupo Fisher and MacGiuitie 266

18. Urechis caupo Fisher and MacGinitie 267

19. Urechis caupo Fisher and MacGinitie 273

20. Skull of Testudo pracextans Lambe 294

21. Skull and lower jaw of Testudo praeextans Lambe 295

22. Palatal view of the skull of Testudo praeextans Lambe 296

23. Lower jaw of Testudo praeextans Lambe 297

24. Carapace of Testudo praeextans Lambe 298

25. Plastron of Testudo pi-aeextans Lambe 303

26,27. Pelvis of Testudo praeextans Lambe 304

28. Sketch of the maxillary, premaxillary, and preorbital bones of

Mugil cephalus 388

29. Sketches of the maxillary, premaxillary, and preorbital bones of

Myxus elonfjatus 390

30. Sketches of the maxillary, premaxillary, and preorbital bones of

Chelon cheio 390

31. Sketches of the maxillary, premaxillary, and preorbital bones of

Trachystoma petardi 393

32. Diagram of the possible I'elationships of genera of the Mugilidae 395

33-35. Phoridae of Guam 400

36-38. Phoridae of Guam 401

39-41. Phoridae of Guam 404

42-45. Phoridae of Guam 405

46-48. Phoridae of Guam 408

49. Genital structures of Eupagoderes cf. deserttis-caUfornicus sec-

tion 486

50. Sapotes puncticollis Casey 489

51. Eupagoderes geminatus Horn 497

52. Eupagoderes griseus, new species 499

53. Eupadogeres aeneus, new species 500

54. Eupagoderes mortivallis Fall 502

55. Eupagoderes rohustus, new species 504

56. Eupagoderes decipiens (LeCoute) 505

57. Eupagoderes spcciosus (LeConte) 510

58. Eupagoderes niarmoratus Fall 512

59. Eupagoderes sordidus (LeOonte) 514

60. Eupagoderes simulans Van Dyke 515

61. Eupagoderes argentatus (LeConte) 516

62. Eupagoderes lucanus Horn 518

63. Amydrogmus variabilis Pierce 521

64. Ophryastes tvickhami Sharp 523

65. Ophryastes latirostris LeConte 526

66. Ophryastes symmetricus Fall 528

67. Ophryastes sulcipennis Casey 529

68. Ophryastes sulcirostris (Say) 530

69. Ophryastes porosiis LeConte 532

70. Ophryastes tuberosus LeConte 534

71. Ophryastes ovipennis Sharp 535

72. Ophryastes vittatus (Say) 537

73. Tosastes coarctatus Champion 540

74. Tosastes globularis Pierce 542

75. Tosastes ovalis Pierce 544

76. Tosastes cinerascens Pierce 546

77. Rhigopsis effracta LeConte 548

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued n^t''t\»>L^JH*j;l ^V <^<

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM

Vol. 96 WMhington: 1944 No. 3186

REVIEW OF THE SPIDER MONKEYS

By Remington Kellogg and E. A. Goldman

Field studies have shown that the forest-inhabiting spider monkeys
of the New World are «nl)jpct to ])lasmodial infections under nat-
ural conditions and consequently, since they are readily obtainable.
are now being used in laboratory studies of the malaria Plasmodium.
Spider monkeys are readily tamed and very adaptable to a life in
captivity. Owing to their d(jcility they are easily handled by labo-
ratory assistants, and it is not difficult to provide suitable vegetable
fofxl for them. For these reasons it is important to establish, if pos-
sible, the valid geographic races of these monkeys, to determine in so
far as available collections permit the geographic ranges of the recog-
nized forms, and to set forth tlie characteristics by which the several
races can be recognized with s(jme degree of certainty.

The spider monkeys form a compact group, or subfamily, the
Ateliiiae, of the primate family Cebidae, 'I'liey are characterized by
slender body, very long, slender limbs, and long tail, naked beneatli
distally and prehensile; the hand has only four functional fingers, the
thumb being usually vestigial or absent.

These monkeys, in structure and hai)its, sliow higli specialization
and adaptation to a strictly arboreal life. They arc limited in range
to tlie unbroken tropical ff)rests from southern Mexico to northern
Matto Gnjsso, Brazil, and central Bolivia in South America. In the
forest areas inhabited, oflen including steep mountain slopes, largo
trees have interlocking branches through which the spider monkeys
are able to travel in long leaps at amazing speed, using the tail con-
stantly as a powerful grasping organ for balancing, and swinging to
differing levels. A spider monkey in a cage is seen at a disadvantage

2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m

compared, for example, with so active an animal as the gibbon, but
the reason for this seems to be that room is lacking for the exercise of
its full powers. It is interesting to speculate on the comparative
speed that might be attained under natural conditions by these widely
different but both outstandingly active New World and Old World
representatives, respectively, of the order to which they belong. In
following spider monkeys in the forest a man may keep up by run-
ning if the ground is clear, but if impeded by having to go around
underbrush he is quickly left behind. Spider monkeys are considered
very good to eat by the native populations of many countries and
are, therefore, in danger of extermination.

Many names for spider monkeys have been based on inadequate de-
scriptions of animals from unknown localities, and if a type specimen
was used it may no longer be extant. The result has been great con-
fusion in the literature bearing on the identification of material, and
of the distribution of the nominal species recognized little has been
known.

The status even of the generic name Ateles has been open to ques-
tion. The spider monkey S[imia\ Sapajus paniscus was included
among the 14 monkeys allocated to Sapajus when this subgenus was
proposed by Kerr (The Animal Kingdom, CI. I, Mammalia, p. 76,
1792). No genotype seems to have been designated for the subgenus
Sapajus Kerr, unless the statement made by J. A. Allen (Bull. Amer.
Mus. Nat. Hist., vol. 7, p. 181, June 20, 1895) that Sapajios of Kerr is
equivalent to Cebus Erxleben, 1777, can be interpreted as restricting
the application of the term Sapajus to the species of Gehus included
among the 14 monkeys mentioned above. In order to eliminate any
possible misinterpretation, the genotype of Sapajus Kerr is here desig-
nated as S\i7nia'\ Sapajus capuclnus Kerr {op. cit., p. 78, 1792), which
is the same as S[i77iia'] capiwina Linnaeus., 1766 {nee 1758). Inasmuch
as the genotype of Gehus Erxleben, 1777, has been fixed (Elliott, Field
Columbian Mus. Publ. 115, zool. ser., vol. 8, p. 560, Mar. 4, 1907) by
subsequent designation as Sitnia capucina Linnaeus, 1766 {nee 1758),
which in turn is equivalent to Gehus nignvittatus Wagner (see
Cabrera, Rev. Soc. Argentina Cienc. Nat., vol. 16, pp. 21-22, 1939),
Sapajus Kerr is herewith relegated to the synonymy of Gehus Erxle-
ben.

Neither opinion 147 (On the principles to be observed in interpreting
article 34 of the International Code in relation to the rejection, as homo-
nyms of generic and subgeneric names of the same origin and meaning
as names previously published. Opinions and declarations rendered
by the International Commission on Zoological Nomenclature, London,
vol. 2, pt. 14, pp. 123-132, Sept. 30, 1943) nor article 34 (International
rules of zoological nomenclature, Proc. Biol. Soc. Washington, vol.
39, p. 86, July 30, 1926) contains an express ruling as to whether a

THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 3

ijiven oreneric nainc is a hoinonyni oi' anolluT piwiously jmblislied
ofenerio name that has the same ori<2:iii and meaning but that dill'ers
from the hitter in more than one letter. If the "one letter" rule is
strictly adhered to, then Sopajou Lacepi'de (Tableau des divisions,
sous-divisions, ordres et genres des mannniferes, p. 4. Published as
supplement to Discours d'ouverture et de cloture du cours d'histoire
naturellc, et tableaux nu'thodiques des mannniferes et des oiseaux, 1799)
is the oldest available name for the spider monkeys. The genotype of
Sapajou Lacepede is Sapaj&u paniscus Lacepede=/S'/wMf paniscus Lin-
naeus. Pending an opinion from the International Commission on
Zoological Xomenclature, the generic name Ateles E. Geoffroy (Ann.
Mus. Hist. Nat. Paris, vol. 7, p. 2G2, 1806) , of which the genotype is
likewise Simla paiiiscus Linnaeus, is here retained for the spider
monkeys.

We have been unable to reach any satisfactory conclusions regarding
the identity of one of the descril)ed spider monkeys allocated to this
genus. The '"Antigua Monkey" was described by Pennant (His-
tory of quadrupeds, vol. 1, p. 206 (no. 212), 1781) as follows :

Mfonkey] with a short nose; black face, hair on each side long; back and sides
orango and black, intimately mixed ; holly white; outside of lops black ; inside ash-
I'olorotl ; tail of a dusky ash; its length 20 inches; that of body oightocn.

Lately in possossion of Richard Morris, Esq. of the Navy-Offico; brought from
Antigua; but its native place uncertain; very good-natured, lively, and full of
tricks; frequently hung by its tail.

This monkey subsecpiently became the basis for [^Sinua] Sapajus
variefjatuH Kerr (The Animal Kingdom, p. 79, 1792) and Simia antiqu-
ana Pechstein (Thomas Pennant's Allgemeine Uebersicht der vierfiis-
sigen Thiere, vol. 1, p. 227, footnote, 1799). In our judgment these
names apply clearly to a spider monkey, and quite likely to one of the
Middle American subspecies, but since the characters given are not
sufficient to identify the race positively, we believe both shoidd be held
unident ifiable. '^\n<ni Atclcs variegatws (Kerr) antedates .1 tchfi va/ic
fjahiJi ^^'agner, the latter being preoccupied must be supplanted by
Ateles hehehutli E. Geoffroy, which also has priority.

Despite strongly contrasting patterns of color, dilfering lengths of
pelage, and to some extent varying tail and foot proportions, the strik-
ing similarity of all the spider monkeys in the more essential cranial
features seems to indicate close reiationsiiips. The nominal sjxries
arc here reduced to foiii-. mimI it seems to us probable that additional
study of the genus based on moiv complete collections may result in a
further le.ssening of the number. The most divergent forms are the
I'lack paniscus of French Ciuiana and the silvery-bellied yucatanensis
(jf Qiiintana Koo. Perhaps the most clearly defined line of demarca-
tion between species, as we understand them, is in eastern Panama,
where the range of the deep reddish panamcnsM, a member of the

4 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

geoffroyi group, meets or closely approaches the range of the nearly all
black Tohustus. The Rio Amazonas, however, seems to be an effective
barrier separating the ranges of A. paniscus paniscus and A. helzebuth
marginatiis.

Several large series of specimens from the same locality have
afforded us much information concerning the range of individual
variation that may be expected in the genus Ateles. The thumb is
normally represented by a short metacarpal, which ordinarily is not
visible externally. When, in occasional individuals, a short proximal
phalanx is retained it constitutes the vestigial nailless thumb. In
two specimens of velleroms (U. S. N. M. Nos. 74662, 74663) from
Santa Efigenia, Oaxaca, and in one of the same form (A. M. N. H.
IS'o. 123282) from Cantoral, Honduras, these thumbs are present on
one hand and not on the other. In one of pananiensis (A. M. N. H.
No. 141980) from Canas Gordas, Costa Rica, the thumbs are present
on both hands. In a specimen of hyhridus from Guaimaral, Colom-
bia, vestigial thumbs on both hands bear small nails. Lonnberg
(Arkiv for Zool., vol. 32A, No. 25, p. 8, July 18, 1940) describes a
well-developed thumb with a well-developed nail on one hand only
in a specimen of paniscus^ a small thumb present on the other hand
being nailless. All three specimens of paniscus (A. M. N. H. Nos.
94134, 94135, 94136) from Rio Jamunda, near Faro, on the north bank
of the Rio Amazonas, have thumbs on both hands, as already re-
corded by Tate (Bull. Amer. Mus. Nat. Hist., vol. 76, p. 215, Oct. 20,
1939). Two others from Lago Cuipeua, farther east on the same
bank of this river, however, are thumbless. Lonnberg (op. cit.)
refers to the absence of thumbs in specimens of Ateles ater
[= chameh] from various localities, but they are present on both
hands in a skin (C. M. No. 2775) from Rio Yapacani, Santa Cruz,
Bolivia. The evidence thus presented indicates that vestigial thumbs
occur irregularly throughout the genus, and no specific significance
is attached to their presence or absence.

Certain cranial details believed by some authors to represent dis-
tinctive characters prove to be too variable to be of much value. As
Lonnberg [op. cit.) has pointed out, some of the features mentioned
as characters by Tate {op. cit.) in comparing species are subject to
a wide range of variation. The antorbital or "malar foramen" varies
from one or two "pin-holes" to rounded openings 5 mm. in diameter
through the jugal. These often differ in size and number on the
two sides of the same skull. Tate also refers to "small, triangular
pterygoids without pointed tips" as a cranial character in paniscus.
In the Cebidae the fossae between the true pterygoids and their ex-
ternal reduplications are large and deep in Cehus., reduced to a ves-
tige in Alouatta., and entirely wanting in Ateles. The external
reduplications of the pterygoids have the tips broken off in the

THE SPIDER MUNKEYS KELLOGG AND GOLDMAX 5

spei-inieiis of paiif.fcus exiuninotl by Tate, but tlu'^^e wiii^like thin
plates are in reality lar<ier and more extended posteriorly in these
and in other specimens of pan>,'<ciis from (he Amazonian refrion than
usual in this genus. Other characters that have been ascribed by
Tate to paniscus but that have not been found to be constant are the
small size of post<ilenoid jjrocesses, unexpanded zy^jfoniata of nearly
uniform depth, and narrow teeth. The maxillary tooth rows in the
^^enus as a whole are normally straight and convergent anteriorly,
although skulls with parallel or arcuate tooth rows are less frequently
observed. All the teeth in the maxillary tooth rows may be large
or small, and the individual teeth vary in size in relation to one
another. The posterior molars appear to be the most variable.
They are usually similar to the first premolars in crown area but
may be either larger or smaller. Examination of a large number of
spider-monkey skulls shows that the cranial and dental features men-
tioned are subject everywhere to about the same wide range of indi-
vidual variation. Schultz (Journ. Mammalogy, vol. 7, No. 4, pp.
28G-305, Nov. 1926) has discussed the relative ranges of variation in
skull measurements and in the formation of the forehead in a series
of Atelen taken in the immediate vicinity of one camp on the bank
of the Rio Yoya in Nicaragua.

Beyond the general similarity and the range of the usual individual
variation in the forms of Atrles there are, however, a few cranial
features that seem to be worthy of consideration. All the races of
Ateles geoffroyi agree closely in the more important cranial details,
but in the more northern subspecies vellerosus and yueatanens-ia the
brain case is slightl}' narrower than in panamensis and other Middle
American races. The South American species A. paniscv-f and ^1.
hrlzehiith have a longer rostrum, a more strongly inclined backward
facial piofile, and moie elongated nasal openings than geoffroyi.
In j)anisf'Uf<, which api)ears to be the most divergent in cranial de-
tails, the pf)stei-ior upi)er molars scarcely reach the transverse plane
of the anterior roots of the zygomata, while in geoffroyi they nonnally
jjass well beyond this plane; other j)ecidiar chaiaeters in pcmisms
are the broad, somewhat swollen base of the rostrum and the poste-
rior exten.sion of the external reduplications of the pterygoids. In
cranial details, as well as in geogi-a|)hic position, At< Us fii.sc/a ps ru-
bu.shiM is somewhat intermediate iM'tween geoffroyi wwd jxniiftcus.

Specinu'ns examined in series have shown tliat altliough sjnder
monkeys present considerable variation in color the vaiiation is
within limits beyond which it does not normally pass, except as the
usual intergradation that may be expected Ix-tween regional races or
subspecies. The coloration, length of ptdage, and othei* external
features, when determined by examination of specimens in suflicient
numbers, prove to be more reliable indices to subspecific relationship

6 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

than any cranial characters found by us. In Middle America, for
example, skulls of vellerosus from Mexico are distinctly narrower,
with less noticeable widening of the brain case, than those of fana-
me^isis, but differ little in other respects and despite great differences
in color are obviously assignable to the same species. Unlike many
other primates the sexes in the spider monkeys differ little, if at all,
in size. One of the largest skulls of geoffroyi is that of an old female
from Lavala, Nicaragua.

For loan of pertinent material our thanks are tendered to the fol-
lowing: Dr. Thomas Barbour, director, and Barbara Lawrence, Mu-
seum of Comparative Zoology, Cambridge, Mass.; Dr. Adolph H.
Schultz, Laboratory of Physical Anthropology, Johns Hopkins
Medical School, Baltimore, Md. ; Dr. H. E. Anthony and George G.
Goodwin, American Museum of Natural History, New York, N. Y. ;
J. Kenneth Doutt, Carnegie Museum, Pittsburgh, Pa.; Dr. H. C.
Oberholser, Cleveland Museum of Natural History, Cleveland, Ohio;
Dr. William H. Burt, University of Michigan, Museum of Zoology,
Ann Arbor, Mich., and to the latter also for the opportunity to
examine specimens in the Donald R. Dickey collection.

In lists of specimens examined, the following abbreviations are
employed :

A. H. S., Collection of Dr. Adolph H. Scbnltz.

A. M. N. H., American Musenm of Natural History.

C. D., Collection of Donald R. Dickey.

C. M., Carnegie Museum.

C. M. N. H., Cleveland Museum of Natural History.

M. C. Z., Museum of Comparative Zoology.

U. M. M. Z., University of Michigan, Museum of Zoology.

U. S. N. M., United States National Museum.

All measurements are in millimeters. The external measurements
were taken in the flesh by the collector as follows : Total lengthy nose
to end of terminal vertebra; tail, upper base of tail to end of terminal
vertebra; hind foot, back of heel to end of longest nail on digit. The
following cranial measurements of typical adults, unless otherwise
stated, were taken with vernier calipers by the authors: Greatest
length, distance from anterior tip of premaxillae to inion or extreme
posterior median point of brain case ; orhital width, distance between
outer margins of orbits; 'postorbital constriction, least width at con-
striction behind orbits; width of hrain case, greatest width of brain
case at or over mastoids ; zygomatic hreadth, greatest distance between
outside surfaces of zygomata ; inaxiUary tooth row, distance from front
of canine to back of posterior upper molar at alveolar borders.

The following treatment is based on a review of the scattered lit-
erature and an examination of 251 specimens representing all the
recognized species. The work has been done at the request of the
Board for the Coordination of Malarial Studies of the National Re-

THE SPIDER MONKEYS KELLOGG AND GOLDMAX 7

search Council in the hiboiiitorics of the Division of Miimnials luid of
the V. S. Fish and Wildlife Service in the United States National
Museum. The nuips were drawn by Mrs. Kiithoryne C. Tubb, of the
Fis-h and Wildlife Service. This review is inteniled to afford a work-
ing knowledge of the genus, pending more exhaustive revision as the
species become better known.

KEY TO SPIDER MONKEYS (GENUS ATELES) '

A. Gi'iieral coloration of entire back ehieflj' black.

B. Back and belly, a.s well as lin/bs ami tail, wbolly deep glossy black.

C. Hairs on bead wholly black (except for short, white mustache whiskers

on upi)er lip and sparse, short, white chin whiskers) ; pelage deep glossy

black.

D. Foot rather larj^e (length, hi^l to end of longest toe, 190-220 mm.).

E. Tail normally bushy in adult (especially on upper ba.sal half, longest

hail's on upperside .50-100 nun. in length) ; face normally flesh color,

occasionally freckled or black; hairs on back silky, lax, and long

(majority of hairs on midline of back 75-150 mm. in length) ; tail

often more than twice length of head and body. (North bank of

Amazon east of Manfios and north to Caribbean coast of Guianas. )

paniscus
EE. Tail much less bushy but more densely furred in adult (longest
hairs on upi>erside 4(M');j mm. in length); face wholly black;
hairs on back .shorter (majority of hairs on midline of back
75 nrm. or less) ; tail often more than twice length of head and
bod J'. (Western Mat to Gros.so, eastern Bolivia, and north-
eastern Peru to Rio Solimoes and Rio Jurua, Brazil.) chamek

DD. Foot smaller (length, heel to end longest toe, 100-170 mm.) ; tail not
noticeably bu.shy (the longest hairs on upperside 30-50 mm. in
length) ; face wholly black, except for short, white nuistache
whiskers on upper lip and sparse, short, white chin whiskers;
hairs on back harsh and of medium length (majority of hairs on
midline of back 40-70 mm. in length) ; tail variable in length,
occasionally nearly twice as long as head and body. (Serranfa del

Darien, Panama, to .southwestern Colombia.) robustus

CC. Hairs on head not wholly black; semilunar forehead patch and side
whi.skers on face white; remainder of body, lind)S, and tail deep
glo.ssy black; foot nre<lium (length, heel to end longest toe, 170-185
mm.) ; tail vari;d»le in length, occasionally more than twice as long
as head and body. (Slate of Parfi. Brazil, .south of Amazon, between

Rios TajiajTiz and Tocatitlns. ) marginatus

JiB. liack and belly, as well as limbs and tall, not wholly deep glossy black;
hairs on back black at hnsi', and either black dislaliy or with lii>s
tinged with either burnt umlter or seal brown.
F. Crown cap tawny-oUve, yellowish wood brown, oi- willi black hairs lipped
with burnt umber; tuiderparts similar in color to \ipjH'riiarts; entire
body, Including limb.s and tail, covered with black hairs tinged dislaliy
with burnt umber; foot small (leiiglli. heel to end longest toe, l.'.O-170
mm.) ; tail variable in length, usually one-flfth or more longer lh:in
head and body. (Pacific side of cordlllera of Ecuador.) fusciceps

' ('..lors niffitlono*! In thU kpy are nhown In Ilotjort RIdKwny'fl 'Tolor SlandnrdH nnd
Color Nomenclature," 4.3 pp.. r.3 pig. WuHliinKton, 1012.

8 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

FF. Crown cap black, not tawny-olive, yellowish wood brown, or with hairs
tipped with burnt umber ; underparts conspicuously lighter in color
than upperparts, line of demarcation between upperparts and under-
parts being sharply defined ; general coloration of crown cap on head
as well as rest of upperparts black ; in some individuals these black
hairs under strong light having a seal-brown tinge ; triangular fore-
head patch yellow, golden, brownish yellow, or white (occasionally
either entirely absent or more or less hidden by anterior stiff black
hairs) ; side whiskers on face, when present, whitish or huffy (not
present in some specirtfens) ; outer surfaces of forelimbs to elbows
black ; outer surfaces of forearms to wrists generally blackish but
occasionally similar to underparts and intermixed with long dusky
overhairs ; outer surfaces of hind limbs usually blackish to knees or
below, but in some specimens (particularly in eastern part of range)
outer surfaces of hind limbs, including thighs and region around base
of tail, distinctly straw-colored in ground color overlaid with sparse
long black hairs (not numerous enough to affect general ground color) ;
hairs on throat normally black, sometimes grayish, more or less
concealed by black hairs ; remainder of underparts, inner surfaces
of fore and hind limbs, as well as under surface of tail, yellow,
cinnamon-buff, dull cream-buff, or pale olive-buff; tail normally
bicolored, upper surface being black and under surface similar to
that of underparts (in occasional specinfens upper surface of tail
approximately same as that of under surface but more heavily inter-
spersed with long black overhairs) ; foot large (length, heel to end
longest toe, 190-200 mm.) ; tail variable in length but often almost
twice length of head and body. (From near junction of Rio Caura
with Rio Orinoco in Venezuela south to Rio Negro in Brazil, westward
to Colombia east of Cordillera Oriental (Mambita), Ecuador east of

Andes, and northeastern Peru.) belzebuth.

AA. General coloration of entire back not chiefly (jet) black.

O. A conspicuous white triangular forehead patch ; general coloration of i:pper-
parts wood brown, darker on head and upper back and lighter on lumbar
region and thighs ; underparts, inner surfaces of fore and hind limbs, and
under surface of tail whitish or buffy ; side whiskers on face variable,
sometimes whitish or buffy and sometimes dark wood brown. (Magda-

lena River Valley and adjacent mountains, Colombia.) hybridus

GO. No conspicuous white triangular forehead patch ; general coloration of
upperparts not wood brown.
H. General coloration of upperparts, except for head and shoulders on some
individuals, tending toward light mahogany red ; underside of tail, at
least on basal half, deep cinnamon-rufous to ferruginous ; back and
shoulders grading from dark rusty reddish to ferruginous or burnt
sienna, with varying admixture of overlying blackish hairs ; underparts
similar in color to back but with less noticeable admixture of blackish
hairs ; face, crown cap on head, and median streak on back of neck
blackish ; sides of neck covered with a mixture of pinkish-bufE, cin-
namon-buff, and blackish hairs ; outer surfaces of fore and hind limbs
either black or blackish to knees and elbows ; outer surfaces of limbs
below knees and elbows either black or with varying admixture of black
and ferruginous hairs ; a distinct ferruginous or cinnamon-buff streak
along inner surface of arm from armpit to elbow. (Western side of

THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 9

central curdillera of Costa Uiea (lilo I'iiris) south to Cordillera de

San Bias (Cerro Azul) of eastern rauama.) panamensis

////. Cieneral coloration of upporparts, except for head and shoulders, not uni-
form light mahogany red ; underside of tail not deep ferruginous.
/. GeruMal coloration of uppt'riiarls as well as underparts light hufC over-
laid with dusky tipiK-d hairs; side whiskers on face silky light buff;
dark markings on head and limbs usually inconspicuous; crown cap
on head grading from black to buff tinged with brownish ; stiff long
black hairs above eyes concealing to a variable extent the white or
buff triangular forehead patch ; elbows and outer sides of forearms
more or less distinctly blackish ; elongate black patches on knees, the
hairs black to roots. (Southern Nicaragua, Greytowu to Lake Mana-
gua.) I . geoffroyi

//. General coloration of upperparts as well as underparts not light buff.
J. Chest and belly normally near tawny or cinnamon-rufous.

K. General coloration of upperparts golden yellowish to cinnamon-
rufous in tone but obscured by numerous overlying black-tipped
hairs; crown cap black, the hairs black to base; underparts and
flanks near tawny or cinnamon-rufous ; tail black above, more or
less mixed with tawny below from base to near callosity; side
whiskers on face blackish. (Eastern side of central cordillora

of Costa Rica. ) ornatus

KK. General coloration of upperparts lighter or duller than preceding.
L. General coloration of upperparts, except for head and shoulders,
clear tawny, intermixed with black-tipped hairs ; sides, thighs,
and belly paler, tawny or cinnamon-buff, with somewhat paler
tone extending downward to wrists and ankles on inner sur-
faces of fore and hind limbs; shoulders and outer surfaces of
fore and hind limbs black; tail .sharply bicolored, black above
to tip, and tawny below; crown cap either black or dusky (in-
dividual hairs buff or cinnamon-buff at base) ; side whiskers on
face cream color. (Azuero Peninsula, Panama. )_-azuerensis
LL. General coloration of upperparts, except for head and shoulders
in some individuals, buckthorn brown to Mars brown, sparsely
or Jioticeably intermixed with blackish hairs (especially on
midline of back in sonit? individuals) ; belly honey yellow to
tawny, «'Xten(liiig downward on inner surfaces r>f hind limbs to
ankles, and a lighter tone extending downward on inner sur-
faces of forellmbs to wrists; outer surfaces of fore and hind
limbs usually black (in some individuals restricted more or
le8.s to elbow and knee patclies) ; tail usimlly sharply bicolored,
bhnk or diisky above to tip and tawny below (when otherwise,
mixed light and dark hairs cover tall above and below) ; an-
terior part of crown jiatch more or less suffused with cliuia-
nion buff, owing to liglil basal color of liairs and tending to
form a transverse band across forehead ; side whiskers on face
light huff to cream-huff. (Northwestern Costa Rica and higlier

p<irtioiis of northern Nicaragua.) frontatus

JJ. Chest and belly normally neither tawny nor orhrnceous-rufous.

J/. Upper back distinctly darker than lumbar region of back; top of
head. nJTk and upper back, and outer surfaces of fore and liind
llmb.s varying from blackish to brownish black.

002420—44 -2

10 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

N. Underi>arts normally silvery whitish ; brownish black of head and
neck passing gradually through cinnamon-drab on upper back
to near olive-buff on lumbar region, the light drab extending
along median line to base of tail ; side whiskers on face whit-
ish. (Quintana Roo, Yucatan, and northeastern Guate-
mala.) yucatanensis

NN. Underparts not normally silvery whitish.

O. Upperparts dark, especially on head, neck, and upper back, and
more uniformly overlaid with black ; lumbar region more or
less noticeably suffused with cinnamon ; pelage typically
longer and denser ; underside of neck and throat mixed brown-
ish and buffy ; rest of underparts including at least inner sur-
faces of upper portions of limbs pinkish buff to cinnamon-buff,
becoming cinnamon or rusty along sides of body ; crown cap
blackish ; side whiskers on face blackish. (Alta Vera Paz,

Guatemala.) pan

00. Upperparts lighter, distinctly bicolored ; head, neck, and
upper back brownish black ; lumbar region cinnamon-buff
or cinnamon, darkened along midline by overlying blackish
hairs ; underside of neck and throat varying from grayish
to a mixture of buff-gray and brown ; underparts, includ-
ing inner surfaces of arms in a strip narrowing from
armpit to a point near elbows and on inner surfaces of
legs to near ankles, varying from pinkish buff to cin-
namon, becoming near cinnamon on sides of body ; crown
cap blackish ; side whiskers on face dull whitish or yellow-
ish. (Honduras and El Salvador northward through
Guatemala to southeastern Oaxaca and the Isthmus of
Tehuantepec and along east coast of Mexico to eastern

San Luis Potosl.) vellerosus

MM. Upper back not distinctly darker than lumbar region of back ;
general coloration of upperparts, including outer surfaces of
fore and hind limbs, as well as upper surface of tail, dusky,
the darker hairs nearly sooty black on distal two-thirds and
old gold on basal third, and thinly interspersed with these
on head, neck, back, thighs, and upper surface of tail are
old gold or silvery hairs ; coloration of upperparts further
modified by lighter hair bases showing through ; a partially
concealed huffy spot on forehead ; hairs on belly, inner sur-
faces of hind limbs, and under surface of tail dull cinnamon-
buff tipped with sooty black. (Rio Tuyra Valley of eastern
Panama.) grisescens

THE SPIDER MONKEYS KELLOtIG AND tlOLDMAX 11

ATELES PANISCUS PANISCUS (Linnacug)

Guiana Black Spider Monkey; Le Coaita

[Siniin] pdiii.srus L^^•NA^n^s, Systoiiia natural, cd. 10, vol. 1, p. 2(5. 1758.

SdiJitjoH pani'^cufi L.vcia'KDt; Tableau dcs divisicms, sous-divisions, ordres et genres
lies inaiimiifOres. p. 4, linit. [Published as supplement to Di.scours d'ouver-
turo et de cloture du cours d'histoire naturelle, et tableaux nu'thodique des
inamniifdres et des oiseaux. ]

Ateles pentadactyltis E. Gewfbroy, Ann. Mus. Hist. Nat. Paris, vol. 7, p. 2G9, 180G.
Type locality, Cayenne, P^rcnch Guiana. (Description based on female col-
lected by Martin.)

Atclea nubpcnfadarti/Iuii Desmakest, Encyclopedie methodique (Zoologie), Mam-
nialogie, pt. 1, p. 73. 1^20. Typo locality, '"La Cuyane fran(;ai.se, la cote de
liancet au P(>rou" (part).

A[t€lc8] utrr F. Cuvieb, Ilistoire naturelle des mammif^^res, vol. 3. livr. X). p. [1 I,
pi. — , Mar. 1823. Type k>cality, Cayenne, French Guiana (see I. Geofifroy,
1851, Catalogue methodique de la collection des mammif«^res, pt. 1 (Catalogue
des Primates), p. 48). ("Le Cayou" is based on a rather young female.)

[Cebus paniscus] siirituiDirnsis Fiscin^. Synopsis niammalium, p. 39, 182f). Type
locality, not designated.

[Cebus panismi] ftnioincnsis Fischer, Synopsis mammaliuni, p. 30, 1820. Type
locality, "Guyana Gallica."

Ti/pe locality. — "South America : Brazil" ["La Guyane" given as the
"Patrie" by E. Geoflfroy, Catalo^ruo Mammi feres Mus. Nat. Hist. Nat.
Paris, p. G, 18U3. Here restricted to French Guiana.]

Type specimen. — Unknown.

Dt.strihution. — North side of Amazon River east of Manaos and
north to Caribbean coast of the Guianas.

General characters. — A black monkey Avilli face normally flesh-
cr)hjred ; pehi^e silky, lax, and very long (majority of hairs on mid-
line of back 75-150 mm. in length, and on upper basal half of tail
50-100 mm. in lenglh) ; hind foot large (190-220 mm.); tail often
more than t\vice length of head and body. Differs from chamek of
Peru mainly in normally flesh-colored instead of normally black face
and in longer pelage (majority of hairs on midline of back less than
75 mm. in length and on iipjwr base of tail less than 50 mm. in length
in cham/'h). Differs from A. hehehuth hclzelntth of Venezuela in
much longer pelage and nearly uniform black instead of contrasting
colors of upi)er and un(lerf)arts. Differs from A. hehehuth maryinafus
')f the Kio Tapajoz region of the southern side of the Kio Amazonas,
Hrazil, in longer pelage, larger foot, and lack of the semilunar white
forehead patch present in maryinatim.

( ''ilor.--VjU\ iic pelage deep ghtssy black ; face noiinall y (lesh-coloied.

t</.ull. — ^kull large, with anteiior profile strongly inclined backward
to vault of brain ca.se; premaxillae, when viewed from above, rela-
tively brf)ad ; nasal opening much elongated. Very similar to skulls
of chamek and helzebrith^ but brain case somewhat less highly arched
than either; rostrum bioailer. the sides more inflalid at base and

PROCEEDINGS OF THE NATIONAL MUSEUM

6
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THE SPIDER MONKEYS KELU)GG AND GOLDMAN 13

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PROCEEDINGS OF THE NATIONAL MUSEUM

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THE SPIDER MONKEYS KELLOGG AND GOLDMAN 15

sloping' uiilward more ^n-;ulually to zy<i()inata; maxillao fonninj; a
more distinct ridpe behind posterior molars; thin oxtoinal redupli-
cations of pteryijfoids cxtciidiii^ farther i^osteriorly. the posterior bor-
ders less deeply concave behind tiie internal spinelike hanuilar proc-
esses; posterior plane of posterior upper molars barely reaching ante-
rior plane of temporal fossae.

Mcasure77wnt,'<. — Two adult females from Rio Janiunda, near Faro,
north bank of Rio Amazonas, Para, Brazil, respectively : Total length,
i;330. 1.389 nmi. ; tail, 8T0. 920; hind foot, 195, 195. Two adult females
from Lago Cuipeua, north bank of Rio Amazonas, Para, Brazil, respec-
tively: Total length, 1,450, 1,413; tail, 880, 753; hind foot, 220, 220.
Skull: Two adult females from Rio Jamundti, near Faro, Brazil, respec-
tively: Greatest length, 118.8, 11G.8; orbital width, G8.3, G3.C; post-
orbital constriction, 48.8, 51.3; width of brain case, 64, 63.3; zygomatic
width. C8.7. 66.7; maxillary tooth row, 29.9, 29.5. Two adult females
from Lago Cuipeua, Brazil, respectively: Greatest length, 113.6, 122.6;
(jrbital width, 60.2, 68.2; postorbital constriction, 51.4, 51.4; width of
brain case, 64.5, 64.6; zygomatic width, 69.4, 70.5; maxillary tooth row,
29.9, 29.5.

Remorks. — Linnaeus, in tlie tenth edition of Systema Naturae (vol.
1, J). 26, 1758) combined the previously published descriptions of a
spider monkey and of a howler monkey in formulating his diagnosis of
ySimia^ paniscus. The spider monkey is described in the following
words: "Simla fusca major, palmis tetradactylis, cauda prehensili ad
apicem subtus nuda," and this is a word-for-word quotation from the
account of the "four-fingered monkey" published by Browne (The civil
and natural history of Jamaica, p. 489, 1789; edition of 1756 not con-
sulted). The longer diagnosis at the end of Linnaeus's account of
prini.scus seems to be based mainly on the accounts of the "Guariba"
publi.shed by Marcgrave (Historiae rerum naturalium, p. 226, 1648;
and Ilistoria natuial do Brasil, p. 226, 1942) and by Ray (Synopsis
melhodica animalium quadrupi'«hnn et serpentini generis, p. 153,
1693), .supplemented by additional anatomical details which are
credited to Hallman and Aymen.

In thf twelfth edition, however, Linnaeus (Systema naturae, ed. 12,
vf)l. 1. J). 37, 1766) seems to have recognized that (he above-mentioned
combination was inaccurate, and of the tenth edition references he
cited only that of Browne (op. rit., 1756, |). 189) in the synonymy of
[Shnia] paiihcuJi. With the exception of "Pedes Sc Cau(hi dimidia
exterior bnninea,'' this revised diagnosis ai)|)lied to the "coaita." Fur-
thermore, in the twelfth edition Linnaeus {i>p. r/'f., p. 37, 1766) i)laced
the references to Marcgrave and Kay. which had l)een cited under
paniy(ru.t in the tenth edition, in the syii'inymy of the howler monkey,
[,S/////V/. I htlzehul. We are therefore of the o[)inion that Limiaeus in

16 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

1766 should be regarded as the first reviser and that the references to
Marcgrave and Ray should definitely be eliminated in reaching a con-
clusion concerning the status of paniscus.

The "Guariba" of both Marcgrave and Ray is unquestionably the
howler monkey, as was recognized by both Pennant (Synopsis of
quadrupeds, p. 122, 1771; "preacher") and Goldfuss (Schreber, Die
Sfiugthiere, Theil 1, Heft 7, p. 112, 1774) . The bases and type localities
of the mammalian species in the tenth edition of Linnaeus were con-
strued as they were by Thomas (Proc. Zool. Soc. London for 1911,
pt. 1, p. 127, Mar. 22, 1911) on the assumption that Linnaeus's "quota-
tion of his own earlier writings should be given absolutely overriding
importance" and that all others should be ignored. The Linnean
quotation under [^Simia] j^cmiscus in the tenth edition is "S. caudata
barbata, cauda prehensili, palmis subtetradactylis. Syst. nat. 3," and
on referring to the sixth edition of Systema Naturae (p. 3, 1748) , it will
be noted that Linnaeus altered his original concept of this species by
adding "palmis subtetradactylis," since the diagnosis under "/S'^m^(^"
in the earlier edition reads "14. Simia caudata barbata, cauda pre-
hensili. Marcgr. bras. 226." On the assumption that the reference to
Marcgrave was the primary reference, Thomas designated Pernambuco
as the type locality for paiiiscus. In view of the fact that Marcgrave's
account of the "Guariba" applies solely to the howler monkey, the
designation of Pernambuco as the type locality for paniscus cannot
be accepted. Browne {op. cit., p. 489, 1789) stated that the "four-
fingered monkey" is an inhabitant of the main continent, Linnaeus in
the tenth edition gave the habitat of paniscus as "America meridionali :
Brasilia," and in the twelfth edition after the removal of the references
to the "Guariba" Linnaeus restricted the habitat to "America meri-
dionali." Since "La Guyane" has been designated by Geoffroy {op.
cit., p. 6, 1803) as the "patrie" of paniscus, we hereby restrict the type
locality to French Guiana.

The Guiana black spider monkey is readily distinguished from all
others by its long, lax, silky pelage and its large foot. It is believed
to be restricted to the area between the north bank of the Amazon River
and the coast of the Guianas. It was met with by various early trav-
elers, and several names proposed seem clearly to belong in synonymy.
In the three specimens from Rio Jamunda, on the north bank of the
Amazon, near Faro, Brazil, vestigial thumbs are present as pointed
out by Tate (Bull. Amer. Mus. Nat. Hist, vol. 76, p. 215, Oct. 20,
1939). In two from Lago Cuipeiia, also on the north bank of the
Amazon, farther to the east, vestigial thumbs are present in the
skeleton, as usual in the group, but do not appear to have been dis-
cernible in the skin. Tate {op. cit.) refers to "small, triangular
pterygoids without pointed tips" as a cranial character in paniscus.
The external reduplications of the pterygoids are broken off in the

THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 17

specimens examined by Tute, but the thin pterygoid plates in these
and in other specimens from that region are in reality larger and
more extended posteriorly than usual in spider monkeys. Other char-
acter ascribed by him to paniacus are the small postglenoid processes
and unexpanded zygomata of nearly uniform depth. Examination
of large series of spider monkey skulls shows that these cranial fea-
tures are everywhere subject to about the same wide range of individual
variation.

In DiUch (luiana, Kappler (Popular Sci. Monthly, vol. 32, No. 3,
pp. 397-31*8, Jan. 1888) states that this monkey does not occur on the
coast and that it is found only in the higher lands. An adult male
collected by Kappler along the Marowijne lliver [ = Rio Maroni],
Dutch (iuiana, is listed by Jentink (Catalogue systematique des
maiiiiniferes, Mus. Hist. Nat. Pays-Bas, vol. 11, p. 41, 1892). Thomas
(Ann. Mag. Nat. Hist., ser. 8, vol. 6, p. 505, Nov. 1910), however,
records this monkey from the River Supinaam, lower Essequibo,
Demerara. British Guiana. Richard Schomburgk (in Roth, Richard
Schomburgk's travels in British Guiana 1840-1844, vol. 2, p. 72, 1923),
states that this monkey is found mostly in troups of 16 to 20 in the
higlust trees and that one was killed in the vicinity of Maripa at the
forks of the upper Rio Waku-wau, a tributary of the Rio Tacutii,
south of the Kaniiku Mountains, British Guiana.

In French Guiana, Menegaux (Bull. Mus. Hist. Nat. Paris, vol. 8,
No. 5, p. 21JG, 1902) has recordetl this si)ider monkey from the Riviere
Camopi, a tributary of the Rio Oyapock, and from the Rio Lunier,
a tributary of the Rio Carsevenne [=Calsoene].

Lonnberg (Arkiv for Zool., vol. 32A, No. 25, p. 8, july 18, 1940)
has recorded specimens from the Igarape [==Rio| Anibii, north bank
of Rio Ama/.onas opj)()sile mouth of Rio Madeira, and from Patua
(or Paitana;, Pani, Brazil.

apecimen^ examined. — Total number, 5, as follows: Buazil: Lago
Cuipei'ia, noi-th bank Riu Amazonas, I'aia, 2 (M. C. Z.) ; Rio jMniuii(l;i
near Faio. iioi I h bank of Ri(» Amazonas, Para, 3 (A. M. \. H.).

ATKLKS PANISCL'S ( HAMKK (Humboldt)
Bl^CK-FACED Bl^CK Sl'IDKR MO.NKKV

isimiii chnnii.h lli MiiM DT, Hfciu-il (r<il)s«'rv!il ions tie '/uolnf^ic ct d'aiintoiiiic

cuiiipur^f, vol. 1. J). .'{.'),'{, ISlli.
AtclcH UiiiiiivuinhriH .J. A. Ai i KN. F?iill. Aiiht. Miih. Nut. Hist , vnl. WW, \\. dril,

Yh'i'. 14, 1!M4. Type Inrjilily. Hiirirn. dc M.lKin.o, licMlwattTs of Uio Cy-

raraii.'i, .Matto (Irosso, I?ruzil.
[AtelcH atrr\ pcruvianux Lii.NNBFJW, Arkiv Mr Zool.. vol. .'{2A. No. 2.">, p. 1."., .Inly

18, 1U40. TjjM? locality, ca.stfrti Peru.

Ti/pe localitij. — Peru. [Here re.stiicitMJ to Uio Coiiilirniato, a
tributary of Rfo Urubamba. Cnzco. Pciii.l

002420 44 3

18 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Type specimen. — Unknown.

Distribution. — Western Matto Grosso, eastern Bolivia, and north-
eastern Peru to the Rio Solimoes and Rio Jurua, Amazonas, Brazil.

General characters. — Entirely black, with pelage of moderate
length, the hairs on midline of back 75 mm. or less, and on upperside
of tail 40-65 mm. Similar in general to paniscus of French Guiana,
but face black instead of flesh color; pelage shorter, the majority of
hairs on midline of back 75 nxm. or less, instead of 75 to 150 ; tail often
more than twice length of head and body and hind foot large as in
paniscus; skull differs from that of paniscus in detail. Similar in
color to A. fusciceps robustus of southeastern Colombia, but pelage
entirely black without the short, sparse white hairs on chin and about
the mouth of the latter that while inconspicuous appear to be impor-
tant as a distinguishing character ; cranial details also different. Dif-
fers from A. belzebuth beJzebuth of Venezuela in uniform black in-
stead of contrasting black and buffy upper and underparts.

Color. — Epidermis of face normally black and entire pelage deep,
glossy black.

Skull. — Similar in size to that of paniscus^ but brain case slightly
more highly arched ; rostrum narrower, less inflated at base and slop-
ing outward more abruptly to zygomata ; premaxillae more distinctly
pinched-in behind incisors, the sides being almost parallel; maxillae
more truncate instead of prolonged in a distinct ridge behind posterior
molars; external reduplications of pterygoids less extended poste-
riorly, the posterior borders more deeply concave; palate narrower;
posterior plane of last molars reaching farther posteriorly into ante-
rior plane of temporal fossae. Differs from that of robustus as fol-
low^s : Premaxillae more extended anteriorly beyond canines, the sides
more constricted and more nearly parallel at diastema between in-
cisors and canines; nasal opening more elongated. Compared with
that of belzebuth the skull is very similar in general, but the premaxil-
lae differ in narrowness in about the same way as from robustus.

Measurements. — An adult male and female from Rio Comberciato,
Peru, respectively : Total length, 1,270, 1,380 mm.; tail, 820, 880; hind
foot, 190, 202. Two adult females from Rio Yapacani, Bolivia, respec-
tively : Total length, 1,320, 1,320 ; tail, 800, 800 ; hind foot, 220, 220.
Skull: Adult male and female from Rio Comberciato, respectively:
Greatest length, 117.5, 117; orbital width, 48.9, 50.8; width of brain
case, 64.7, 59.8 ; zygomatic breadth, 68, 67.8 ; maxillary tooth row, 33,
29. Two adult females from Rio Yapacani, respectively : Greatest
length, 121.1, 118.8; orbital width, 63.8, 59.2; postorbital constriction,
50.1, 48.7; width of brain case, 65.4, 64.7; zygomatic breadth, 71.3,
68.2; maxillary tooth row, 31.4, 31.5.

Remarks. — The large black spider monkey here treated as A. panis-
cus chamek appears to be distinguished as the most completely black

THE SPIDER MONKEYS KELLOGG AM) (JOLDMAX 19

form ill tlie group. It afrroes closely in larfijc size with paniscus but
dilFers in having black instead of flesh-colored face, shorter pelage, and
cranial details pointed out. The general resemblance of chamek to
A. fu^ciceps rohuyfiis of Colombia is very close, but the short white
hairs on the chin and about the mouth, together with the smaller foot
of the latter, are distinctive.

Specimens collected by Johann Xalterer in 1825 on the Rio Sarare
while en route from Matto Grosso (Villa Bella de Santissima Trini-
dade) to Sao Vincente and in 1829 on the Rio Madeira above the mouth
of the Rio Abuna probably are referable to this race (Wagner, Abh.
math.-phys. CI. bayer. Akad. Wiss. Miinchen, vol. 5, Abt. 2, p. 418,
1848). Similarly the specimens described by Miranda Ribeiro (Com-
mi'^sao de Linhas Telegraphicas Estrategicas de Matto Grosso ao
Aniazonas, Annexo No. 5, Hist. Nat., Zool., Manmiiferos, p. 8, May
1914) from Cabeceiras do Pirocoluina and Jani on Rio Jarua, a tribu-
tar}' of the G)--Parana, are likewise referred to this race. This spider
monkey is rejiorted to occur also in the Serra dos Parecis in central
Matto Grosso. The specimens mentioned by Lonnberg (Arkiv for
Zool., vol. 32 A, No. 25, p. 10. July 18, 1940) from Puerto Salinas and
Desierto on the Rio Beni, Bolivia, may belong here since three speci-
mens (C. M. Nos. 2772, 2774, 2775) taken by Jose Steinbach on the Rio
Yapacani, a tributary of the Rio Mamore, seem referable to this form.

Under the name of Atcles ater \^=paniscus']^ Thomas (Ann. Mag.
Nat. Hi.st., ser. 10, vol. 2, p. 251, Sept. 1928) listed specimens from
Cerro Azul, altitude 2,000 feet about 35 miles west of Rio Ucayali, and
from Chicosa, upper Rio Ucayali, Peru, about 35 miles below junc-
lioii of I'rugamba and Tambo Rivers, altitude 1,500 feet. A similar
allocation has been made i)y Thomas (Proc. U. S. Nat. Mus., vol. 58,
p. 2:^:0, Nov. 10, 1920) of specimens (U. S. N. M. Nos. 194337-38 and
-40) from llio Comberciato, a tributaiy of Rio Urubamba, Cuzco,
Peru, altitude 3.O00 feet, and from Paciiitea.

AccfU'ding to liartlett (Proc. Zool. Soc. L<jndon for 1871, pt. 1, p.
218, June 1871) the black-faced spider monkey inhabits the forests on
the Rio Ucayali and tlie low districts of the valley of the Ama/onas
ill I*eru. A specimen from Peruate on Rio Maranon is recorded by
Cabrera (Trabajos Mus. Nat. Cienc. Nat. IMadrid, ser. zool., No. 31,
j». 1<'>. Oct. 28, 1!»17). A female obtained by Oastelnau and Devilje on
tiie banks of the Jii(; Javari near its mouth on the Rio Amazonas is
listed by I. Geolfroy (Catalogue methodique de la collection des mam-
miferes, pt. 1 (('atalogue (h's I'rimales), [). 18, IS51 ) .

It is possible that this bhick-faccd spider monkey may range down
the Rio Maranon and the Solimoes to the Rio .Jurua in I>ia/,il since
I^iinnlxMg (Arkiv f<ir Zool., vol. 32A, No. 25, \\ ]0, July IS, |!M0) lihts
as ater .specimens from Lago (Jrande west of mouth of Rio Jurua on
Rio Solimoes, from Jaburu and Joao Pessoa on the lower Hio Jurua

20 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

and from Santo Antonio on the Kio Eiru, a tributary of the upper Kio
Jurua. The specimens described by Von Ihering (Rev. Mus. Paulista,
vol. 6, p. 409, 1904) from below Sao Felipe on the Rio Jurua should
belong to the same race.

Although Wallace (Proc. Zool. Soc. London for 1852, pt. 20, p. 108,
May 23, 1854) remarks that another spider monkey, "probably Ateles
oier^ inhabits the West Brazil district on the river Purus," recent work
in Brazil has not confirmed this statement.

Specimens examined. — Total number, 9, as follows : Bolivia : Ibon
[ = Ivon], Rio Beni, 1; Rio Yapacani, 4 (1 skull only; C. M.) ; Santa
Helena (upstream from Rurrenagaque), Rio Beni, La Paz, 1. Peru:
Rio Comberciato, a tributary of Rio Urubamba, Cuzco, 3.

ATELES BELZEBUTH BELZEBUTH E. Geoffrey

Marimonda Spider Monkey

Ateles helseluth E. Geoffroy, Ann. Mus. Hist. Nat., Paris, voL 7, p. 272, pi. 16,
1806.

Le chuva de Bracamorros Humboldt, Recueil d'observatious de zoologie et
d'anatomie comparee, vol. 1, p. 8, 1812; "Ateles nuirginatiis," p. 341 (Prov-
ince de Ja4n de Bracamoros, on the banks of Rio Santiago and Rio Ama-
zonas [=Rio Maranon] betvpeen the cataracts of Yariquisa and of Pato-
rumi, Departamento de Amazonas, northern Peru; specimen brought by
Indians from Tutumbero, opposite Pongo de Cacangares, seen in house of
governor at Tomependa, on Rio Maranon near mouth of Rio Chinchipe.)

A[teles^ marimonda Oken, Lehrbuch der Naturgeschichte, Theil 3, Zool., Abt. 2,
p. 1201, 1816. Type locality, Orinoco.

Ateles fuligiiiosus Kuhl, Beitriige znr Zoologie uud vergleichenden Anatomie,
Abth. 1, p. 25, 1820. Type locality, unknown. [Based on specimen in
Mus. Nat. Hist. Nat. Paris.]

Cebus hrissonii Fischer, Synopsis mammalium, p. 40, 1829. Type locality, RIa
Orinoco, Venezuela.

A\_teles'\ variegaius Wagner, Schreber's Die Siiugthiere, Suppl., Abt. 1, p. 313,
1840. Type locality, Cocuy [=Cucui or Cucuhy], Rio Negro, northern
Amazonas, Brazil, at the Venezuelan boundary (^dc Wagner, 1847, Abh,
math.-phys. CI. bayer. Akad. Wiss. Miinchen, vol. 5, Abth. 1, p. 421) =
Serra de Cocoi, upper Rio Negro {pie Sclater, Ann. Mag. Nat. Hist., ser. 4,
vol. 6, p. 472, Dec. 1870). (Preoccupied by Simia variegatus Kerr, 1792=
Ateles variegatus (Kerr), unidentifiable.)

Ateles bartlettii Gray. Ann. Mag. Nat. Hist., ser. 3, vol. 20, No. 118, p. 300,
Oct. 1867. Type locality, Brazil, the upper part of the Amazons ["Eastern
Perti, near Xeberos"=Jeberos or Jeveros, Loreto (see Gray, Proc. ZooL
Soc. London for 1867, pt. 3, No. 63, p. 992, pi. 47, Apr. 1868) =Forests of
northwestern Perfi at highest point of mountains on forest trail between
Moyobamba and Chayavitos, three days' journey from the latter (see
Bartlett, Proc. Zool. Soc. London for 1871, pt. 1, No. 14, p. 217, June 1871).]

Ateles chuva Schlegel, Mus. Hist. Nat. Pays-Bas, vol. 7, livr. 12 (Monogr. 40:
Simiae), p. 175, 1876. Type locality, not designated. [Based in part on.
"Le chuva de Bracamorros" of Humboldt, 1812 ; Ateles variegatus of Wag-
ner, 1840; Ateles bartlettii of Gray, 1868; and two mounted female speci-
mens, one from Peru, in the Leiden Museum.]

THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 21

Atelcs prnblema Schtjcoel MS., in Jontink, Catalogue systi'matique dos mam-
miferes, Mus. Hist. Nat. Pays-Bas, Leiden, vol. 11, p. I-, ]8;»2 {nomcn vudiiin).

Type loeaT/fij. — I'nknown. [Iloro restricted to E>ineralda. west of
(he mouth of Kio Giinpo, on Rio Orinoco, and south of JNIount Duida,
Venezuela (see Humboldt, Recueil d'observations de zoologie et
d'anatomie comparee, vol. 1, livr. 7, p. 3'2G, 1812).]

Type .ypec'unvn. — A .specimen exhibited at public fairs until alter
its death, when it passed into the possession of Reaumer, constituted
the basis of "Le Belzebut" of Brisson (Rcfjnum animale, p. 211, 1756) ;
this .stulFed specimen, the description of Builon, and two livin<i: spider
monkeys in the menagerie were combined as the basis for Geotfroy's
name.

Distribution. — Ranges in the vast lowlands from near the junction
of the Rio Orinoco and Rio Caura in central Venezuela* south to the
valley of the Rio Xegro, westward to Colombia east of the Cordillera
Oriental (Mambita), Ecuador east of the crest of the Andes, and to
northeasteni Peru (Sarayacu).

General characters. — Distinguished by unusual color pattern, the
black upperparts contrasting strongly with i)ale buify undorparts,
there being a sharp line of demarcation along lower part of sides.
This marked contrast in coloration extends to the outer and inner sur-
faces of its liml)s, although these vary in detail, and a triangular fore-
head patch varying from white to golden or brownish yellow is nor-
mally but not invariably present. Foot large (190-200 mm.) ; tail
variable in length but often almost twice the length of head and body.
Dilfers conspicuously in color, but similarity in cranial details indi-
cates close relationship to hyhridus of Colombia. Differs notably from
A. panlsru.<i poftiscu.s of French Guiana in much slioi-tcr polago and in
contrasting colors of upper parts and underparts.

Color. — Face and eyebrows black, a triangular forehead patch vary-
ing from white to golden, yellow, or brownish yellow (occasionally
entirely ab.'^ent or more or less hidden by stiff uj)t iirned black hairs over
eyes and opposing black frontal tuft from crown) ; crown cap and
upperparts generally black to a sharp line of demarcation with under-
j)arts (in some individuals the black hairs under strong light have a
.seal-brown tinge) ; underparts, in general, including inner surfaces of
fore anfl hind limi)s, and underside of tail varying from cinnamon-bufT,
yellow, or dull cream-buff to pale olive-buff; side whiskers on face,
when present, whitish or buffy (absent in some specimens) ; ou(<'i-
surfaces of foi-elimbs to elltows black; outer surfaces of forearms to
wri.sts generally bbicjcish, liut occasioiuilly similar to undeiparts and
intermixed with long dusky overhairs. Outer surface,s of liind limbs
usually blackisli to knees or 1)elow, but in some specimens (j)articularly
in eastern part of range) the outer suifaces of jiind limbs, as well as
thighs and region arf)und base of tail, are distinctly st law-colored in

22 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

ground color thinlj^ overlaid with long black hairs (not numerous
enough to materially affect the general ground color) ; hairs on throat
normally black, sometimes .grayish, more or less concealed by black
hairs ; tail normally bicolored, the upper surface black and the under
surface similar to that of underparts (in occasional specimens the
upper surface is about the same as that of the under surface, but more
heavily interspersed with long black overhairs).

Skull. — About as in that of hyhridus. Very similar in general to
that of fanhcus but brain case somewhat narrower ; posterior borders
of external reduplications of pterygoids more deeply concave.

Measurements. — ^An adult male and female from El Llagual, Caura
District, Venezuela, respectively : Total length, 1,257, 1,317 mm. ; tail,
823, 810; hind foot, — , 193. An adult male and female from Valle de
los Monos, Mount Duida, Venezuela, respectively: Total length, 1,110,
1,330; tail, 695, 840; hind foot, 185, 195. Skull: An adult male and
female from El Llagual, Venezuela, respectively: Greatest length,
109.5, 117.7; orbital width, 60.2, 61.8; postorbital constriction, 50, 51;
width of brain case, 59.7, 59.8; zygomatic width, 68.3, 66.7; maxillary
tooth row, 29.8, 28.8. An adult male and female from Valle de los
Monos, Venezuela, respectively : Greatest length, 119.4, 113.2 ; orbital
width, 61.6, 61; postorbital constriction, 52, 52.1; width of brain case,
61, 63.2 ; zygomatic width, 71.9, 68.4 ; maxillary tooth row, 29.9, 30.

Remarks. — This well-marked and widely dispersed spider monkey
has evidently been described under various names which appear to be
assignable to the synonymy of hehehuth. Humboldt (Recueil d'obser-
vations de zoologie et d'anatomie comparee, vol. 1, livr. 7, p. 353, 1812)
observed that it was found along the banks of the Rio Orinoco but
always above the Eaudal de Atures and the Raudal de Maipures in
Venezuela. Allen records specimens from La Union on the Rio Caura
(Bull. Amer. Mus. Nat. Hist., vol. 20, No. 29, p. 344, Oct. 8, 1904) as
well as from the Rio Mato, a tributary of the Rio Caura (Bull. Amer.
Mus. Nat. Hist., vol. 28, No. 12, p. 148, May 27, 1910) and from El
Llagual on the Rio Caura, Venezuela (Bull. Amer. Mus. Nat. Hist.,
vol. 30, No. 10, p. 272, Dec. 2, 1911). Johann Natterer in February
1831 obtained a male, three females, and a young of this spider monkey
at Serra de Cocoi on the upper Rio Negro, Brazil, one female of which
was exchanged with the British Museum of Natural History (Sclater,
Proc. Zool. Soc. London for 1871, pp. 39, 225, 1871).

According to Bartlett (Proc. Zool. Soc. London for 1871, p. 217),
this spider monkey is found in Peru in the interior forest on the moun-
tain range between Lamas (north of Rio Mayo, a tributary of Rio Hual-
laga) and Saravacu on Rio Ucayali; on the lower spurs of the moun-
tains between Moyobamba and the Rio Huallaga ; at Cahuapanas on
the headwaters of the Rio Cahaupanas, a tributary of the Rio Ma-
raiion; near the native village of Chamicuros (Parinari District) on

THE SPIDER MONKEYS — KELLOGG AXD GOLDMAN 23

the Rio Cliamicuros [ = Rio Siiniiria], a tiibiitai\v of the Rio Marafion;
and on the Rj'o Ti<rre, a northern trihutarv of the Rio Maranon west
of Nauta. It has been reported also from Puerto Indiana on the Ri'o
Marafion just west of mouth of the Rio Napo by Tate (Bull. Amer.
Mus. Nat. Hist., vol. 7G, p. 21G, Oct. 20. 1939), and from Elvira on
the Rio Maranon above the mouth of the Rio Chanibira by Bartlett
(Proc. Zool. Sec. London for 1882, p. 373),

Sl)eciniens are also listed by Gooldi and Hajjjmann (Bol. Mus. Goeldi,
vol. 4, p. 42, 1904) from Iquitos, on the Rio Marafion above mouth of
the Rio Napo, by Cabrera (Trabajos Mus. Nat. Cienc. Nat. Madrid,
ser. zool.. No. 31, p. 4G, Oct. 28, 1917) from Tarapoto (Nueva Floren-
cia) on the Rio Nape, and by Loiniberg (Arkiv for Zool., vol. 14, No.
4, p. 6, June 7, 1921) from below Baeza on the Rio Quijos, a tributary
of the Rio Coca in eastern Ecuador. Festa (Boll. Mus. Zool. ed Anat.
Conip. Univ. Torino, vol. 18, No, 435, p. 4, Feb. 11, 1903) identifies as
Ateles variegatus several specimens collected by him in Ecuador in
tlie valley of the Rio Santia^jo and at San Jose on the Rio Snni and
remarks that he had found this spider monkey to be abundant in the
forests bordering the Paute, Zamora, and Santiago Rivers.

A specimen of this monkey (U. S. N. M. No. 3332) collected by
William E, Moore in 1857 somewhere along the Rio Napo in eastern
Ecuador has yellowish underparts and a golden brow patch.

Spcehnens examined. — Total number, 9, as follows: Colombia:
Mambita (Llanos) on Rio Guavio, a tributary of Rio Meta, 1
(M. C. Z.); Villavicencio, western Rio Meta, 1. Ecuador: Rio Napo,
1; Jinia (southwest of Sigsig) on headwaters of Rio Pamar, a tribu-
tary of Rio Paute, 1 (skull only, M. C. Z.). Peru : No definite locality,
1 (U. M. M. Z.). Venezuela: El Lhigual [ = Yagual], 2
(A. M, N. H.) ; Valle de los Monos, Mount Duida, 2 (A, M, N. H,),

ATELES BELZEBUTH MARGINATUS E. GcofTroy
WniTE-WniSKERED SPIDER MONKEY

Ateles maryinatuH E. GeomtOY, Ann. Mus. Hist. Niif. Paris, vol. 13, p. 92, pi. 10,
Mar. 1809.

AtelcH front (din Uhnnetit, Proc. Znol. Hoc. London for 1830-31, pt. 1, No. 4, p. 38,
Apr. C, 1H;{1. TyiK' locality, unknown.

Atclcti (ilbifroHH IL Sciiinz, Systcniail.sches Verzcicbni.ss aller bis jctzt bckannleu
Siiugethlen? od»?r Synopsis Mamnialinrn, vol. 1, p. OS, 1841. [This name ap-
pears In the Hynonymy of Atelca margitiatus and apparently Is based on
Coaltn A fnmt blanc, AIcIch maryinatux Kr. (Jnvier, Ilistuiro nalun'll»> dcs
niainniifcMcs, vol. 7, llvr. 01*, piiRcs untintMbi'rcd, Apr. is;{(). )

Type locality. — "Los parties dii I5iesil. iiiro.sees par le Rio-
Janeiro" [=-Para, Brazil, and the borders of the Rio Orinoco, Vene-
zuela; see E. GeofTroy, Ann. Mus. Hist. Nat. Paris, vol. 10, p, 100,
1812. Here restricted to Cametfi, Rio Tocantins, Pani, Brazil. |

Type specimen. — Museum National d'Histoire Natiirelle, Paris.

24 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

Distribution. — South side of Eio Amazonas, between the Rio Tapa-
joz and Rio Tocantins, state of Para, Brazil.

General characters. — A deep, glossy-black spider monkey, with a
white semilunar forehead patch and a few white side whiskers on
face. Similar in general to helzehuth., which it resembles in the pos-
session of a conspicuous forehead patch, but the pelage is longer (long-
est hairs on back about 90 instead of 50 mm.) and it differs notably in
the extension of black over entire body and limbs. Differs from panis-
cus of the opposite side of the Rio Amazonas in smaller size, especially
the smaller foot, shorter pelage, and in the conspicuous white forehead
patch (absent in paniscus) ; cranial details also different.

Color. — An adult male from Caxiricatuba, east bank of Rio Tapajoz,
Para, Brazil : Face flesh-colored, at least around eyes and on nose ;
side whiskers white mixed with black; a narrow strip across brow just
above eyes thinly clothed with erect black hairs behind which is a
white semilunar forehead patch overlapped behind by black hairs from
crown ; rest of upper and underparts deep, glossy black.

Skull. — Very similar to that of hehehuth but interorbital region
more depressed. Compared with that of paniscus the interorbital
region is more depressed and the anterior profile rises more steeply to
vault of brain case; it also differs notably in the lesser posterior exten-
sion of the external reduplications of the pterygoids and the smaller
winglike tips.

Measurements. — An adult male from Caxiricatuba, east bank of
Rio Tapajoz, Para, Brazil: Total length, 1,250 mm.; tail, 750; hind
foot, 108 (188 ?). Three females from Marai, right bank of Rio
Tapajoz as stated by Lonnberg (Arkiv for ZooL, vol. 32A, No. 25, p. 7,
July 18, 1940), respectively : Total length, 1,114, 1,190, 1,245; tail, 770,
613, 740; hind foot, 190, 189, 190. Skull: Adult male from Caxirica-
tuba, Rio Tapajoz, Para, Brazil : Greatest length, 114.1 ; orbital width,
61.3; postorbital constriction, 49.5; width of brain case, 59.6; zygo-
matic breadth, 69.7; maxillary tooth row, 31.9,

Remarks. — A. h. marginatu^s is readily distinguished by the white
semilunar forehead patch combined with black general coloration. It
appears to be most closely allied to helzehuth but differs in color pat-
tern and in longer pelage. In length of pelage it suggests paniscus,
which inhabits the opposite bank of the Rio Amazonas, but is less
extreme, as the longer hairs on the middle of the back in that form
exceed 100 mm. in length. In cranial characters also it agrees more
closely with helzehuth than paniscus.

This black spider monkey with white semilunar patch on forehead
and white side whiskers seems to be most abundant between the Rio
Tapajoz and the Rio Xingii. Specimens in the Museu Goeldi at
Belem were obtained at Altamira on the Rio Xingu and at Santarem at
the mouth of the Rio Tapajoz. Lonnberg (Arkiv for ZooL, vol. 32A,

THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 25

Xo. 2"), p. T. July 18, 1040) iwords this monkey from Marai on the
right bunk of the Rio Tapajoz, and Bates (The naturalist on the
river Amazona, p. 217, IST;")) found it at Aracu on the Rio Cupary, a
tributary of the Rio Tapajoz. It was found by Sieber at Cametii on
the left bank of Rio Tocantins.

Specimens e.vamhird. — Tt)tal number. 3, as follows: Brazil: Caxi-
ricatuba, east bank of Rio Tapajoz, a few miles below Tauary, Pani,
1 (M. C. Z.) ; no definite locality, 2 (1, C. M.).

ATELES BELZEBUTH HYBRIDUS I. Geoffrey

Hybrid Spider Monkey

Atclcfi hi/hridiis I. Gvxhfroy, Mem. Mus. Hist. Nat. I'aris. vol. 17, p. 168. 1829.

Aides albifrons (Jeay. Catalogue of monkeys, lemurs and fruit-eating bats in the
collection of the British Museum, p. 44, 1870. Type locality. South America.

[Atclcn bclscbnth] bnunieus Gr-w, Catalogue of monkeys, lemurs and fruit-
eating bats in the collection of the British Museum, p. 44, 1870. Type locality,
"Brazil."

Amcr-anihropoidrs Joysi Montandon, Comptes Rendus Acad. Sci., I'aris, vol. 1S8,
No. 11, p. S17, Mar. 11, 1929. Type locality, left affluent of upper Kio Tarra,
a tributary of tiie liio Catatumbo, which flows into Lake Maracaibo, Vene-
zuela.

Type locality. — Valley of Rio Mao;dalena, Colombia. Here re-
.<5trictcd to La Gloria on the Rio Magdalena, southern ]\Iagdalena De-
j)artment, Colombia.

Type specimen. — Cotypes, 2 male?5 and 1 female, Museum National
d' Histoire Naturelle, Paris; presented by M. Plee in 182C.

Distribution. — Serrania de Valledupar (Las Marimondas) and
southward along the Rio Cesar to the Rio Magdalena (Puerto
Estrella), and south in the Cordillera Oriental at least to latitude 6°
N. in Santander (Bolivar), Colombia.

General cluiraeters. — Distinguished by wood-brown general colora-
tion of upperparts and a while triangular forehead patch. Apparently
clo-sely allied to helzebuth of Venezuela, but contrasting strongly in
brownish in.stead of black upperjiart.s. Differs from its geographic
neighbor, A. jiiaciceps rohuntuH., of western Colombia, in brownish up-
perparts and white forehead patch instead of nearly all black colora-
tion, larger foot, and cranial details.

Color. — Face and upturned hail's on anterior part of forehead just
above eyes black, partially concealing a white triangular forehead
patch; coloi-ation of ui)perj)arts in general wood brown, darker on
head and upper back, becoming lighter and near avellaneous on lumbar
region and hips; outer surfaces of forelimbs, thighs, and upperside of
tail wood brown; underparts, inner surfaces of fore and hind limbs,
and under surface of tail whiti.sh or buify ; siilc whiskers on face vary-
ing from whiti.sh or buffy to daik wood brown.

Skull. — About as in hrlzrhuth. Similar to that of A. pani^sciis panis-
cus but variable, the premaxillae in some individuals less ])roduced

26 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

anteriorly and the nasal opening less elongated. Compared with that
of A. fusciceps robustus the premaxillae tend to be more produced
anteriorly beyond plane of canines and the nasal opening more elon-
gated.

Measurements. — Three adult males from K,io Guaimaral, Valledu-
par, Magdalena, Colombia, respectively: Total length, 1,248, 1,238,
1,211 mm. ; tail vertebrae, 750, 739, 741 ; hind foot, 177, 178, 172. Three
adult females from same locality, respectively: Total length, 1,330,
1,247, 1267; tail vertebrae, 856, 778, 809; hind foot, 185, 187, 187.
Skull: Three adult males already mentioned, respectively: Greatest
length, 113.9, 114.4, 116.6; orbital width, 60, 65, 63.3; postorbital con-
striction, 50.9, 49.2, 49.4; width of brain case, 65.6, 59.7, 65.3; zygomatic
breadth, 71.4, 728, 74.1 ; maxillary tooth row, 30.3, 30.9, 33.3. Three
adult females already mentioned, respectively : Greatest length, 116.9,
109.9, 110.7; orbital width, 64.8, 59.9, 61.3; postorbital constriction,
50.3, 48.1, 49.2; width of brain case, 61.6, 60.7, 60.8; zygomatic breadth,
70.2, 69, 69 ; maxillary tooth row, 32.2, 32.6, 29.2.

Remarhs. — Ateles liybridus was based on specimens presented to the
Paris Museum by M. Plee in 1826. They were collected in the valley of
the Rio Magdalena, Colombia, but no definite locality was given. It
is now known to be a well-marked form with an extensive range, and
for precision we restrict the type locality to La Gloria, on the Rio
Magdalena in the southern part of Magdalena Department, Colombia,
where the animal appears to be typical. Ateles albifrons Gray was
described from South America without definite locality. According to
Elliot (A review of the Primates, vol. 2, p. 45, June 15, 1913) , the type
in the British Museum is stated on the ticket to be from Medellin,
Colombia. That important town was for many years a headquarters
for dealers in natural-history specimens, many of which labeled as
from there are known to have been collected elsewhere, and it is not
improbable that the type of albifrons may have been taken at some
other locality. At any rate, descriptions of the type agree so well with
hyhridus that the name may be assumed to belong in synonymy under
it.

Another spider monkey, [^Ateles helsehuthl bininneus^ was described
by Gray (Catalogue of the monkeys, lemurs and fruit-eating bats in
the collection of the British Museum, p. 44, 1870) as the fourth color
variety of helzehuth. The original description of this specimen, which
was stated to have come from Brazil, is as follows : "Brown, or brown-
washed grey; cheek, loins, and outside of the thighs whiter; chest,
throat, inside of limbs pale grey ; crown, outside of limbs, and upper
surface of tail darker brown." Philip Hershkovitz has recently exam-
ined the type of hrunneiis at our request and writes that it is identical
with hyhridus. The type has been exhibited as a mounted specimen
and is considerably faded.

In cranial characters hyhridus seems to be identical with helzehuth.,
and the similarity in color pattern strongly indicates that the two are

THE SPIDER MONKEYS — KELLOGG AND GOLDMAX 27

conspecific. On the other hand, the cranial details as ^vell as color
ditFerences point to a departure from robu^tus, a j^eographic neighbor
on the west. This monkey has been reported from as far south as
Bolivar in the Cordillera Oriental of Santander Department.

Montandon (La Nature, Taris, No. 2809, vol. 1, p. 440, May 15, 1929;
Coiiiptes Rendus Acad. Sci. Paris, vol. 188, No. 11, p. 817, March 11,
1929) has described a monkey under the name of Amer-anthvopoldes
luysi, typifying an assumed new family Amer-anthropoidae, from a
left aftlucnt of the u|)per Rio Tarra, a tributary of the Rio Catatumbo,
which flows into Lake Maracaibo, Venezuela. The collector. Dr. Fran-
cois de Loys, observed two individuals and killed the female, and al-
though an effort was made to save the skin and the skull these were
subsequently spoiled by humidity. The description was based on notes
and a photograph (Montandon, Journ. Soc. Amer. Paris, new ser.,
vol. 21, pp. 18:3-19;"). pi. 5, 1929) of the animal. The absurdity of the
conclusions reached by Montundon is pointed out in detail by Cabrera
(Rev. Soc. Argentina"^Cienc. Nat., vol. 10, pp. 204-209, July 12, 1930).
The animal photographed seems to be unquestionably an Afclcs with
a trianguhir white patch on the forehead. Specimens examined by us
from tlie San Calisto district of the upper Rio Tarra proved to be
A teles hclzchuth hi/hndus, to which the name Amer-anthropoules loysi
is here relegated in synonymy.

Specimens examined. — Total number, 17, as follows : Colombia : La
Gloria, Rio Magdalena, 2; Las Marimondas, eastern Andes, Fonseca,
Magdalena, o: Puerto Estrella, Rio Magdalena, Magdalena, 2; Rio
Guaimaral, Valledupar, Magdalena, 6; Rio Tarra, San Calisto, San-
tander, 2.

ATELES FUSCICEPS FUSCICEPS Gray

Brown-headed Spider Monkey

Atclfn fusrircps Gray, rn.c. Zool. Soc. London for 18(i5, pt. 3, No. 47, p. TA'.i, Apr.

1806.
Atcles fuscicepi Sclateu. Proc. Zool. Soc. London for 1872, pt. 2. No. 42, p. GGIi,

pi. 54 (col.). Nov. 187"J. (Specimen collcftod |»y (Jlarcnce Buckley in Trans-

undeun Ecuador.)

Type locality. — South America. [Here restricted to Hacienda Chi-
nipamba,near Pefiaherrcra (west of Ibarra), Tntag Disti-ict, Tnibabura
Province, ncn-thwestern Ecuador; allilnde 1,500 meters.]

Type specimen. — Briti.sh Museum (Natural History) No. .

Di-strihufifni. — Pacific side of cordilleia of TOcuador.

General characters. — A black or bnnvnish-black si)ider monkey, willi
top of head more distinctly brownish than body; foot small (150-170
mm.) ; pelage rather eoapse; tail variable in length, usually one-fifth
or more longer than head and body, clothed witii hair of moderate
length. Closely allied to rohimtus of Colomi)ia but more brownish,
especially on the head. Compared with paniscus, especially as repre-

28 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

sented by A. paniscus chamek of Peru, the brownish head is distinc-
tive: other differences are the short whitish hairs on chin (chin nearly
naked and white hairs absent in chamek) ; shorter, coarser pelage, the
crown patch less projecting over forehead, the shorter-haired tail,
smaller foot, and cranial details.

Color. — Face and upturned hairs on forehead black; crown cap
tawny-olive, yellowish wood brown, or with black hairs tipped with
burnt umber; remainder of body, including limbs and tail, covered
with black hairs tinged with burnt umber.

ShuU. — Skull of medium size and proportions; premaxillae mod-
erately extended anteriorly; rostrum rather short and nasal opening
of medium length ; auditory bullae somewhat flattened. Skull similar
to that of the related form rohustus. Compared with that of panisciis :
Premaxillae less produced anteriorly beyond plane of canines and
anterior nasal opening less elongated.

Measure^nents. — An adult male and female from Hacienda Chini-
pamba, near Penaherrera, Ecuador, respectively : Total length, 1,090,
1,281 mm.; tail, 720, 770; hind foot, 160, li69. An adult female from
Cordillera de Chilluri, Ecuador: Total length, 1,194; tail, 655; hind
foot, 152. Skull: An adult male and female from Hacienda Chini-
pamba, near Penaherrera, Ecuador, respectively : Greatest length,
113.7, 117.2; orbital width, 63.6, 62.5; postorbital constriction, 50.4,
48.9 ; width of brain case, 61.2, 61.7 ; zygomatic breadth, 73, 71.2 ; maxil-
lary tooth row, 32.6, 32.2. An adult female from Cordillera de Chil-
luri, Ecuador: Greatest length, 113.8; orbital width, 58; postorbital
constriction, 47; width of brain case, 59.7; zygomatic breadth, 69.6;
maxillary tooth row, 30.2.

Remarks. — Ateles fusciceps was based on a specimen from an un-
known locality. Another specimen of this well-marked monkey was
collected by Clarence Buckley in Transandean Ecuador, as recorded
by Sclater {op. cif.)^ who also published a colored plate. For pre-
cision in regard to the type locality, however, the name is restricted by
us to the form occurring at Hacienda Chinipamba, near Peiiaherrera
(west of Ibarra), Intag District, Imbabura Province, northwestern
Ecuador, where the altitude is given as 1,500 meters.

Lonnberg (Arkiv for Zool., vol. 14, No. 4, p. 5, June 7, 1921) has
recorded specimens from near Gualea (alt. 4,000 feet) and Santo
Domingo de los Colorados (alt. 2,000 feet) on the upper tributaries of
Rio Esmeraldas, northwestern Ecuador.

Specim^n^ examined. — Totalnumber, 8, as follows: Echjador: Caro-
lina Ibarra, Imbabura Province, 1 (skull only, U. M. M. Z.) ; Cordillera
de Chilluri, 1 (U. M. M. Z.) ; Gualea (near), Pichincha Province, alti-
tude 3,000-4,000 feet, 1 (M. C. Z.) ; Hacienda Chinipamba, near Pena-
herrera, Intag District, Imbabura Province, 2 (U. M. M. Z.) ; Mindo
(below), Pichincha Province, altitude 4,000 feet, 1 (M. C. Z.) ; Par-

THE SPIDER MONKEYS KELLOGC! AND GOLDMAN 29

amba, Imbabura Province, 1 (skull only, U. M. M. Z.) ; no definite
locality, 1 (U.M.M. Z.).

ATELES FUSCICEPS ROBUSTUS J. A. Allen

Colombian Black Spider Monkey

Atclcs robiintu.'! J. A. Ali.i;n, Bull. Aiiier. Mus. Nat. Hist., vol. 33, art. 43, p. 052,

Dee. 14, 1914.
Atclcs dwicnsiii Goi.uman, Proc. Binl. Soe. Washington, vol. 28, j). lUl, Apr. 13,

1915. Type healliy, near head t>l' Kin Llnion, Mount Pure, eastern Panama ;

altitude 5,200 leet.

2ype locality. — Gallera, Department of Cauca, western Andes, Co-
lombia; altitude, 5,0(J0 feet.

Type specimen. — Male adult, skin and skull, American Museum of
Natural History No. 32354, collected July 13, 1911, by Leo E. Miller.

Diatiibution. — Western cordillera of Andes from southwestern Co-
lombia northward on west side of Rio Cauca to eastern Panama

(.Mount Pirre).

General characters. — All black, except for a slight brownish tinge on
forehead of one individual and a few inconspicuous white hairs on
chin and about mouth; hairs on back harsh and of medium length
(majority of hairs on midline 40-70 mm. in length) ; tail variable in
length, occasionally nearly twice as long as head and body ; hind
foot .-nuiil (100-170 mm.) ; tail hairs of moderate length (30-50 mm.
on ui)perside) . CIo.sely allied to fiusciceps of Ecuador but more nearly
uniform black in color, without the distinctly brownish head of
fuHciceps. Similar in color to paniaciLs of northern Brazil, but with
a few white hairs on chin (chin nearly naked and white hairs absent
in paniJiCws) ; foot .^mailer; hair on back and tail shorter; skidl dif-
fering in derail. DilFers from (/eoffroyi and sub.species of Middle
America in nearly uniform black, instead of diverse coloration, vary-
ing from light buff to fenuginous.

< olor. — Face and entire pelage deep glossy black, exeei)t in sonn'
.''I)eciniens having a slight browni.sh tinge on the forehead and a few
^slliti.'<ll hairs on chin and about mouth.

SL-uU. — About a.s in the related form fuJ^clceps. Vei-y similar to that
nf paniscu-s. but pninaxijlao less produced anteriorly beyond plane of
canines; anterior nasal opening less elongated. Closely resembling
that of f/ro/frt^y/^ but rostrum longer; anterior jjiofile risinir h'ss
steeply from ends of premaxillae; auditory hiillar u-iially more
flattened.

.)f'ffMi/r/ments. — Type ( f roni original description): Total length,
l.-"JO mill.: tail. r;J0; hind foot, IGO. Two adult fenuile topotypes,
respect iv«'ly: Total length, 1.150, l.LiOO: tail, GOO, 750; hind foot, 155,
lOO. ^7•////,• 'l\vo adult female topotypes, respectively: Greate.st
length. ll.-).3, 111.7; (»rhital width, 58.9, 62.2; postorbital constriction,

30 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 98

47.9, 44.7; width of brain case, 59.1, 57.2; zygomatic breadth, 65.9,
69.1 ; maxillary tooth row, 28.8, 31.1.

Remarks. — Ateles danensis was based on a single specimen from
eastern Panama, exhibiting characters that prove to be inconstant
in additional specimens examined, and the name must be relegated to
the synonymy of rohustus. Despite the marked contrast in color be-
tween this black form and the red monkey of eastern Panama, the
agreement in nearly all cranial details suggests close relationship.
The auditory bullae are usually more flattened in rohustus, but even
this feature is not always distinctive. To the southward, on the other
hand, close alliance with fmciceps is indicated by the slight brownish
tinge on the forehead in at least one individual, and apparent identity
in cranial details. Allen (Bull. Amer. Mus. Nat. Hist., vol. 35, p. 235,
May 31, 1916) records a specimen of this spider monkey from Barba-
coas, Narino Department,, southwestern Colombia.

Gray (Ann. Mag. Nat. Hist., ser. 4, vol. 11, No. 66, p. 468, June 1873)
has recorded '■'■ Ateles ater'''' from the vicinity of Concordia, Antioquia,
Colombia. Elliot (A review of the Primates, vol. 2, p. 30, June 15,
1913) under the same name lists Cerete on the Rio Sinii, Bolivar,
Colombia, as a locality record. Festa (Boll. Mus. Zool. ed Anat.
Comp. Univ. Torino, vol. 18, No. 435, p. 4, Feb. 11, 1903) likewise iden-
tifies a female taken in forest near the Rio Lara, which empties into an
arm of Golfo de San Miguel, Panama, as Ateles ater. Anthony (Bull.
Amer. Mus. Nat. Hist., vol. 35, No. 20, p. 375, June 9, 1916) refers to
two specimens collected at Tapalisa, Panama.

It is therefore evident that this black spider monkey ranges from
the Serrania del Darion (Cerro Pirre and Tapalisa) southward along
the mountain ranges bordering the Rio Sinii in Bolivar to the moun-
tain range west of the Rio Cauca (Concordia), and southward at least
to southwestern Colombia (Barbacoas).

Specimens examined. — Total number, 7, as follows: Colombia:
Gallera (type locality), 2 (A. M. N. H.). Panama: Cituro, 1 (A. M.
N. H.) ; Mount Pirre, 1; Tapalisa, 1 (A. M. N. H.) : Rio Bayano, 1
(skull only, M. C. Z.) ; Rio Tuyra, 1 (skull only, M. C. Z.).

ATELES GEOFFROYI GEOFFROYI Kuhl

NiCARAGUAN SpIDER MONKEY

Atele[s'\ geoffroiji Kuhl, Beitrage zui' Zoologie nnd vergleichenden Anatomie,

Abth. 1, p. 2G, 1S20'. (Printed also on same page as "Ateles Geoffrey, viihi

species inedita.")
Ateles melanochir Dksmarest, Encyclopedie methodique (Zoologie), Mammal-

ogie, pt. 1, p. 76, 1820. [Based on specimen in Mus. Nat. Hist. Nat. Paris.]

Type locality, unknown.
Ateles melanochir Sclater, Proc. Zool. Soc. London for 1875, pt. 3, No. 27, p. 419,

pi. 48 (col.), Oct. 1875.
Ateles melanocerciis Scht.egel MS., in Jentink, Catalogue systematique des niam-

mif^res, Mus. Hist. Nat. Pays-Bas Leiden, vol. 11, p. 43, 1802 {nomen riudum).

THE SPIDER MONKEYS KELLOGG AND GOLDMAN 31

T'lpc local'itij. — Unknown [Here restricted to San Juan del Norte
(Gi-eytown), Nicaragua; see '•'•AteJc^ hyhrulus''' Sclater, Proc. Zool.
Soc. London for 1862, pt. 2, No. 12, p. 186, Sept. 1862 (specimen from
Rio R.ina, Gorgon Bay. near San Juan del Xorte) : and Salvin In
Alston, Biologia Centrali-Americana, Mammalia, vol. 1, pp. 9-10, Sept.
1879.]

Type specimen. — Female adult, jVIuscum National d'Histoire Natu-
relle, Paris (menagerie specimen acquired in 1819; see I. (icollroy,
Catalogue methodique de la collection des mammiferes, pt. 1 (Cata-
logue des Primates), p. 49, 1851.)

Distribution. — Coastal region bordering San Juan del Norte or Ma-
rina Bay, southeastern Nicaragua; probably ranging across through
the lowlands to the Pacific coast.

General characters. — Distinguished by light bulY, overlaid with
dusky tipped hairs, in general coloration; dark markings on head and
limbs variable and inconspicuous as a rule. Most closely resembling
frantatus of northwestern Costa Rica, but lighter and dark markings
much more restricted. Contrasts strongly with omatus of eastern
Costa Rica in light buff instead of rich rufescent coloration. Differs
from .1. fusciceps rohiLstus of eastern Panama and western Colombia
in diverse instead of nearly uniform black coloration.

Color. — Face and eyebrows varjnng from blackish to a mixture of
black and buffy hairs, those on the brows directed upward to form a
thin ruff; top of head thinly overlaid with blackish or brownish hairs
directed forward, the under color usually light buff. Crown cap
gi'ading from black to buff tinged with brownish; stiff long black
hairs above eyes concealing to variable extent the white or buff tri-
angular forehead patch ; upperside of neck, entire dorsal area, upper-
part of arms all around and of legs to ankles all around, except knees,
light buff tliinly intermixed with long black or brownish hairs; nar-
low areas usually extending 2 or 3 inches up and down over knees
black, the hairs black to roots, but in one specimen the knee patches are
limited to a few overlying dusky hairs; elbows and outer sides of
forearms more or less distinctly blackish; throat and sides of face
and neck silky light buff to light ochraceous-buff ; underparts dull
liglil buff; hands and feet blackish; tail above about like back, below
.somewhat paler along basal portion, but with a narrow line of dusky
hairs bordering callosity jiear tip. In one specimen all the darker
areas are reduced and the general pelage is suffused with liglit oclira-
ceous-buff, iMH-oming near cinnamon-buff on inguinal icgioii. lu an-
other individual small cinnamon-buir ai-cas appear on shouldei-s, inner
sides o[ wrists, sides of feet, and n<'ar middle of underside oi tail.

SkitJl. — Very similar to that of ornntxiH but apparently smalhr.

Mra,'<vrrmrnt}<. — No external measurements available. Skvll : An
adult and a subadult fenialo from Nicaragua, resi)ectively : Greatest

32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. »►:

length, 103.7, 101.5 mm.; orbital width, 58.7, 52.7; postorbital constric-
tion, 46.8, 45.9; width of brain case, 55.7, 56.2; zygomatic width, 61.6,
56.8 ; maxillary tooth row, 28. 26.2.

Remarks. — A. geoffroyi appears to be the first identifiable name
applied to a spider monkey from Middle America. At the time it
was described the locality from which the type specimen came was
unlmown. Much general evidence, however, indicates that this is the
so-called gray spider monkey, also described as A. melanoGhir, now
known to inhabit southeastern Nicaragua, and that this general gray
(really light buff) style of monkey may range across through the
lowlands of the Rio San Juan to the vicinity of Lake Managua and
Lake Nicaragua. It is interesting to note that a very young example
of front atus as figured by Gray (Zool. Voy. Sulphur, vol. 1, No. 1,
Mammalia, pt. 1, pi. 1, Apr. 1843) has essentially the same light-gray
color pattern as that of the adult of geoffroyi. It is probable that
the type of geoffroyi came from San Juan del Norte, and for preci-
sion we restrict the name to the animal that occurs in the vicinity of
that locality. A specimen from Rio Rana, Gorgon Bay, near San
Juan del Norte (Sclater, op. cit., p. 186, 1862) was subsequently listed
by Gray (Catalogue of monkeys, lemurs and fruit-eating bats in the
collection of the British Museum, p. 43, 1870) as ''Ateles hyhridus"
from "St. Juan, Nicaragua." A very good description of this spider
monkey, based on field observations, was given by Salvin to Alston
(Biologia Centrali- Americana, Mammalia, vol. 1, pp. 9-10, Sept.
1879). Confusion of the Nicaraguan race with A. hyhridus by Gray
was undoubtedly due to external resemblance, which is rather strik-
ing; but cranial characters indicate close alliance with geographic
neighbors that are markedly different in coloration. A specimen
(M. C. Z. No. 29626) from El Valle del Anton, near the Pacific coast
east of the Azuero Peninsula in Panama, agrees closely in general
coloration with those from Nicaragua, but the black on the crown
patch and on forearms is more extensive. Whether this animal was
native to the locality or brought from some other region seems some-
what uncertain.

Speciinens examined. — Total number, 10, as follows: Nicaragua:
Managua, 6 ; no definite locality, 4 ( 1, A. M. N. H.) .

ATELES GEOFFROYI VELLEROSUS Gray

Mexican Spider Monkey

Ateles vellerosus Gray, Proc. Zool. Soc. London for 1865, pt. 3, No. 47, p. 773,

Apr. 1866.
Ateles veUerosus Sclateb, Proc. Zool. Soc. London for 1872, pt. 1, No. 1, p. 5, pi. 2

(col.), June 1S72. (Believed by A. Boucard to have been procured near

Acapulco, Mexico.)
Ateles neglectus Reinhakdt, Vid. Medd. Nat. Foren. Kjobenbavn, ser. 3, vol. 4

(1872), Nos. 6-9, p. 150, 1873. Type locality, Mirador, Veracruz, Mexico.

THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 33

Atdcs tricolor IIoujstek, Proc. Bicil. Soc Washington, vol. 27, p. 141, July 10,
1914. Type locality. Hacienda Santa Ktigonia. S miles north of Tapanatepec,
southeastern Oaxacn.

Type localify. — ?Biazil. [Here restricted to Mirador, about 15 miles
northeast of Huatusco, Veracruz, Mexico; altitude 2,000 feet.]

Type spccitncn. — No. . British Museum (Natural History).

DistHbution. — Unbroken forests of Veracruz and eastern San Luis
Potosi and southeastward! lhr()U<xh Tabasco, across the Isthmus of
Teluiantepec in eastern Oaxaca, to Honduras and El Salvador, except
for highlands of Guatemala.

General characters. — A subspecies distinguished by combination of
black or brownish-black top of head, neck, and shoulders, in contrast
with buffy lumbar region, and pinkish-buff to cinnamon-butf under-
parts. Differs from yucatafnensis of Quintana Roo in deeper buff
underparts (underparts in yucatanensis silvery white or light buff).
Differs from pan of Guatemala in greater contrast in colors of upper-
parts (in pan the upper back and lumbar region are more nearly uni-
form blackish in tone).

Color. — Top of head, neck, shoulders, outer surfaces of fore and
hind limbs, and forearms all around varjnng from black to brownish
black, passing near middle of back into cinnamon-buff or cinnamon and
extending across lumbar region and including hips, the latter colors
somewhat darkened along median line by overlying brownish hairs;
underparts in general, including inner sides of arms in a strip nar-
rowing from armpit to a point near elbows and on inner sides of
legs to near ankles, varying from pinkish buff to cinnamon-buff, be-
coming near cinnamon on sides of body; underside of neck varying
from grayish to a mixture of buffy-gray and brown; face blackish,
but lips and small tufts near sides of mouth are didl whitish in some
specimens; hands and feet black; tail above black or brownish black,
mixed below with buffy Juiirs in a median lijio gradually narrowing
from base to near callosity.

SkxiU. — Rather narrow, but closely similar in general to those of
the other subspecies. Comi)ared witli yurafanrvsf.s the frontal region
of the skull is slightly flatter, less convex along median line anteriorly,
as shown in profile.

Mea.'<nrem/'nt!<. — Two adult males from Sania Kfigenia, Oaxaca,
resi)ectively: Total length, 1,200, 1,260 mm.; tail vertebrae, 7G0, 705;
hind foot, 17G, 178. Average of 7 adult females from Santa Kligenia,
Oaxaca: Total length, 1,204 ( 1, 140-1, .'^01 ) ; tail vertebrae, 751 (700-
7ft0) ; hind foot, 170 (17H-1H5). SJcnll (three adult males from Sanla
Kfigenia. respectively): Greatest lengtli. IlO.l. 112.5, 110.9; orbital
width, 60, 56.3, 57.2; post orbital const i-iet ion, 51.3, 47.6, 47; width
of brain case, 59.3, .58.5, .59.6; /ygoniatic wi.Ith, 70.5, 63.9, 68.r);
maxillary tooth row, 28.9, 29.9, 29.9. Average of 7 adult females

34 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

from Santa Efigenia, Oaxaca: Greatest length, 108.2 (105.2-113.4);
orbital width, 56.2 (54.5-58.8) ; postorbital constriction, 46.7 (45.5-
48.7) ; width of brain case, 58 (55.9-60.2) ; zygomatic width, 63.5
(59.2-66.9) ; maxillary tooth row, 28.5 (27.2-29).

Remarks. — A. velUrosus was originally assigned by Gray to Brazil,
but this appears to have been an error, as pointed out by Alston
(Biologia Centrali-Americana, Mammalia, vol. 1, p. 10, Sept. 1879).
By some subsequent authors, including Alston, the name was applied
to spider monkeys from Mexico without definite locality. Inciden-
tally, the plate illustration of Sclater {op. cit.) closely resembles this
form. It represents a specimen believed by Boucard to have been pro-
cured near Acapulco, Guerrero, where it could not have been native,
as no monkeys occur in that general region. Under the name velJerosus
Thomas (Proc. Zool. Soc. London for 1890, pt. 1, p. 72, June 1890)
recorded specimens taken in Veracruz at the following localities : Raya
de Boca Agustin, Misantla ; boundaries of Misantla and Jalapa ; and
Hacienda de Tortugas, Jalapa. At the time A. tricolor from south-
eastern Oaxaca was described (op. cit) by Hollister, veUerosus was not
recognized by him, and the name A. neglect us of Veracruz was over-
looked. The following year, however, Hollister (Proc. Biol. Soc.
Washington, vol. 28, p. 142, 1915) regarded tricolor and neglectus as
identical. The various descriptions of veUerosus apply well to the
series of specimens now available from both Veracruz and Oaxaca,
under which we, therefore, include both neglectus and tricolor as
synonyms. For precision we would restrict the name veUerosus to the
form known to occur near Mirador, Veracruz, the type locality, inci-
dentally of the later name, neglectus.

The range of subspecies veUerosus marks tlie northern limit of the
group in Middle America. It inhabits unbroken forested regions
from near sea level to about 4,000 feet altitude, including some of
the areas of heaviest and most continuous rainfall in Mexico. William
Lloyd, a field agent of the former Biological Survey, writing of the
mammals of the vicinity of Victoria, Tamaulipas, in 1891, stated
that "a monkey has been said to have been taken a little south of here,
but the most authentic information says they are not found north of a
point west of Escandon, rather more than 50 leagues south." It is
possible that spider monkeys ranged north into southern Tamaulipas,
but information obtained in 1898 indicated that the northern limit was
then near Xilitla, southeastern San Luis Potosi.

The German naturalist Ferdinand Deppe, in 1825, purchased a live
Ateles in Alvarado, Veracruz, which had been caught by a Mexican
about 20 hours distant from that city. Afterward, Deppe (Sclater,
Nat. Hist. Rev., vol. 1, No. 4, pp. 508-509, Oct. 1861) while en route
from Caxaia to Alvarado, observed a "great number" of Ateles in a
forest near Valle Real.

THE SPIDER MONKEYS KELLOGG AND GOLDMAN 35

One specinien fn)iu Juchitan, Oaxaca, has silvery whitish under-
parts, and another from the same locality is normally colored. A
very y()un<2: individual from Santa Eli<r('nia, Oaxaca, does not differ
aj>preciably in color from adults from the same locality. One very
younjnr one from Tehuantepec has the upperparts normally colored,
and another from the same locality is uiuisually daik. tlie luinl)ar re-
gion nearly as dark as the upper back.

Specimens examined. — Total numl)er, 59, as follows: Mexico:
Oaxaca : Juchitsin, 3 (A.M.X.H.) ; Oaxaca, without delinite locality,
4 (A. M. X. H.) ; Santa Eligenia, 13; Tehuantepec, 4 (one skull without
skin. A.M.X.H.; one skull without skin, U.M.M.Z.) ; Tuxtepec, 2.
Tahax'o: Teapa. 1. Veracruz : Barranca de Boca, Canton de Jalapa,
1 (skin only) ; Cuatotolapan,2 (skulls only, U. M. M. Z.) ; Pasa Xueva,
VJ (2 skulls without skins, A. M. N. H.). Hondukas: Tegucigalpa:
Cantoral. 1 (A.M.X.H): Guaymaca, 2 (A.M.X.H.). Olancho:
C'atacamas, 1 (A.M.X.H.). Ocotepeque : El Chorro, east side of
Ocotepeque, 4.500 feet, 1 (skin only, A.M.X'^.H). El Salvador: San
Mi^nicl : Lake Olomega [=Laguna Lomego], 5 (C. D.),

ATELES GEOFFROYI YUCATANENSIS, new subspecies

Yucatan Spider Monkey

Tj/pe locaVti}/. — Puerto Morelos, northea.st coast of Quintana Roo,
Mexico; altitude 100 feet.

Type specimen. — Male adult, skin and skull; U. S. N. M. No. 108531
(Tiiological Surveys collection) ; collecled April 2, 11)01, by E. AV.
X(Is(jn and E. A. Goldman; original number 14052.

Distribution. — Forests of the Yucatan Peninsula, northeastern
Guatemala, and ])robably adjoining parts of British Honduras;
doubtless intergrading to the .southward with vcUcrosus.

General characters. — A rather small, slender, light-colored race,
with entire underparts silvery whitish or very pale buff, pelage short
and thin. Size about as in vrUiVoimfi of Veracruz but decidedly paler,
especially on underparts where in typical specimens a whitish silvery
tone extends to neck and inner sides of limbs; underside of tail
'•ream-buff to near callosity; frontal outline of skull more i)rominent.

Color. — Type: Top of head, neck, and outer surfaces of fordimbs
brownish black, pa.s.sing gradually through cinnamon-drab and light
drab on shoulders and anterior part of back to near olive-bull on
hips, the light drab extending along nie<lian line to base of tail: fa<e
mainly dusky, but cheeks whitish; knees brownish black: res( of outer
surfaces of hind limbs cinnamon-drab; entire under|)arts inebHJing
underside of neck, inner surfaces of limbs to wrists and ankles thinly
haired, between crcam-buir and silvery whili.sh; hands and feet black-
ish; tail cinnamon-diab above and on sides. becoming cream-bulf below
along median line from base to near callosity.

36 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

SktiJl. — Similar in size to that of veUerosus^ but frontal profile more
convex anteriorly; dentition rather light.

Measurements. — Type : Total length, 1,150 mm. ; tail vertebrae, 766 ;
hind foot, 181. Two adult male topotypes, respectively : Total length,
1,141, 1,090; tail vertebrae, 744, 817; hind foot, 167, 174. Three adult
female topotypes: Total length, 1,17^, 1,150, 1,056; tail vertebrae, 756,
766, 713; hind foot, 169, 171, 165.

Skull (type and two adult male topotypes, respectively) : Greatest
length, 109.7, 105.7, 107.8; orbital width, 57.2, 53.9, 55.6; postorbital
constriction, 47.8, 44.8, 45.8; width of brain case, 59.5, 56.8, 57.3;
zygomatic width, 65.3, 61, 66.3; maxillary tooth row, 27.3, 27.5, 28.
Three adult female topotypes, respectively: Greatest length, 104.8,
108.5, 104.8; orbital width, 52.8, 54.1, 53.1; postorbital constriction,
44.6, 45.8, 45.6; width of brain case, 55.1, 57.2, 57; zygomatic width,
58.3, 60.2, 60.3; maxillary tooth row, 25.7, 29.8, 28.7.

Remarks. — The spider monkey of the Yucatan Peninsula is readily
distinguished by the silvery whitish underparts. Specimens from
Apazote, Campeche, and Uaxactum, Guatemala, are referable to the
present form, but in slightly darker and more buffy underparts they
indicate gradation toward veUerosiis. One topotype and a young
without definite locality have sparse white hairs on the anterior part
of the crown and a vestige of a white forehead patch partially con-
cealed by stiff black hairs.

Specimens examined. — Total number, 16, as follows : Mexico : Cam-
peche : Apazote, 1. Quintana Roo : Puerto Morelos, 6. Guatemala :
Uaxactum, 8 (U. M. M. Z.) . No definite locality, 1 (A. M. N. H.) .

ATELES GEOFFROYI PAN Schlegel

Guatemalan Spider Monkey

Ateles pan Schlegel, Mus. Hist. Nat. Pnys-Bas Leiden, vol. 7, livr. 12 (Monogr.
40, Simiae), p. 180, 1876.

Type locality. — Coban, Alta Vera Paz, Guatemala.

Type speehnen. — Cotypes, adult male and two adult females,
Museum d'Histoire Naturelle des Pays-Bas, Leiden.

Distribution, — Mountains of central Guatemala; doubtless inter-
grading with vellerosus.

General characters. — A very dark spider monkey with lumbar region
only slightly, if at all, lighter than shoulders and thus lacking the
contrasting dorsal areas presented by vellerosus and other Middle
American races ; pelage long and dense.

Color. — Face, top of head, shoulders, outer surfaces of limbs, fore-
arms all around, feet, and entire tail black or brownish black. In one
specimen the dark color extends nearly uniformly over all of the
upperparts, but in two others the lower part of the back is suffused
with cinnamon partially concealed by overlying dusky hairs; under-

THE SProER MONKEYS — KELLOGG AND GOLDMAN 37

side of neck and throat thinly covered with mixed brownish and buffy
liairs; rest of underparts, incKiding a strip narrowing from armpit
to a point near inner side of elbow, and inner sides of hind limbs to
ankles between pinkish butf and cinnamon-bufT, becoming deep cinna-
mon or rusty along sides of body.

ShuII. — About as in vcllerosus.

Mea-s'iircin-cnts. — No external measurements available. Skull: Two
adult females from Guatemala, respectively: Greatest length, 1G9.1,
lO.'i,!; orbital width, 57.3, 59.4; postorbital constriction, 48, 50; width
of brain case, C0.*2, 62.4; zygomatic width, G6.8, 68.8; maxillary tooth
row. 27.8. 27.5.

Remai'ks. — Three specimens exhibited by the Guatemalan Govern-
ment at the World's Columbian Exposition were presented to the U. S.
National Museum. These Avere labeled ''Guatemala" without definite
locality but are believed to be from Alta Vera Paz and probably repre-
sent typical Ateles pan. Aside from the very dark coloration, the coat
is long and heavy, indicating that they came from a high elevation in
the mountains. The skulls indicate close agreement with vellerosus
in size and general details.

Specimens examined. — Total number, 3, as follow's: Guatemala:
Alta Vera Paz, 3.

ATELES GEOFFROYI FRONTATUS (Gray)

Black-foreheaded Miriki or Spider Monkey

EriodcH fronintun C;il.vy. Ann. Miig. N.it. Hist., ser. 1, vol. 10, No. G'), p. 250,

Dec. 1842.
lirarhj/tclrs frontatun Or,\y, Zoology of the voyage of H. M. S. Sulphur, vol. 1,

No. 1, Maiumalla, pt. 1, p. 9, pi. 1 ( col. ) , Apr. 1843.

Type locality. — South America [ = harbor of Culebra, Le6n = Cule-
bra. Bay of Culebra, (iuanacaste, northwestern Costa Rica, fide Gray,
Zool. Voy. II. M.S. Sulphur, vol. 1, No. 1, Mannnalia, pt. 1, p. 10, Apr.
1843.]

Type specimen. — Female adult wit h young shot by Capt. Sir Edward
Hr-lcher; British Museum (Natural History) No. .

Dint rihut ion. — Noi'tli western Costa Rica and cxtrcinc western and
northern Nicaiagua.

General cJuiracters. — Similar in j)att<'ni of coloration, that is, the
normal restriction of bhuk areas to top of head and irregularly to
•juter surfaces of limbs, to yeoffroyi of .southea.stern Nicaragua, but
bo<ly darker, the ground coloi* of iipperparts near buckthorn brown
or Mars l)rown and of underi)aits honey yellow to tawny, instead of
light buir. DifTers frf)m p(rn<nnei)»iH of Panama in brownish instead
of deep ferruginous general body color. Differs from vellerosv^ of
Veracruz in the restriction of black areas on anterior ])ait of back and
more yellowish tone of lumbar regic^n.

38 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Color. — Face, forehead, top of head, and in some specimens back
of neck and shoulders black; anterior part of crown patch more or
less suffused with cinnamon-buff, owing to light basal color of hairs,
and tending to form a transverse band across forehead ; side whiskers
on face light buff to cream-buff; upperparts in general buckthorn
brown to Mars brown, sparsely or noticeably intermixed with blackish
hairs (especially on midline of back in some individuals) ; belly honey
yellow to tawny, extending downward on inner surfaces of legs to
ankles, and a lighter tone near cinnamon-buff extending downward
on inner surfaces of arms to wrists; outer surfaces of arms and legs
usually black (in some individuals restricted more or less to elbow
and knee patches) ; side whiskers on face light buff to cream buff;
tail usually sharply bicolor, black or dusky above to tip and tawny
below to callosity (when otherwise, mixed light and dark hairs cover
the tail above and below) .

Skull. — About as in fanmnensis., averaging somewhat broader than
in geoffroyi or vellerosus.

Meamremsnts. — No external measurements available. Skull: An
adult male from Peiia Blanca : Greatest length, 109.3 ; orbital breadth,
59.4; postorbital constriction, 49; width of brain case, 58.6; zygomatic
breadth, 69.6; maxillary tooth row, 28.4. Two adult females from
Lavala and Tuma, Nicaragua, respectively : Greatest length, 117.3, 112;
orbital width, 64.2, 60.2; postorbital constriction, 51.1, 51; width of
brain case, 62.8, 59.8 ; zygomatic breadth, 71.4, 65 ; maxillary tooth row,
29.6, 28.4.

Remarks. — A. geoffroyi front atus is assumed to be a recognizable
subspecies, although aside from the type in the British Museum, from
northwestern Costa Rica, little is definitely known of its characters or
distribution. Certain specimens from Costa Rica without definite
locality, and from various localities in Nicaragua, are tentatively re-
ferred to frontatus., although in none of these is the black on the head
so restricted as in the type as figured by Gray {op. cit.). The color
pattern in this form suggests close relationship to geoffroyi^ which
might also be expected on geographic grounds. A very young individ-
ual (U. S. N. M. No. 61208) from Costa Rica, without definite locality,
closely resembles Gray's figure just mentioned in light coloration. A
skull of an old female from Lavala, Nicaragua (A. M. N. H. No.
28419), is one of the largest examined from anywhere in Middle
America.

Specimens examined. — Total number, 38, as folloAvs : Costa Rica : 3
(without definite locality) . Nicaragua : Lavala, 2 (skulls only, A. M.
N. H.) ; Peiia Blanca, 3 (A. M. N. H.) ; Rio Siquia, 2 (skulls only,
U. M. M. Z.) ; Rio Yoya, a tributary of Rio Princapolca, 25 (skulls
only, A. H. S.) ; Tuma, 2 (skulls only, A. M. N. H.) ; Uluce, 1
(A. M. N. H.).

THE SPIDEK MONKEYS KELLOGG AND GOLDMAN 39

ATELES GEOFFROYI ORNATUS Gray

Costa Rica Ornate Spider Monkey

Hrlcs ornatuK Guay, Cntalopiie of monkeys, It'inurs and frnit-catitif; l)ats in the
collection of the British Museum, p. 44, 1870.

7'i/pc locality. — Unknown. [Here restricted to Cual)ro, Tahunanca
region, southeastern Costa Kica.]

Type apecimen. — British Museum (Natural History) No. .

Distribution. — Eastern sh)pe of central cordillera of Costa Rica;
doubtless intergrading ^vith panamensis on Pacific side of the central
mountain range.

General characters. — A dark golden-yellowish subspecies, the upper-
parts in strong light having a glossy, golden-yellow sheen, owing to the
yellowish subterminal bands of hairs. Resembling panamensis of
I*anama and western Costa Rica but brighter, more golden yellowish in
tone, the back, however, more obscured by overlying black-tip])ed hairs,
throat lighter, underside of tail and inner side of upper arm lacking the
deep '"ferruginous" of panamensis. Differs from frontatus of Guana-
caste, northwestern Costa Rica, in greater extension of black area on
limbs and over back of neck and shoulders, and black-tipped hairs
overlying and tending to modify the golden-yellowish tones on back.
Contrasting strongly in golden yellowish, instead of light bully colora-
tion, with geofroyi of the coast region of southeastern Nicaragua.

Color. — Face, top of head, forearms all around, outer sides of legs,
hands, and feet black; upperpai-ts from shoulders to hips and base of
tail glossy golden-yellow or apricot buff, varying in some specimens
to cinnamon-rufous, more or less overlaid with blackish hairs; chest
near cinnamon-bull", the rest of underparts and Hanks becoming near
tawny or cinnamon-rufous, this general color extending down along
inner sides of legs in some sperimens nearly to ankles; throat and under-
side of neck near pinkisli buff; tail black ai)<)ve, more or less mixed
with tawny below from base to near callosity.

S/ntll. — About like that of panamensis-^ similar to thai of geoffroyi.,
but apparently .somewhat larger.

M eajturemenis. — No external measurement s ava i lable. Skull : Three
adult males from Talamanca i-cgion. Costa Rica, resjx'ctivdy : Cxreatest
length, 110.2, 111.2. 1 i:',.H mm. ; ori)ital width, r.:}.;^ (',1.7, C.'i; postorbital
constriction, 45.2, 45.7, 49.0; width of brain case. Gl.O. 58. C1.7; zygo-
matic bivadth, 71.5, CH.2. 71.0; maxillary tooth row, 27. G, 30.f), 27.9.
Average of adult females from Talanianca region, Costa Rica:
CM-eatest length, 108.1 (104.5-114.4); orbital width. C.l.G (5G.4-64.1);
postorbital constriction. 47.4 (4-1.H-49.1 ) ; width of brain case, .5H.7
(55.7-61.3); zygomatic width, GG.8 rG;'..7-70.7) : niaxilhirv tooth i-ow.
27.5 (26.3-29..5).

Rryturrkx. — Much confusion has existed regarding the status of this
brightly colored sul)species. Specimens that we believe belong here

40 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

have commonly been referred somewhat doubtfully to A. geofroyi
Kuhl. The description of A. omatus by Gray {op. cit.), from an
unknown locality, applies so well to specimens from eastern Costa Rica
that their identity seems unmistakable. A more detailed description
of the type than that published by Gray is given by Elliot (A review
of the Primates, vol. 2, p. 45, June 1913) . The glossy quality as well as
bright colors of the pelage suggests the propriet}^ of the name omatus.
The contrast in color with plain buffy geo-ffroyl of southeastern Nica-
ragua is very great, but close agreement in cranial characters and other
evidence indicate probable intergradation.

Specimens examined. — Total number, 14, all from Costa Eica as
follows: Cataratas, Rio San Carlos, 2 (A. M. N. H.) ; Cuabre, Tala-
manca region 1 (A. M. N. H.) ; Guapiles, Limon Province, 1 (A. M.
N. H.) ; Santa Maria, 37 miles south of San Jose, 3 (skulls only, A. M.
N. H.) ; Talamanca region, 7.

ATELES GEOFFROYI PANAMENSIS, new subspecies

Red Spider Monkey; Mono Colorado

Type locality. — Cerro Brujo, about 15 miles southeast of Portobello,
Province of Colon, Panama ; altitude 2,000 feet.

Type specimen. — Female adult, skin and skull, U. S. N. M. No. 171489
(Biological Surveys collection) ; collected June 8, 1911, by E. A. Gold-
man ; original number, 21165.

Distribution. — Forested regions of Panama east of the Canal Zone
(Cordillera de San Bias) , and west through Chiriqui to central western
Costa Rica.

General characters. — A. rather large, deeply rufescent race. Very
similar to omatus of the Caribbean slope of Costa Rica, but reddish
tone more intense, the back less obscured by overlying dusky hairs;
inner side of upper arm pinkish cinnamon to ferruginous. Differs
from azuerensis of Azuero Peninsula, Panama, in deep reddish instead
of cinnamon or tawny general coloration.

Color. — Type: Upperparts from back of shoulders to base of tail,
backs of thighs, and sides of body ferruginous, slightly obscured by
black tips of hairs; outer surfaces of fore and hind limbs either black
or blackish to knees and elbows ; outer surfaces of limbs below knees
and elbows either black or with varying admixture of black and ferru-
ginous hairs, the hairs banded but black at tips; underparts thinly
haired, pinkish cinnamon on chest and along inner sides of hind limbs
to ankles, becoming deep, clear ferruginous or dark dusty reddish over
abdomen ; face, crown cap on head, and median streak on back of neck
blackish ; sides of neck covered w^ith a mixture of pinkish buff, cinna-
mon-buff, and blackish hairs ; chin nearly naked ; throat and under side
of neck thinly clothed with light cinnamon-drab or light brownish
hairs; tail above ferruginous heavily mixed with black on basal two-

THE SPIDER MONKEYS — KELLOGG AND GOLD^LVX 41

thirds, bec<)inin<r clear black toward tip, bolow deep ferruginous to near
callosity where blackish hairs extend all around. The ji;round color
of body in other specimens grades from dark rusty reddish to ferru-
<:in(»us or burnt sienna, and a narrow ferrurrinous or cinnamon-buff
streak usually extends long inner side of upper arm from armpit to
elbow.

/Sk-i/IL — Very similar to that of groffroyi, but brain case rather broad.

Measurements. — Type: Total lenirth, 1.-2H0 mm.; tail vertebrae, 840;
hind foot, 194. An adult male and ftinale from Cerro Azul, respec-
tively: Total leuirth, 1.280. 1.225; tail vertebrae. 786, 785; hind foot,
183, 176. Skull (type): Greatest length, 112.8; orbital width, 62;
postorbital constriction, 52.4: wiilth of brain case, 61.2; zygomatic
breatlth, 68: maxillary tooth row, '^7.3. An adult nuile and female
from Cerro Azul. respectively: Greatest length, 114.2, 109.1; orbital
width, 62.7, 59.4; postorbital constriction, 53, 50.3; width of brain case,
63.2. 61.4: zygomatic width, 70.2, 69.6; maxillary tooth row 31, 28.1.

/i'> nurrkft. — This is the monkey conunoidy known as ''mono colo-
ra<l<i" in Panama and western Costa Rica. It is the most intensely
reel of all the races of Middle America, but despite this it has been
generally confused with the liglit bulFy form, geoffroiji, of Nicaragua.
While the ground color of most of the body appears to be ferruginous
to burnt sienna in ordinary light, in strong light and particularly in
direct sunlight individual hairs on the upperjKirts are .seen to have
glos.sy subtcrminal bands giving that part of the pelage a golden-
yellowish sheen, a peculiarity still more pronounced in ornaPufi and
not shared with the more northern races. The specimens from
Chiriqui and as far west as central-western Costa Rica agi'ee closely
in color with those from the (^anal Zone in the vicinity of the tyi>e
locality. In one sjiecimen of a series of six skins from Pozo Azul,
Costa Rica, a white patch on the forehead is nearly concealed in front
by loiiL'er black-tipped hairs and the cheeks are dull whitish or ]>alo
huffy.

Specimens examined. — Totiil mimbci-, 21. as follows: Cosia Rir.v :
Canas Gordas. Puntarenas, 3 (A. M. X. II.) ; Kl Pozo, 12 niiles above
n)oiifb of Ri'o Grande de Terraba, 1 (slcin only, A. M. N. II.) ; Pozo
A/nl. 10 miles above mouth of Rio Grande de Pirris, 12 (3, C. M.; 4
skins with skulls and 5 extra .skulls, A. M. N. II.). P.\nam.\: Cerro
Azid, near head of Chagres River, 2; Cerro Brujo {{\\)v locality), 1;
Coto, Chiriqui, 2 (M. C. Z.) ; Rfo Pequeni, Canal Z(me, 2 (M. C. Z.) ;
Snn Juan. Chagres River. 1.

ATELE.S GROFPROVI A7.URRKNSIS Bole

Azi'ERO Spider Monkey

Atrlrs n:urrfn.«iif Hoik. S<1. I'lihl. flfvi-liiinl Mus. Nut. Illst., vol. 7, p. 149, AuR.
.31. i;t.37.

42 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Type locality. — xVltos Negritos, 10 miles east of Monti jo Bay (part
of the spur forming south drainage divide of Kio Negro), Mariato
Suay Lands, Azuero Peninsula, Veraguas Province, Panama; alti-
tude, 1.500 feet.

Type specimen. — Female adult, Cleveland Museum Natural His-
tory No. 1235.

Distribution. — Forested mountains of the Azuero Peninsula, Pan-
ama, probably in deeper forests on both slopes, but known only from
the western (Veraguas) side from the vicinity of Ponuga southward.
Possibly ranging west to Burica Peninsula on Panama-Costa Rican
boundary.

General characters. — A subspecies distinguished from neighboring
forms by lighter tawny or ochraceous-tawny general coloration.

Color. — Face blackish ; top of head and back of neck varying from
deep black to a mixture of black and cinnamon-buff, with black pre-
dominating; upperparts in general ochraceous-tawny thinly inter-
mixed with black-tipped hairs; shoulders and outer surfaces of arms
and legs black ; belly similar to back, but paler owing to absence of
black-tipped hairs; throat and chest cinnamon-buff, this general tone
extending downward to wrists and ankles on inner surfaces of limbs;
tail above black, below tawny to proximal end of callosity.

Skull. — Rather small and narrow, but very similar otherwise to
that of panamensis.

Measurements. — An adult male and female from type locality, re-
spectively: Total length, l.lYO, 1,166 mm.; tail, 720, YOl; hind foot,
177, 168. Skull (t3^pe, from original description) : Greatest length,
110.8; width of brain case, 56.8; zygomatic breadth, 65.1; maxillary
tooth row, 27.7. An adult male from type locality : Greatest length,
109.8 ; orbital width, 57.7 ; postorbital constriction, 44.1 ; width of brain
case, 58.7 ; zygomatic breadth, 67 ; maxillary tooth row, 29.7.

Remarks. — This subspecies is definitely known only from the heavily
forested mountains of the Azuero Peninsula, where it appears to be
isolated, but may prove to have a more extended range near the Pacific
coast of western Panama. A series of 25 skulls from the Burica Penin-
sula, near the Panama-Costa Rican boundary, said to be from light-
colored skins not saved and, therefore, not available for examination
by us, are indistinguishable from those inhabiting regions as far north
as Nicaragua. For reasons of geography and light color reported we
refer them, tentatively, to as/uerensis.

Specimens examAned. — Total number, 27, all from Panama, as fol-
lows: Altos Negritos, Azuero Peninsula (type locality), 2 (CM. N.
H.) ; Rio La Vaca, near Puerto Armuelles, Burica Peninsula, 25 (skulls
only,A.H. S.).

THE SPIDER MUX KEYS — KELLOGO AND GOLDMAN 43

ATELES GEOFFUOYI GRISESCENS Gray

Hooded Spider Monkey

Atclfs griscscens Gray. rroe. Zool. Soc. Lnmloii for 1SG5, pt. 3, p. 732, Apr. 1806.
Atil> s cumJlatu-^ Quay. Troc. Zool. Soe. London for l.S(>"), pt. 3, p. 733, Apr. ISfiO.

Type locality, unknown.
Atclvs oucullatiis Sclatix, rmc. Zool. Soc. London for 1871, pt. 1, p. 223, pi. 14

(col.), June 1871.
Alclct mclatiochir Sclater, Proc. Z<iol. Soc. London for 1875, pt. 3, p. 419, pi. 49

(col.), Oct. 1875.

Type 7o(?a7^7^/.— Unknown. [Here restricted to Rio Tuyra, south-
eastern Panama.]

7\)/pe si>eci77un. — Bi-iti.-«li Aruseum ( Xatural History) No. .

D'.Mtributioj}. — Presumably the valley of Rio Tuyra and probably
southeastward through the Serrania del Sapo of extreme southeastern
Panama and the Cordillera do Baudo of northwestern Colombia.

Gineral characters. — Adults apparently characterized l)y long, lax
pelage and peculiar du.sky coloration, with a general admixture of
yellowish gray or golden hairs, the hairs on upperparts golden at the
base. The skull indicates close relationship to panarrieiisis^ despite the
contrast in the deep reddish color of the latter.

Color (type of nwuU-atufi). — Skin around orbits and on nose bare
and of a browni.sh flesh color with darker freckles intermixed; cheeks
and lower jaw nearly bare of hair, but .skin more decidedly of a black
shade (Murie, Proc. Zool. Soc. London for 18G5, pt. 3, p. 739, Apr. 18G6) .
Hairs very long and lax, those on head projecting forward over fore-
head; crown and neck black; rest of body above and below, limbs, and
tail black intermixed with yellowish-gray or golden hairs, so numer-
ous as to give a pale yellowish-brown coloration to back (not far
removed from color on the bases of the hairs in similar parts of A.
f/riJiescens) ; hands and feet black, bases of hairs on hands golden, but
not on those of feet which are black to the roots (Elliot. A review of
the Primates, vol. 2, pp. 38-39, June 1913). Barbour (Journ. Mam-
malogj'. vol. 13, Xo. 4, pp. 307-3G8, Nov. 1932) lias conunented on the
.skin and skull of a spider monk(n' (M. C. Z. No. 27490) obtained at
Chepigana in Darien. Panama. In this inunature specimen, the entire
upperj)arts, including (he outer .surfaces of fore and hind limbs as well
as the upl)er surface of the tail, are dusky, the darker hairs nearly sooty
black on the distal two-thirds and thinly interspersed with these on
head, neck, back, thighs, and upper surface of tail are old gold or silvery
hairs; the cf)loration of the uj)perparts is further modified by a lighter
sulTusion resulting from (he old-gold color of the basal third of these
liairs showing through. A partially concealed bulfy spot is present on
the forehead. Tlie thrf)at and chest are thiidy covered with sooty
black hairs. On the belly, inner surfa<-es of hind limbs, and under
surface of tail the hairs art- ilull ciniiamon-bufT tipped with sooty

44 PROCEEDINGS OF THE NATIONAL MUSEUM vol.86

black. The inner surfaces of fore limbs are noticeably lighter than
the outer and also somewhat lighter than chest. The hairs on hands
and feet are lighter at the base than at the tip. The hairs on the
forehead project forward as on the type specimen.

Skull. — Similar to that of panamensis.

Measureme^its. — An adult male (type of cueuUatiis) : Head and
body, 431±mm.; tail, 698; hind foot, 159 (fkle Murie, 1866, op. cit., p.
T39). jSkuIl: Immature specimen from Chepigana: Greatest length,
103; orbital breadth, 52.3; postorbital constriction, 47.6; width of
brain case, 55.6 ; zygomatic breadth, 60 ; maxillary tooth row, 26.8.-

Remarks. — A somewhat grizzled spider monkey obtained by E.
Greey, an officer on the Eoyal West-Indian Mail Co.'s steamship
Shannon at St. Thomas, Virgin Islands, for the Zoological Society of
London became the basis of A. grisescens of Gray. After the death of
the animal, the skin and skull were acquired by the British Museum
(Natural History), and the skin was compared by Sclater (Proc. Zool.
Soc. London for 1871, pt. 1, p. 223, June 1871) with the skin of a some-
what similar half -grown male purchased from a London dealer in 1869.
This specimen, also evidently of unknown native origin, he regarded
as probably identical. The type of this spider monkey, according to
Elliot (A Review of the Primates, vol. 2, p. 37, June 1913), is entirely
black except for the yellowish-brown tint of the underside of the tail,
but intermixed with the hairs of the upperparts are long, gray silvery
or golden hairs. The latter are not sufficiently numerous to affect the
general blackish coloration except on the shoulders, lower back, and
the limbs where the hairs are yellow or golden at the base.

The type of cucullatus^ a male now in the British Museum (Natural
History), had a small tubercle representing the thumb, and Sclater
{op. cit..f 1871, p. 224) remarked that he had "some reason to suppose
it may be from the northern coast of Colombia, as ... a black spider
monkey with long hair over its head is occasionally brought for sale
into Cartagena."

Attention is here directed to two black-handed spider monkeys fig-
ured by Sclater {op. cH.^ 1875, pi. 49), which were received likewise
from officers of the West Indian Mail Co. The color of the face and
nose closely resembles that described for grisesoens^ and in addition the
distal black color of the hairs on the upperparts does not entirely con-
ceal the yellowish basal color of these hairs. If these specimens are
correctly allocated to grisescens, it would seem that this spider monkey
is subject to considerable variation in external appearance. Owing to
the lack of adequate material for study the status of this form remains
uncertain.

Specimens examined. — Total number, 2, as follows : Panama : Che-
pigana, Darien, 1 (M. C. Z.) No definite locality, 1 (yg., A. M. N. H.) .

• Hindermost molar has not erupted.

THE SPIDER MONKEYS — KELLOGG AND GOLDMAN 45

INCERTAE SEDIS

(?) ATELES RUFIVENTKIS Sclatcr

Atih\s viUn-o.stci ScL-MKR, I'mc. ZuoL Soc. London for 1871, pt. 2, p. 478, Aug. 1.S71

{tier Gray).
Atrhs rufii'ftitrL'* Sci.ateu. Troc. Zool. See. London for 1872, pt. 2, p. G88, pi. 57

(coL), Nov. 1872.

Type locaJf'fr/. — Rio Atrato, Darien, Colombia.

Type spi'chnrii. — Britisli Miisoinn (Xatural History) No. .

Remarks. — Sclater (Ptoc. Z(K)1. Soc. London for 1871, pt. 2, p. 478,
Au^. 1871) stated that the type of Afeles iniflventris^ a young female,
with body length 12 inches, tail 15 inches, was obtained at Colon, Pan-
ama, by an officer of the Royal West Indian Mail Co. and tJiat it had
been brought from the Rio Atrato by one of the American party (pre-
sumably the Dan'en E.xploring Expedition under the command of
Comdr. T. O. Self ridge. U. S. X.) engaged on the survey of the isthmus
for a ship canal. The color of this monkey was described as uniform
black excejit for the flesh-colored face and muzzle and the pale fulvous
belly; the color of the latter, however, hardly extending on the inner
surfaces of the limbs. A colored illustration for this monkey was sub-
sequently published by Sclater (Proc. Zool. Soc. London for 1872, })t.
2, p. 688, pi. 57 (col.), Nov. 1872). Elliot (A review of the l^rinuit.es,
vol. 2, p. 30, June 15, 1913), however, described this monkey as having
the "underi)arts extending a short distance on inner side of arms and
leg.s, bright rufous." and the line between the color of the underparts
and the black of the body as sharply defined.

Dt't<M-mination of the status of the name AfeJcs rufivnitrh presents
some ditiiculty at the present time. The relative lengths of head and
body to that of the .short-haired tail, the color pattern, and direction
of hair on crown and foi-ehead corresp(jnd closely with these features
in Ahtvattn paUiata ar<juatoriallH^ young females of which also lack
tlio bushy beard present on the throat in the males. These resem-
blances are shown very well by two .slightly larger young howlers
{V. 8. N. M. Nos. 3349, 3359) also from the Rio Atrato. It .seems
unlikely that representatives of both Ateles and Alouatta with the
same color i)attern would be found in the same region. Sclater states
that the thumb is absent, but examination of skins in the collection
shows that the thumb in skins of some young individuals is incon-
spicuous and might have been overlooked. The hand and feet in the
illustration published by Sclater {ojk <it.) seem more clo.sely to re-
semble those of AUmatta than those of Atrlrs. Elliot {op. c'li.) states
that no skull .s^mmus to have been preserved. While all these points
may be given consideration final generic allocation nuist await critical
examination of the type, now not available.

u. 1. «oviiii(MiNT fmNTiNa orrict: l>«4

PROCEEDINGS OF THE UNITED STATE5 NATIONAL MUSEUM

i33ueJ ff^jKyL Q*Ml hy the

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM

Vol. % Wathington: 1944 No. 3187

A EEVISIOX OF THE AMERICAN CLIXGFISHES, FAMILY
GOBIESOCIDAE, WITH DESCRIPTIONS OF NEW GEN-
ERA AND FORMS

By Leonard P. Schultz

Recently, while attempting to identify some specimens of cling-
fishes from the fresh waters of Venezuela and Colombia, South
America, it became clear to me that the American Gobiesocidae were
in a state of confusion greater than I had suspected from previous
work on the group. A search of the literature did not reveal any
attempt to strai<i:hten out the classification or nomenclature of this
family. Mew species have been described with regularity, but in
many cases the doscribers have ignored the same species long ago
named from nearly the same localities. I have made no attempt in
this revision to record all the miscellaneous references to American
Gobiesocidae, but I have included the most important contributions.
To record every reference in the literature would require one to
examine the specimens in most of the important museums of the
worM, which is not possible at the present time. However, that will
eventually iiave to be done if the identifi«ations recorded in numerous
instances are to be corrected.

It was found necessary to dissect tlic skin away from the front of
all median fins in order to count all the fin rays. Vei*y few authors
lia\t' counted tlie first one or two rays at the beginning of both anal
and doisal fins. The shoi-t stubby ray on the dorsal edge of I he
pectoral fin also is included in my counts. (See table L)

The coloration is variable and cannot be used to separate species
except to a limited extent.

47
G04148 — «4 1

48 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

In this study of the American Gobiesocidae, 9 genera and 25 species
and subspecies are recognized. By far the greatest majority of these
occur in the tropical Pacific. Although the Gobiesocidae are mostly
marine fishes, a few species have been taken in fresh-water streams
a short distance above brackish water, clinging to the stones.

Rimicola of the Pacific is represented in the Atlantic by the genus
Acyrtus^ but the genus Infratridens has no known Atlantic counter-
part. Arhaciosa has five forms in the Pacific and but one in the
Atlantic. Sicyases occurs in the Pacific only at the southern part
of South America. Cotylis has four species in the Pacific and one
in the Atlantic. Sicyogaster has an interrupted distribution, with
one species along the Pacific coast of southern South America and
another along the west coast of the United States and British Colum-
bia. Arcos has one species on each side of Central America.
Gohiesox has two species in the Atlantic, and three in the Pacific
confined to tropical waters.

Cocos Island presents a problem that needs further study. So far,
the Pacific representatives of Gohiesox cephalus, and Cotylis nigripin-
nis from the Atlantic, have been taken only on Cocos Island. They
are Gohiesox fulvus and Cotylis nigripinnis woodsi.

The records of Gohiesox adustus (Pellegrin, Bull. Mus. Paris, vol. 7,
p. 206, 1901, and Giinther, Biologia Centrali-Americana, Pisces, p. 4,
1906) in the Rio Chapalagana at Tepico in the Rio Grande de San-
tiago may be some other species. No description is given, and thus
it is not possible to place the above record with any species until the
specimens have been re-examined.

While this study was being made, Dr. S. F. Hildebrand kindly
turned over to me some notes made by Dr. W. H. Longley on types in
certain museums of Europe. Though most difficult to read, these
proved of considerable value, and it was a pleasure to note that I
came to the same conclusions that the late Dr. Longley had inde-
pendently arrived at in regard to referring certain species to the
synonymy of others. Although his notes were never published, some
of the conclusions appeared during 1933 and 1934.

The following key was prepared after examining the American
clingfishes in the collections of the United States National Museum,
as well as specimens lent by the Chicago Natural History Museum
(F. M. N. H.) through the courtesy of Dr. K. P. Schmidt and Mrs.
Marion Grey, and others lent by the University of Michigan Museum
of Zoology through the courtesy of Dr. Carl L. Hubbs. Dr. C. M.
Breder, Jr., of the American Museum of Natural History, kindly
allowed me to examine the holotype of Gohiesox yuma Nichols.

REVISION OF AMERICAN CLINGFISHES — SCHULTZ 49

KEY TO THE GENERA AND SPECIES OF AMERICAN GOBIESOCIDAE

1'/. Groove bt'twivn tip of snout iiiul upiht lip of prt'iuaxilluries extonding around
front of snout and not forming a convex curve dorsally over tip of snout;
width of middle of upper lip narrow, about the same as hiterally, and ap-
proximately equal to width of pupil; gill membranes attached opposite
tliird to fifth upper pectoral fin rays; axial flap of skin behind pectoral fin
with its upi>er edge attached at midbase of pectoral fin or below midbase;
fleshy pad on outer pectoral base present only ventrally, without a free
margin posteriorly and enlarged or swollen at lower posterior corner of
l)ectoral fin base: lower first to fifth pectoral raj's short, about half length
of longest pectoral ray, eighth and ninth much longer than lower pectoral
rays; anal rays G to 8; dorsal rays G or 7 (all rudiments counted as
one ray).
2rt. Incisorlike teeth at front of lower jaw with 4 minute points,^ these at front
of upper jaw mostly conical ; each jaw with 1 or 2 inner rows of minute
conical teeth ; axial fiap of skin behind pectoral fin attached at lower part
of pectoral fin base; anal origin a little behind a vertical line through
dorsal origin ; greatest depth of body 5V_> to 6Vj, length of head 3 to 3%.
greatest width of head 4i/j to 5, length of disk 5 to 5Vj, all in standard
length ; length of disk about equal to distance from tip of snout to front
of disk ; pectoral rays about 1!) to 21 ; color when alive green or reddish,
with or without light spots {Acyrtus, new genus) (Florida Keys and

West Indies) Acyrtus rubiginosus (Poey)

26. Incisorlike teeth at front of lower jaw with smooth tips ; middle front teeth
of upper jaw conical ; teeth in inner rows of both jaws shorter, smaller,
and conical : axial fiap of skin behind pectoral fin attached opposite middle
of pectoral base; greatest depth of body 8 or 9, length of head 3Vj to 3%,
greatest wiilth of head 5, length of disk 5Vo, all in standard length ; anal
origin a little in advance of dorsal origin ; interorbital space 3^/^ in head,
eye 1V> in interorbital space; length of disk about equal to caudal
I)eduncle; lower pectoral rays shorter, second and third from bottom about
half length of longest pectoral fin rays ; pectoral fin rays about IG or 17
(Rimicola Jordan and Evermann) (Todos Santos Bay, Baja California
to Monterey Bay and west coast of Vancouver Island, British Colum-
bia) Rimicola eigenmanni (Gilbert)

lb. Tip of snout formed by premaxillaries, which are much wider at middle of

snout than laterally, groove arched dorsally over tip of snout ; axial Hap of

skin behind p<'Ctoral fin with its upper etlge attached much above midbase of

this fin ; lower first to seventh pectoral fin rays not shortened, about as long

as eighth or ninth from bottonj.

3fl. Anterior twth of lower Jaw trlfid incisors, trifid tips usually evident, except

middle 2 or 3 sometimes worn off smooth although 1 or 2 of more laterally

placed incisors at front of lower jaw always trifid.

An. CJlll membrane attached opjiosite third to fifth jx-cloral lln rays; front

teeth of upper Jaw smooth liiiped incl.sors (sometimes fiattened-cor)l-

fonn) ; iront of both Jaws with 1 or 2 inner rows of small conical teeth

behind outer row of enlarged Incl.sorlike teeth, sometimes these inner

rows apparently represented by oidy 2 or 3 teeth ; fleshy pad on outer

base of pectoral fin with free posterior margin ending a little below

attachment of gill membraneH ; greatest width of head 3, length of head

' SoiiiPtiiiKfl the middle two tpctli arc worn down n<'nily HnirMdli. jih In tlio type of '/.
hrrtillinun Illldclirand and OlnxliiirK.

50 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 93

2% to 2%, greatest depth of body 5 to 6, length of disk 31/2, all in stand-
ard length ; length of disk much greater than distance from tip of snout
to front of disk ; distance from dorsal origin to midbase of caudal fin
contained 1% to 1% times in snout tip to dorsal origin; anal origin
under base of the third or fourth dorsal fin ray ; caudal peduncle short,
its depth about equal to its length and about 3 times in base of dorsal
fin; dorsal fin rays 11 to 13, anal 10 or 11, pectoral 18 to 21 (usually
19 or 20) {Infratridens, new genus) (Gulf of California; southern

California) Infratridens rhessodon (Rosa Smith)

4&. Gill membranes joined opposite upper edge of pectoral fin base ; incisorlike
teeth of both jaws with trifid tips, except middle pair or two sometimes
smooth-tipped ; teeth in both jaws in a single row, lateral 2 to 4 conical
and last 1 or 2 sometimes strong canines ; outer lower base of pectoral
fin with fleshy pad poorly developed and without any trace of a free
margin ; pelvic fins joined about halfway out fourth to sixth pectoral
fin rays and not near base ; dermal flap in axile of pectoral fin joins oppo-
site fourth to tenth pectoral fin ray ; width of head BV2 to 6, length of
head 2% to 5, greatest depth of body 6 to 10 (except in eos), length of
disk 4 to 6 (except in eos), all in standard length; opercular spine
not strongly developed and not reaching to rear of head {Arhaciosa
Jordan and Evermann).
5a. A pair of black spots (more or less ocellate) on back behind head over
pectorals usually distinct, each spot well separated ; dorsal surface of
back in front of dorsal origin variously barred or mottled or dark
spotted but without 3 hourglass-shaped large dark blotches. ( Species
inhabiting waters of the Pacific coast and offshore islands.)
6a. Distance from base of last dorsal ray to midcaudal fin base con-
tained l%o to 1% times in length of dorsal fin base; least depth
of caudal peduncle about 1.0 to 1% times in length of caudal
peduncle (from base of last anal ray to midcaudal fin base) ; inter-
orbital space longer than length of ^out ; dermal flap of skin
in axis of pectoral fin with its upper edge joined to pectoral fin
base opposite fifth to eleventh pectoral ray; dorsal rays 10 or 11
(usually 10), anal rays 7 to 9 (usually 8 or 9) ; pectoral fin rays
19 or 20 ; middle teeth of both jaws with trifid tips, middle denticle
usually longest on lateral teeth, worn down in adults (Gulf

of California) Arbaciosa humeralis (Gilbert)

6b. Distance from base of last dorsal ray to midcaudal fin base con-
tained 0.75 to 0.9 in length of dorsal fin base ; least depth of caudal
peduncle 1.6 to 2 times in length of caudal peduncle.
7a. Pectoral fin rays 22 to 24 (usually 23) ; dorsal rays usually 8 (7
to 9), anal 7 or 8 (usually 8) ; greatest width of head contained
Sy2, length of head 2% to 2%, in standard length ; snout a little
longer than width of interorbital space; middle teeth of both
jaws with trifid tips (Mazatlan, Mexico).

Arbaciosa eos (Jordan and Gilbert)

7b. Pectoral fin rays usually 19 to 21 (rarely 22).

8a. Middle incisorlike teeth of both jaws (at least on adults) with

smooth tips, the lateral incisors trifid ; pectoral fin rays usually

about 21 ; bony ridges on snout weakly developed.

9a. Dorsal rays 8 to 10 (usually 9) ; anal 8 or 9 (usually 8)

(Peru) Arbaciosa pyrrhocincla pyrrhocincla (Cope)

9b. Dorsal rays 8 ; anal 7 or 8 (Galapagos Islands).

Arbaciosa pyrrhocincla truncata Heller and Snodgrass

REVISIOX OF AMERICAN CLINGFISHES — SCHULTZ 51

8b. Middle Incisorlike teeth of both jaws usually trifid, seldom worn
ciCf smooth even on adults; dorsal rays 6 to 8; anal G or 7;
jiectoral 19 to 21 (usually IJ) or 20) ; bony ridges on upper
part of snout rather well developed (Ecuador to Gulf of

California) Arbaciosa rhodospila ((Jiinther)

56. Three or four larixe hourglass-shaped daric brown or blackish blotches
from in front of dorsal fin to rear of head ; a fainter one sometimes
on top of head ; side of head with four oblique bars and sides of body
with dark bars ; incisors with trifid tips; dorsal rays 7 to 9; anal 6 to
9 (rarely G or 9) ; pectoral 18 to 23 (West Indies; Guatemala to

Brazil) Arbaciosa fasciata (Peters)

36. None of the teeth with trifid tips.

10a. Middle pair of incisors on both jaws much broader and longer than

adjoining pairs ; posterolateral teeth small and conical ; rims of orbits

bony, elevated ; opercular spine strongly developed and forming

posteriormost tip of head ; valvular flap and margin of anterior nostril

with its margin finely fringed with short cirri ; gill membrane attached

at upper anterior edge of pectoral fin base; fleshy pad well developed

on outer lower surface of pectoral base, with a free membranous

edge posteriorly ending at ba.se of tenth to twelfth pectoral ray ;

shoulder girdle with a free dermal flap extending dorsally nearly to

attachment of gill membrane; anal ori^rin under base of second or

third from last dorsal fin ray; disk large, its length about equal to

head and contained about 2% to 2-i/f, in standard length ; anus just

behind rear margin of disk; origin of dorsal fin a trifle closer to tip

of opercular spine than midcaudal fin base; dorsal rays 10 or 11

(usually 11) ; anal 8 or 9 ; pectoral 24 or 25 {Sici/ascH Miiller and

Troschel ) .

llo. Dorsal origin equidistant between midcaudal fin base and upper edge

of gill opening to middle of length of upper pectoral rays; distance

from base of last dorsal ray to midcaudal fin base in upper edge of

gill opening to dorsal origin 1.80 to 2.35; length of caudal peduncle

in snout tip to anal origin 15.50 to 7.36; base of dorsal fin in head

1.90 to 2.40; ba.se of anal fin in bead 2.83 to 4.06; base of dorsal fin

In snout tip to dorsal origin 3.70 to 4.G7 ; base of anal fin in snout

tip to anal origin G.70 to 8.8.'i (Chile and I'eru).

Sicyases sanguineus Miiller and Troschel
116. Dorsal origin equidistant between iiiidcaudal fin base and middle
of postorbital length of head ; base of last dorsal ray to midcaudal
fin base In upper edge of gill opening to dorsal (trigin 1.(55; length
of caudal iH?duncle in snout tip to anal origin 5.26; base of dorsal
fln in head 1.55; base of anal fin in head 2.14; base of dorsal fin in
snout tip to dorsal origin 3.04; iiasi' of anal fln in snout tip to anal
origin 5.10 (Juan Ffrnrmdcz Island).

Sicyases hildebrandi, new species
106. Middle pair of Incisors not enlarged, all the Inci.sorllke or conical teeth
at front of both Jaws of nearly same size and length; front of lower
Jaw with small incisors In 2 or 3 pairs, with smooth tips; posterolat-
eral teeth smaller, conical, sometimes one or two a little enlarged
and almost cardnelike; ustuilly a small patch of very short conical
teeth behind outer row of larger teeth at front of jaws b\it sometimes
lacking fir reduced to 1 or 2 teeth ; rims of orbits not elevated or bony ;
anterior nostril with a dermal flap, sometimes with liifid or even

52 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

multifid tips arising on posterior rim, but nostrils not fringed witli
short cirri.
12a. Short blunt papillae on lips and around mouth generally, these in
form of short barblets, arrangement as follows : Median part of chin
and lower jaw with 2 or 3 rows of papillae, or chin anteriorly with
a pair of low lobes in form of reversed parentheses [ ) ( ] and
sometimes at their inner tips a pair of papillae (more or less fused
with anterior lobes in nigripinnis and in pinniger) ; an inner row
of barblets lateral to median lobes, one pair on each side ; lower lip
at each side of median part of chin lobelike, sometimes bearing 2
small papillae ; along inner edge of groove of lower jaw are 2 or 3
large papillae or knobs on each side; upper lip with a median
papilla or knob and 5 more on each side ; front edge of snout above
groove without papillae but laterally 3 to 5 knobs or papillae
present or absent; sometimes another papilla occurring behind
rictus and still another below rictus; gill membranes joined oppo-
site fifth to seventh upper rays of pectoral fin ; fleshy pad on outer
base of pectoral fin with a free posterior membranous margin
extending dor.sally to opposite attachment of gill membranes ; dorsal
rays 10 to 19 ; pectoral fin rays 21 to 27 ; anus closer to anal origin
than to rear margin of disk {Cotylis Miiller and Troschel).
13a. Dorsal rays fewer than 15, counting all rudiments.
14a. Dorsal rays 13 or 14; anal 10; upper lip with papillae.

15a. Pectoral fin rays 22; papillae around mouth short and knob-
like (Gulf of California) Cotylis papillifer (Gilbert)

15&. Pectoral fin rays 24 to 27; papillae around moiith more numer-
ous, better developed, barbellike; lobe of lower lip next to
middle of chin with two barbels (pi. 1, A) (Panama Bay to

Ecuador and northern Peru) Cotylis microspilus (Fowler)

14&. Dorsal rays 10 to 12 ; anal 8 to 10 ; papillae on upper lip, lobelike ;
lobe of lower lip next to middle of chin without barbels (pi. 1,
B) ; dorsal origin equidistant between midbase of caudal fin
and middle of postorbital length of head to equidistant be-
tween midcaudal base and upper base of pectoral fin ; color
pattern variable ; median fins mottled, barred, or blackish with
tips of rays white.
16a. Depth 41/2 to 61/2 ; eye 3.1 to 3.6 in length of base of dorsal fin ;
dorsal rays usually 11, anal usually 9, pectoral 22 to 26
(Maryland to West Indies to Brazil).

Cotylis nigripinnis nigripinnis Peters
166. Depth about 6% or 7; eye 2.8 in length of dorsal fin base;
dorsal rays 10, anal 8, pectoral 22 (Cocos Island).

Cotylis nigripinnis woodsi, new subspecies

13&. Dorsal rays 17 to 19 (counting all rudiments) ; anal 10; anal

origin under base of ninth or tenth dorsal fin ray or under middle

of base of dorsal fin ; origin of dorsal fin a little closer to tip of

snout than midbase of caudal fin ; papillae on upper lip knoblike

(Gulf of California) Cotylis pinniger (Gilbert)

12&. No papilla on upper lip, lobelike structures occurring around lips of
lower jaw when best developed being low knobs or ridges, chin
lacking inner series of papillae as described for Cotylis.
17a. Gill membranes joined at upper edge of pectoral fin base, some-
times a little anteriorly, giving appearance of being opposite

REVISION' OF AMERICAN CLINGFISIIES — SCIIULTZ 53

bases of uppor flrst to third pectoral fin rnys or the orbits larper

than iiiterorbifal space; liioisorlike teeth at front of lower jaw

projeotinp forward in a nearly horizontal or oblique direction,

middle pair a little larger than those laterally.

18o. Anal rays 10 to 14; dorsal 12 to 16, pectoral 10 to 23 (counting

all rudiments) ; fleshy pad on outer margin of pectoral fin

base very well developed and free membranous border along

its posterior edge extending up to or beyond twelfth pectoral

ray from dorsal edge; interorbital space equal to or wider than

eye; least depth of caudal peduncle 4^^ to 5Vi times in dorsal

origin to midcaudal tin base ; anal origin under anterior third of

dorsal fln base {Sicyof/ai^tcr Brisout de Barneville).

19a. Anal rays 10 or 11 ; dorsal 12 or 13 ; free margin of fleshy pad

on pectoral fin base ending abruptly opposite ninth to twelfth

ray from upper edge of pectoral fin base; eye 1 to l^^ In

interorbital space and 4 or 5 in head ; anus a little closer

to i-ear margin of disk than to anus; origin of dors&l fln

equidistant between midbase of caudal fin and anterior half

of postorbltal length of head (rem and Chile).

Sicyog-aster marmoratus (Jenyns)
196. Anal rays 12 to 14 ; dorsal 13 to 16 ; fleshy pad on outer base
of pectoral fin with free posterior margin ending gradually,
about opposite flrst to third upper pectoral fin ray ; eye 1%
to 2 in interorbital space; length of disk about 3, head about
214. greatest depth 4% to 5, all in standard length ; anus
much closer to anal origin than rear margin of disk; origin
of dorsal equidistant between midbase of caudal fln and
rear of head; interorbital space about equal to snout (San
Diego to Queen Charlotte Islands, British Columbia ; I'uget

Sound) Sicyogaster maeandrlcus (Girard)

186. Anal rays 7 or 8; dorsal 7 to 9; pectoral 22 to 25; diameter of
eyes greater than interorbital space, the latter about % to %
in eye; color usually reddish when alive (Arco/<, new genus).
20a. Free margin of fleshy pad on pectoral fln base ending op-
posite thirteenth ray from uj^per edge of pectoral fin base;
Interorltital space G or 7 in head; least depth of caudal
peduncle 3Vi' times in distance from midcaudal fin base to
dcrsal origin and 1 '/•• in Its length; iielvi<-s fastened nearer
base of r)ectoral rays than one-third way out (Oalapii^os
Islands; Panama Bay; and MazatlAn, Gulf of California).

Arcos poecilophthalmus (Jenyns)

20^. Free margin of fiesby pad on pectoral fin liase ending opi)oslte

sixteenth to nineteenth r;iy from upiK-r edge of pectoral fln

base; interorbital ~> or <i limes In bead; least deptii of caudal

peduncle 4 times in flistance from midcaudal base to dorsal

origin and r4ft In its b-ngth; pelvics fastened about on<'-thlrd

way out lower i)ectoral rays (Bahama Islands; West Indies).

Arcos macrophthalmua (G(lnther)

176. 0111 membranes Joined opposite third to seventh upp<'r pectoral fln

rays .somewhat more anteriorly than In Cotyliit; liiclsorllke teeth

at front of lower Jaw not prnj»H'tlng horizontally forward but

curved oblifpiely upward so as to nearly opjio.se those In upper

Jaw, the pair of incisors at middle of lower Jaw nearly same size

as a«lJoinlng ones; outer surface of p<'<Moral fln base with a dls-

54 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

tinctly fleshy pad, posterior margin free and joined opposite at-
tachment of gill membranes {Gobiesox Lacepede).
21ffi. Disk much greater than distance from tip of chin to front of disk.
22a. Origin of dorsal fin equidistant between midcaudal fin base and
rear one-third of pectoral fin rays or a little behind them ; anal
origin under fifth dorsal fin ray, behind middle of base of rays
of that fin; teeth of lower jaw not projecting forward in a
nearly horizontal position but directed nearly straight up-
ward in adults, a little more oblique in young specimens;
head 2.2 to 2.7, disk 2.6 to 3.3, and depth 4 to 5.5, all in
standard length ; dorsal rays 8 or 9, anal 5 to 7, i)ectoral 18
to 21 ; anus equidistant between anal origin and rear margin
of disk or a little nearer to anal origin ; eye 1% (young) to 5
(adults) times in interorbital space.
23a. Length of disk when measured from its rear margin reaches
nearly to end of anal fin usually from midbase to base of
last anal ray ; small dark spot often present near front of
base of dorsal fin (Costa Rica, West Indies, to Brazil).

Gobiesox cephalus Lacepede

23&. Length of disk when measured from its rear margin reaches

only to base of first or second anal fin ray ; front of dorsal

with a large dark blotch not at base of fin (Cocos Island).

Gobiesox fulvus Meek

226. Dorsal origin equidistant between midcaudal fin base and upper

base of pectoral fin or rear of head ; dorsal rays 10 or 11, anal

8, pectoral 19 to 21 ; gill membrane attached opposite fourth

to seventh upper pectoral fin rays ; interorbital 3% to 4i/^ in

head ; distance from base of last dorsal ray to midcaudal base

2% to Sli times in distance from dorsal origin to rear of head ;

anus equidistant between or closer to anal origin than rear

margin of disk ; fleshy pad on pectoral fin base with posterior

margin free all way up to attachment of gill membrane ; anal

origin under fifth dorsal ray ; depth of caudal peduncle equals

its length.

24a. Eye IV2 to 2 in interorbital space (Texas; British Honduras;

Bahamas and West Indies). Gobiesox punctulatus (Poey)

246. Eye 0.9 to 1.1 in interorbital space (Pacific-Mazatlan).

Gobiesox adustus Jordan and Gilbert
216. Disk about equal to distance from tip of chin to front of disk;
pelvic fins attached about one-third way out pectoral fin rays;
length of disk equal to distance from rear margin of disk to
anus or 1% times from disk to anal origin; head 2.9, disk 3.8
to 4.2, depth 5 or 6, width of head 3%, all in standard length ;
eye 1% to 2 in interorbital space; interorbital 3^ and disk
1% in head ; dorsal origin equidistant from midcaudal fin base
and middle of length of pectoral fin ; distance from last dorsal
ray to midcaudal fin base 4 times in distance from rear of head
to dorsal origin ; anus closer to anal origin than rear margin
of disk ; least depth of caudal peduncle greater than length of
caudal peduncle from base of last anal ray to midcaudal fin
base; opercular spine not well developed; anal origin under
base of about the sixth dorsal ray ; dorsal rays 11 or 12 ; anal
7 or 8; pectoral 18 to 20 (Gulf of California).

Gobiesox funebris Gilbert

rev:

K 1

;

S 1

: ; ;

M ^ I

;

; ;

^ 1

a> C4

^ «

;

eo •

N 1

*" :

•O 1

M lO 1

^ <o

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;

CO c» ««< ■>!• e>«

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- :

00 ca r^ w lO CO

;

e» ;

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00 ■*

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s ;

00 1

a 1

S " :

00 lO ;

'* 1

<«•

00 1

•" ;

"^ ;

t^ 1

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2 1

ro 1

Pectoral fln rays abovo
attachment of gill
membranes

r« 1

CO m

e 1

•H eo r^ c«

N »< e«

•« 1

«< M CO

•4< to C4

•*

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ei -H

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004148 — 44-

56 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

ACYRTUS, new genus

Genotype. — Sicyases ntbiginosus Poey.

This new genus is characterized by the lack of a dorsal curve of the
premaxillary groove along upper lip over the snout tip in conjunction
with the incisorlike teeth of the lower jaw having four minute joints.
It may be distinguished from all other genera of American Gobieso-
cidae by the key on page 49. Other characters are those of the
genotype.

Named Acyrtus in reference to the absence of the dorsal curve of the
premaxillary groove over front of snout so common in all other Ameri-
can clingfishes except Bimicola, to which this new genus is most closely
related.

ACYRTUS RUBIGINOSUS (Poey)

Sicyases rubiginosus Poet, Synopsis piscium Cubensium, p. 391, 1868 (Palmasola,

Cuba); Enumeratio piscium Cubensium, p. 124, 1876 (Cuba). — Jordan,

EvEBMANN, and Clabk, Rep. U. S. Comm.. Fish, for 1928, pt. 2, p. 490, 1930

(Matanzas, Cuba).
Sicyases carneus Poey, Synopsis piscium Cubensium, p. 392, 1868 (Palmasola,

Cuba) ; Enumeratio piscium Cubensium, p. 124, 1876 (Cuba). — Joed an,

EvERMANN, and Clark, Rep U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930

(Matanzas, Cuba).
Sicyases carneus Voey—S. rubiginosus Poey, Longley, Carnegie Inst. Washington

Year Book, No. 33, p. 271, 1934.
fArbaciosa sp. Beebe and Tee-Van, Zoologica, vol. 10, No. 1, p. 252, 1928 (Lamentin

Reef, Port-au-Prince Bay, Haiti).
Gobiesox rubiginosus Jordan and Evermann, Rep. U. S. Comm. Fish and Fish.

for 1895, App., p. 492, 1896 (Matanzas, Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3,

p. 2337, 1898 (Cuba).
Oobiesox carneus Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for

1895, App., p. 492, 1896 (Matanzas, Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3, p.

2337, 1898 (Matanzas).
Gobiesox (Rimicola) beryllimis Hildeibrand and Ginsbttbg, Bull. U. S. Bur. Fish.,

vol. 42, p. 213, fig. 5, 1927 (Key West, Boca Chica, Fla. ) .
Rimicola beryllinus Jordan, Evermann, and Clark, Rep. U. S. Comm. F^sh. for

1928, pt. 2, p. 490, 1930 (Key West).

Remarks. — An examination of the type of hei^yllinus indicates that
the incisorlike teeth at the front of the lower jaw have four minute
points, although the middle ones are worn down a little. The teeth
along with other characters in the description of heryUmus are in need
of rechecking, but the type appears to have been dried out sometime
and is not in first-class condition. Hildebrand and Ginsburg's draw-
ing, "figure 5," has the lower rays of the pectoral fin twice too long and
the disk is not quite long enough. The fin rays were not correctly
counted in the original description.

Material examined.— Cvba : U.S.N.M. Nos. 82581 and 82582, totaling
12 specimens. Florida : U.S.N.M. Nos. 87533 (holotype of beryllimis)
and 116936, one specimen.

Range. — Florida Keys and West Indies. ,

REVISION OF AMERICAN CLINGFISHES — SCIIULTZ 57

Genus RIMICOLA Jordan and Evermann

Rirnirola Jokuan and Evkkx[an.n, in Jordan, Pr«)<-. California Acad. Sci., vol. 6,
p. 231, 1S96. (Genotype: Oobiesox muscarttm Moek and Plerson [ = Oobie80X
eigenmanni Gilbert].)

RIMICOLA EIGENMANNI (Gilbert)

Gobicsox eigenmanni GiLBtJiT. Proc. U. S. Nat. Mus., vol. 13, p. 90, 1890 (Point
Loma, near San Diego, Calif.).

Oobiesox rhcssodon Rosa Smith, Proc. U. S. Nat. Mus., vol. 7, p. 553, 1885 (San
Cri.stubal. P.aja California t.

liimicola cifjenmanni Jokdan. Proc. California Acad. Sci., vol. 6, p. 231, pi. 32,
1S9C. — Jordan and Evebmann, Rep. U. S. Conuu. Fish and Fish, for 1895, App.,
p. 4rt2, 189G (Point Loina) : U. S. Nat. Uus. Bull. 47, pt. 3, p. 2339, 1898
(Point Loma, San Cri.«^tObal Bay).— Snyder, Proc. U. S. Nat. Mus., vol. 35,
p. 183, 1908 (Todos Santos Bay, Baja California, northward to Pacific
Grove, Calif.). — Jordan, Evekmann. and O.ark, Rep. U. S. Comm. Fish, for
1928. pt. 2, p. 490, 1930 (Point Loma to Monterey).— Wiijjt, Copeia, 1936,
No. 2, p. IIG (Round Island Flats, Clayoquot Sound, British Columbia).

Rimicola muxraruyn Jordan, Proc. California Acad. Sci., vol. 6, p. 231, 1896. —
Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for ISOn, App.,
p. 492, 1896 (Monterey Bay) ; U. S. Nat. Mas. Bull. 47, pt. 3. p. 2338, 1898
(Monterey Bay).

Oobiesox muscantm Meek and Pierson, Proc. California Acad. Sci., vol. 5, p. 571,
pi. 71, 1895 (Monterey Bay, Calif.).

Remarks. — My counts appear to disagree with those made by
Snyder, Wilby, and other authors because I have inchided all the
rudimentary fin rays at the beginning of each fin, apparently not
counted previously. To be certain of my fin ray counts, the skin
was di.s.sected away from one side of each fin at the base of the an-
terior fin rays. This revealed usually one or sometijnes two rudi-
mentar}' fin rays, and thus the counts by Snyder and by Wilby should
be increa.sed by one or two to bring them into line with my counts.

Material examined. — British Columiua (Vancouver Island) :
U.S.X.M. No. 53800. California: U.S.N.M. Nos. 44372 (holotype
of riqrnmnnni), 48875 (cotype of rnvsranivi) ., 49570, and G1055;
U.M..M.Z. Xos. 64260, 64261, 642(52, 6364S, 63646, 63647, totaling 17
specimens.

Range. — Todo.s Santos Bay, Baja Califomia, to jMonterey Bay, and
west coast of Vancouver Ishind.

INFRATRIDENS, new genus

Genotype. — Oobiesox rliessodon Rosa Smith.

This new genus may be recognized from all otjier genera of Gobie-
.socidae by the characters descriFn'd in the key on |)ag{'s 40-.50. It is
di.stinguislu'd by its trifid incisorlikc lc(>th at front of lower jaw, the
smooth-tipped teeth in upper jaw, and the convex premaxillury groove
across front of snout. Otber characters are those of the genotype.

58 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Named hifratridens in reference to the trifid teeth at front of
lower jaw.

INFRATRIDENS RHESSODON (Rosa Smith)

GoMesox rhessodon Rosa Smith, in Jordan and Gilbert, Proc. U. S. Nat. Mus.,
vol. 4, p. 63, 1881 (Point Loma) (nomen nudum). — Rosa Smith, Proc. U. S.
Nat. Mus., vol. 4, p. 140, 1881 (San Diego, Calif.).

Arbaciosa rhessodon Jordan, Proc. California Acad. Sci., vol. 6, p. 230, pi. 36,
1896. — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 1895,
App., p. 492, 1896 (San Diego; Gulf of California) ; U. S. Nat. Mus. Bull.
47, pt. 3, p. 2340, 1898 (San Diego; Gulf of California).— Jordan, Evermann,
and Clark, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930 (San Diego;
Gulf of California).

Material examined. — Caufornia: U.S.N.M. Nos. 5246, 28396 (3
types of rhessodon), 34765, 41975, 49574, 67312, 104193, 117642, total-
ing 24 specnnens; U.M.M.Z. Nos. 63650, 63653, 63651, 63649, 63652,
totaling 69 specimens. Catalina Island: U.S.N.M. No. 121964 and
U.M.M.Z. No. 64263, totaling 8 specimens. Baja California:
U.S.N.M. Nos. 36948 and 79149, totaling 2 specimens.

Range. — Southern California to Baja California.

Genus ARBACIOSA Jordan and Evermann

Artaciosa Jordan and Evermann, in Jordan, Proc. California Acad. Sci., vol. 6,
p. 230, 1896. (Genotype: OoUesox JiumeraUs Gilbert.)

arbaciosa HUMERALIS (Gilbert)

Oobiesox humeralis Gilbert, Proc. U. S. Nat. Mus.. vol. 13, p. 95, 1890 (Puerto
Refugio, Angel Island). — Pellegrin, Bull. Mus. Hist. Nat. Paris, vol. 7,
p. 162, 1901 (Gulf of California).

Arbaciosa humeralis Jordan, Proc. California Acad. Sci., vol. 6, p. 230, pi. 35,
1896. — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for 1895,
App., p. 491, 1896 (Gulf of California) ; U. S. Nat. Mus. Bull. 47, pt. 3, p.
2341, 1898 (Angel Island; La Paz). — Jordan, E\t:rmann, and Clark, Rep.
U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930 (Gulf of California).

Material examined. — Gulf of California : U.S.N.M. Nos. 44374
(cotype of humeralis), 125008 (cotype of humeralis), 46693, 48259
(4 cotypes of humeralis), totaling 8 specimens; F.M.N.H. No. 3336,
44 specimens ; U.M.M.Z. No. 136128, 1 specimen.

Range. — Gulf of California.

arbaciosa EOS (Jordan and Gilbert)

Oobiesox eos Jordan and Gilbert, Proc. U. S. Nat. Mus., vol. 4, p. 360, 1882
(Mazatlan).

Arbaciosa eos Jordan, Proc. California Acad. Sci., vol. 6, p. 230, pi. 37, 1896
(Mazatlan). — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for
1895, App., p. 491, 1896 (Pacific coast of Mexico) ; U. S. Nat. Mus. Bull. 47,
pt. 3, p. 2343, 1898 (Mazatlan). — Jordan, Evermann, and Clark, Rep. U. S.
Comm. Fish, for 1928, pt. 2, p. 491, 1930 (Pacific coast of Mexico).

Material examined.— M-exico (Mazatlan) : U.S.N.M. No. 30889 (18
cotypes of Gohiesox eos), C. H. Gilbert.

REVISION OF AMERICAN CLINGFISHES — SCHULTZ 59

Range. — Mazatlan, Mexico.

ARBACIOSA PYRRHOCINCLA I'YUKHOCINCLA (Cope)

Sicyases pyrrhorincUis Cope, Proc. Anier. Pliilos. Soc, vol. 17, p. 43, 1S77 (Peru).
Arhaciosa pitrrhocinchis Abbott, Proc. Acad. Nat. Sci. Philadelphia, 1S99, p. 363

(Peru. Pwasmayu Bay ?). — E\T':i£MAnn and R.vdcuite, U. S. Nat. Miis.

Bull. 95, p. 155, 1917.
Arhaciosa hicroglnphka Evp^imanx and Raucliffe, U. S. Nat. Mus. Bull. 9.', p. 155,

pi. 14, fig. L', 1917 (Ldbo.s de Afuera, Porn).
? Arbariofia petcrsii (nan Garniau) AiiBorr, Proc. Acad. Nat. Sci. Philadelphia,

1S99, p. 363 (Peru) [not based on any .spociuicu].
? Oobicsox zebra (non Jordan and Gilbert) Rkoan, Ann. Mag. Nat. Hist., ser. 8,

vol. 12, p. 280, 1933 (Lobos de Tierra, Peru).

Material examined. — Peru (Lobos de Afuera Islands) : U.S.N.M.
Nos. 77561 (type of hierogly phica) ^ 77565 (10 cotypes of hieroglyph-
ica), 101703, 101704, 101705, 12-8175, totaling 21 spcciineii.s. Also from
Peru : U.S.N.M. Nos. 88817, 88827, 88828, 119753, 128174, totaling 19
specimens.

Range. — Peru.

ARBACIOSA PYRRHOCINCLA TRUNCATA Heller and Snodgrass

Arhaciosa trunrata IIkixitk and Snodcr-vss, Proc. Washington Acad. Sci., vol. 5, p.
210, pi. 14, liX)3 (Tagu.s Cove, Albemarle Island, Galdpagos). — Kendall and
lUDCLUFE, Mem. Mu.s. Comp. Zool., vol. 35, No. 3, p. 160, 1912 (Chatham
Island).— IlKKUK, Publ. Field Mus. Nat. Hist., zool. ser., vol. 21, p. 391, 1936
(South Seymour Island ; Eden Island).

Oohieaox zebra (non Jordan and Gilbert) Gilbkrt, Proc. U. S. Nat. Mus., vol. 13, p.
452, 1890 (Duncan Island, Gal.lpagos).

Material eu^amined. — Galatagos Islands: U.S.N.M. Nos. 65427,
101702, 101710, 101715, 101716, 101718, 109419, 116202, 119337, totaling
36 specimens; F.M.N.H. No.s. 25105-25215, 41301-41303, 41214, 41215,
41641^1646, totaling 62 specimens.

Range. — Galapagos Islands.

ARBACIOSA RHODOSriLA (Gunther)

Oobicsox rhoflosj)ilus G: nthkk, Proc. Zool. Soc. London, vol. 0, p. 25, 1864 (Pan-
ama) ; Trans. Zool. Soc. lyindon, vol. 6, pp. 300. 445, 1S09 (Panama). — Jordan
and KVKBMANN, Rep. U. S. Comm. Fish and Fish, for 18JK>, App., p. 492, 1896
(Panama) ; U. 8. Nat. Mus. Bull. 47, pt. 3. p. 2335, 1898 (Panama).— Bou-
i,KN(iKB. Bol. Mus. Zool. Anat. (."omp. Iniv, Torino, vol. 11, No. 335, p. 8, 1S99
(Santa Kl«'na Bay, Iv-uador).— GiiJjarr and Stauks, Mem. California Acad.
Sci., vol. 4, p. 1S9, 1904 (Panan)a).- Mkkk and IIildkiika.m), The marinr
flslicH of Panama, pt. 3. p. 926, 192S (Panama and Santa EIrna Bay). — Joudan,
EvKUMA.NN, and Cr.AiiK, Hep. U. S. Comm. Fish, for 192M. j.t. 2, p. 489, 1930
(i'anama).

Sicyases petersii Garman, Proc. Boston Soc. Nnt. Hist., vol. 18, p. 203, 1875 (San
Joa6, San .Miffjifl, and Sabnga. all Pj-arl Island.s, Panama Bay).

Oobicsox zebra Jokhan and Ghhfst, I'ror. U. S. Nnt. Mus., vol. 4, p. 359, 1882
(Mnzatlfin).

60 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Arhaciosa sehra Jordan and Eveemann, Rep. U. S. Comm. Fish and Fish, for 1895,
App., p. 403, 1896 (Mazatlan) ; U. S. Nat. Mus, Bull. 47, pt. 3, p. 2341, 1898,
(MazatMn). — Mebsc and Hildebkand, Marine fishes of Panama, pt. 3, p. 927,
1928 (Toboguilla Island). — Kendall and Radcliffe, Mem. Mus. Comp. Zool.,
vol. 35, No. 3, p. 160, 1912 (Toboguilla Island). — Jordan, Evermann, and
Clabk, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 491, 1930 (MazatlSn.)

Material examined. — Gui:f op CAiiiFORNiA : U.S.N.M. Nos. 29250 (47
cotypes of seSra), 47496, 119716, 119717, 119754, totaling 56 specimens.
Costa Eica: U.S.N.M. Nos. 92119 and 101711, totaling 3 specimens.
Panama Bay: U.S.N.M. Nos. 65428 and 120435 (3 cotypes of petersii),
totaling 8 specimens. Colombia : U.S.N.M. No. 101712, one specimen.
Ecuador : U.S.N.M. Nos. 88826 and 101717, totaling 3 specimens.

Range. — Gulf of California to Ecuador.

ARBACIOSA FASCIATA (Peters)

Sicyases fasciatus Peters, Monatsb. Akad. Wiss. Berlin, 1859, p. 412, May 8, 1860
(Puerto Cabello [probably Venezuela]). — Gunthek, Catalogue of the fishes
in the British Museum, vol. 3, p. 497, 1861 (Puerto Cabello) ; Trans. Zool.
Soc. London, vol. 6, p. 390, 1869 (Puerto Cabello). — Guitel, Arch. Zo-ol. Exper.,
vol. 5, No. 5, pp. 645-652, 1906 (anatomy). — Jordan, Evermann, and Clark,
Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 490, 1930 (Puerto Cabello).

Oohiesow rupestris Poey, Memorias sobre la historia natural de la isla de Cuba,
vol. 2, p. 283, pi. 18, fig. 6, July 1860.

Oobiesox fasciatus Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for
1895, App., p. 492, 1896 (Puerto Cabello) ; U. S. Nat. Mus. Bull. 47, pt. 3, p.
2338, 1898 (Puerto Cabello).

Arhaciosa rupestris Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for
1895, App., p. 492, 1896 (Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2341, 1898
(Cuba). — Beebe and Tee- Van, Zoologica, vol. 10, No. 1, p. 252, fig., 1928
(Port-au-Prince Bay, Haiti). — Jordan, Evermann, and Clark, Rep. U. S.
Comm. Fish, for 1928, pt. 2, p. 491, 1930 ( Cuba ) .—Parr, Bull. Bingham
Oceanogr. Coll., vol. 3, art. 4, p. 136, 1930 (Green Cay, Bahamas).

Sicyases rupestris Poey, Synopsis piscium Cubensium, p. 391, 1868 (Cuba) ;
Enumeratio piscium Cubensium, pt. 2, p. 124, 1876 (Cuba).

Arhaciosa minuta Meek and Hxldebrand, The marine fishes of Panama, pt. 3,
p. 928, pi. 92, 1928 (Colon, Panama).

Material examined. — St. Croix Island: U.S.N.M. Nos. 15382 and
15431, totaling 13 specimens. Barbados : U.S.N.M. No. 86752, 1 speci-
men. Cuba : U.S.N.M. Nos. 37414, 37421, 82580, totaling 15 specimens.
Guatemala: U.M.M.Z. Field No. H35-138a, 6 specimens. Panama:
U.S.N.M. No. 81523 (type of minuta). Brazil: U.S.N.M. Nos. 87799,
87800-87803, 88042, totaling 19 specimens.

Ra/nge. — ^West Indies ; Guatemala to Brazil.

Genus SICYASES Muller and Troschel

Sicyases Muxlee and Teosohel, in Miiller, Arch, fiir Naturg. (Wiegmaun), 9th
year, vol. 1, pp. 297, 298, 1843 (genotype: Sicyases sanguineus Miiller and
Troschel) ; Ber. Verb, preuss. Akad. Wiss., 1843, p. 212 (genotype: Sicyases
sanguineus Miiller and Troschel) (ref. copied) ; Horae ichthyologicae. Be-
schreibung und abbildung neuer Fische, pt. 3, p. 19, 1849.

REVISION OF AMERICAN' CLINGFISHES — SCHULTZ 61

Tomicodon Brisout de Barnb\jlle, Rpv. Zool. Soc. Cuv., vol. 9, p. 144, 1846 {Tomi-
codun chilen^fis Brisout de Barueville) ; Echo Monde Savant, vol. 13, p. 535,
184G.

SICYASES SANGUINEUS MUIIer and Troschel

Sicya^cs sanguineus Mi;ixt3 and Troschfx, in Miiller, Arch, filr Naturg. (Wieg-
mann), 9th year. vol. 1, p. 298, 1843 (Chile) ; Horae ichthyologlcae, pt. 3, p. 19,
pi. 3, fig. 1, 1849 (Chile). — e}i3NTHKK, Catalogue of the fishes in the British
Mu.seuni, vol. 3, p. 404, ISOl (Chile: Valparaiso). — Delfin, CatAlogo de los
peces de Chile, Valparaiso, p. 90, llMll (Bahia de Concepci6n ; Cavancha;
Isla de Juan Fcrnaiulez ; Toni6; Talcahuaiio).

Totnicodon chilcnxis BitisouT de Bauneaille, Rev. Zool. Soc. Cuv., vol. 0, p. 144,
1846 (Valparaiso).

? Oobicsox brcvirostris Gay, Ilistoria flsica y polilica de Chile . . . Zoologia,
vol. 2, p. 335, pi. 9, fig. 1, 1848 (ref. copied).

Gobiesox sanguineus Abbott, Proc. Acad. Nat. Sci. Philadelphia, 1899, p. 363,
(coasts of Peru and Chile). — E\-ebuann and Radci.iffe, U. S. Nat. Mus.
Bull. 95. p. 153. 1917 (Peru and Chile).

Sicj/ases chilensis GI'-nther, CataloRuo of the fishes in the British Museum, vol.
3, p. 497. ISGl (Valpar.ii.xo). — Delfin. CatAlogo de los peces de Chile, Val-
paraiso, p. 90, 1901 (Valparaiso).

Material examined.— Perv: U.S.N.M. Nos. 44130, 77512, 83029,
91557. totalino: 11 .specimens. Chile (Valparaiso) : U.S.N.M. No.
121950 and F.M.N.H. No. 32994, 1 specimen each number.

Raiifje. — Peru and Chile.

SICYASES HILDEBRANDI. new species

Holotype. — U.S.N.M. No. 88818, the only known specimen, 69 mm.
in stan(hinl lenfrth, collected by Dr. W. L. Schmitt at Cumberland Bay,
Juan Fernandez Island, off Chile, December 1920.

Description. — Certain measurements were made on the holotj'pe,
and these alon^r with others made on three specimens of S. sanguineus
are recorded in table 2.

The following counts were made on the holotype: Dorsal rays 11;
anal rays 9; pectoral rays 25-25; free edge of pectoral pad ends oppo-
site 11-11 pectoral raj's counting down from the dorsal edge; gill
membranes attached opposite upper edge of pectoral fin base.

Head about 31/0, depth 0.9, di.sk 2.9, all in .standard length; eye V^/o
in intt'rorbital space, the latter 2% in head (to upper edge of gill
opening); dorsal origin equidistant between niidcaudal fin base and
middle of postorbital length of head; anal origin under bases of
second and third from last dorsal (in rays; tip of snout to tip of
opercular spine equal to distance from upper edge of gill opening to
dorsal origin; base of last dor.sal ray to niidcaudal fin ba.se V/^ in
upper edge gill opening to dorsal origin; ba.se of dorsal fin l"/,o and
l>ase of anal fin 2.1, lK)th in gill oj)eMing to dor.sal origin; ;iniis just
hcliincl (lie icar niai-;:iii of the disk: lice iioslcrior miiiiriii of the

PROCEEDINGS OF THE NATIONAL MUSEUM

fleshy pad on lower pectoral base ending opposite the eleventh ray
from dorsal edge of pectoral fin; the middle incisors of both jaws
much wider than the adjoining pair and those on lower jaw longer,
but the two middle pairs of upper jaw about the same length; the
interorbital space slightly concave ; both nostrils close in front of eye,
the anterior one with a small fringed flap arising on its posterior edge.

Table 2. — Measurements oj the two species of Sicyases, in hundredths of the

standard length

hildebrandi]

Characters

Holotype

U.S.N.M. Nos.

77512 77512 77512 83029

Standard length (in millimeters)

Length of head to -upper edge of gill opening

Length of head to tip of opercular spine

Greatest depth of body

Greatest width of head

Least depth of caudal peduncle

Length of caudal peduncle

Length of snout

Diameter of eye

Width of bony interorbital

Postorbital length of head to upper edge of gill open-

Postorbital length of head to tip of opercular spine..

Length of disk

Distance from base of last dorsal ray to midcaudal

base

Disk to anus

Anus to anal origin

Snout tip to dorsal origin

Snout tip to anal origin -.

Snout tip to center of anus.

Length of base of dorsal fin

Length of base of anal fin

Longest ray of dorsal fin

Longest ray of anal fin —

Longest ray of caudal fin

Longest ray of pectoral fin

Tip of snout to disk

Origin of dorsal to upper edge of gill opening

28.3

33.3

14.5

27.5

7.82
13.8
10.6

7.54
12.2

14.9
19.1
35.6

21.7
3.91
19.6
59.4
72.5
52.6
19.5
14.2
12.5
11.7
24.5
14.5
13.3
36.0

32.2

35.6

14.9

29.1

8.30
13.9
11.7

7.63
12.2

16.4
20.2
36.4

20.7
3.90
23.4
62.7
76.3
53.2
17.0
11.4
12.2
11.2
23.6
15.3
14.2
37.3

78.5
33.8
37.6
15.9
29.9

7.77
10.8
12.2

8.02
14.0

16.7
21.9

38.8

17.5
4.20
20.4
66.2
79.7
58.1
15.5
9.30
12.5
11.5

14.3
14.3
37.6

35.3

38.3

16.1

33.9

8.33
11.1
13.1

7.56
16.0

17.2
23.3
40.6

19.1
3.67
21.6
68.4
80.0
57.2
14.7
10.0
13.3
11.7
23.5
16.7
15.5
38.6

159
37.1

44.3
21.7
37.1

7.86
13.8
13.8

6.2S
18.2

20.4
28.0
45.9

16
0.94
20.4
69.2
80.5
59.2
17.9
9.12
14.1

22.0
18.2
15.4
37.7

Remarks. — The chief differences between this new species and
Sicyases sanguineus are in the more anterior position of the dorsal fin
and the length of the bases of the dorsal and anal fins. The following
measurements indicate the amount of the above differences, first for
the new species then for sanguineus : Distance from base of last dorsal
ray to midcaudal fin base in upper edge of gill opening to dorsal origin
1.65 and 1.80-2.35; length of caudal peduncle in tip of snout to anal
origin 5.26 and 5.50-T.36 ; base of dorsal fin in head 1.55 and 1.90-2.40;
base of anal fin in head 2.14 and 2.8'3^.06 ; base of dorsal fin in snout to

REVISION OF AMERICAN CLIXGFISHES — SCHULTZ G3

dorsal origin 3.04 and 3.70^.67; ba>e of anal fin in snout to :inal orij^in
5.10 and C.70-8.tv3 ; dorsal origin equidistant between niidcaudal fin base
and middle of postorbital length of head for hildebrandi, and upper
edge of gill ojiening to middle of length of upper pectoral rays for
sanguhu'us. In general, the new species appears to be a little slenderer
than sang^uineus. Presumably, when an adequate series of hildebrandi
is collecteil from the Juan Fernandez Islands and studied, this new
species may be best treated as a subspecies of sanguineus.

Named hildebrandi for Dr. Samuel F. Hildebrand, senior ichthy-
ologist, United States Fish and "Wildlife Service, who while working
up a monograph of the fishes of Peru noticed this new fish and sug-
gested that I describe it. It is with great pleasure that I name this
new species in his honor and in recognition of his numerous and
valuable contributions in ichthyologj'.

Genus COTYLIS MuUer and Troschel

Cotylis MCllkk and Troscuel, in Miiller, Arch, fiir Naturg. ( Wiegmann) , 9tb year,
vol. 1, p. 297, 1S43 (genotype: Cotylis nnda Miiller and Troschel=Lcpado-
gaKternudux Bl<»ch and Schneider=Go?>/r.«(oj?/7.i/n/iH.s' Jordan and Evermann =
Gobiesox nigripinnis Peters) {Cycloptcrus nudus Linnaeus not identified) ;
Horae iehthyalogicae, pt. 3, p. IT, pi. 3, lig. 2, 1840 (genotype: Cotylis nuda
Bloch and Schneider). [Cotylis (non Midler and Troschel) Giiulher, Cata-
logue of the fishes in the British Museum, vol. 3, p. 498, 1861, hut restricted to
Cotylis fimbriata Miiller and Troschel, 1849, from Red Sea, and not described
In 1843. Cotylis (liinther has the substitute name Cotylichthys Jordan, Proc.
Acad. Nat. Sci. Philadelphia, 1919, p. 341.]

liryssctacres Jokdan and Evkkmann, ih Jordan, Proc. California Acad. Sci., vol.
«;. p. 230, 189G (genotype: Gobiesox pinniycr Gilbert) ; U. S. Nat. Mus. Bull.
47, pt. 3, p. 2328, 1898 (genotype: G. pin«jV/cr Gilbert).

Caulistius Joroan and Evehman.n, Rep. U. S. Comrn. Fish and Fish, for 1895, App.
p. 491, 1896. (Genotype: Gobiesox papillifer Gilbert.)

Ilryssophilus Jordan and Evermann, U. S. Nat. Mus. Bull. 47, pt. 3, pp. 2329, 2330,
1898. (Genotype: Gobiesox papillifer Gilljert.)

COTYLIS PAPILLIFER (Gilbert)

Gobiesox papillifer Gimiebt, Proc. U. S. Nat. Mus., vol. 13, p. 90, 1890 (Magdalena
Bay, Lower (.'alifornla). — Jokuan and Evi:u.\iann, R(»p. U. S. Comni. Fish
and Fish, for lS9rj, App.. p. 4!»1, 1S96 (Magdalena Bay) ; U. S. Nat. Mus. Bull.
47, pt. 3, p. 2330, 1898 (Magdalena Bay).

Caulistius papillifer Jobda.n, Eveicmann, and Ci.abk, Rep. U. S. Conini. Fish, for
1928, pt. 2, p. 488, 1930 (Magdalena Bay).

Material examined. — Baja California (Magdalena Bay) : U.S.N. M.
No. 44.'J7G (type of papillifer), collected by the Albatross.
Range. — Magdalena Bay, Baja California.

64 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

COTYLIS MICROSPILUS (Fowler)

Plate 1, A

CauUstiiis microspilvs Fowlee, Proc. Acad. Nat. Sci. Philadelphia, 1916, p. 412,
fig. 4 (Panama Bay).

Description. — Head contained about 2%, disk 2.9 or 3, greatest
depth of body 5 to 6, in standard length; eye 21/2 in interorbital
space ; disk a little shorter than length of head ; tip of chin to front
of disk about % to % length of disk ; distance from base of last dorsal
fin ray to midbase of caudal fin contained nearly three times in base
of dorsal fin; dorsal origin nearly an eye diameter closer to mid-
base of caudal fin than to rear of orbit; depressed anal fin reaching
a trifle past a line through caudal fin base, and depressed dorsal fin
reaching to opposite caudal fin base; upper lip on premaxillary with
a median papilla and five more on each side, but none posteriorly on
upper lip; middle of snout with three short papillae or knobs, but
edge of snout above groove without papillae anteriorly but about five
well-developed ones laterally ; another papilla behind rictus and one
on lower lip below rictus; median part of chin and lower jaw with
three rows of papillae, the most anterior being a pair of low lobes,
next a pair of papillae, and the inner row consisting of two pairs
of papillae, with the outer pair posterior to the anterior pair ; lower
lip at each side of median part of chin forming a small lobe bearing
two small papillae ; three large papillae along the inner edge of the
groove along edge of lower lip on each side ; preopercular spine well
developed; three or four pairs of small incisorlike teeth at front of
lower jaw in outer row; teeth at front of upper jaw nearly conical;
lateral teeth of both jaws conical; a small patch of teeth behind outer
teeth at front of both jaws; interorbital space flat; anterior nostril
tubular with a short dermal flap, sometimes branched, arising at the
posterior rim of this nostril; shoulder girdle with a fleshy lobe and
a shallow groove along its lower edge separating it from the lower
less fleshy lobe; base of pectoral fin with a fleshy lobe, the posterior
and ventral margins free, this free margin beginning at point where
gill membrane is fused opposite base of sixth or seventh pectoral
ray from dorsal edge of that fin; upper edge of axial dermal flap
behind pectoral fin is fused to base of fin opposite ninth or tenth ray
from dorsal edge of pectoral fin ; pelvic fin attached to near base of
pectoral fin rays ; lower rays of pectoral fin nearly as long as middle
pectoral fin rays ; margins of disk and pelvic pads of disk all covered
with low flattened papillae; anus much closer to anal origin than
to rear margin of disk.

Coloration. — In alcohol, pale brownish everywhere on dorsal sur-
faces of head and anterior parts of body profusely brown-spotted,
these spots small and rather close together ; tips of all rays of median

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL 96 PLATE t

A, IxjAcr side of head of Colylis microjpiluj (F*'owlcr), V. S. N. M. No. 1U7H2. H, l/)wtr
side of head of Cotylii nigripinnis nif,ripinnis Peters, U. S. N. M. No. 58829. Drawn
bv Mrs \;,„r M Awl.

REVISION' OF AMERICAX CLINGFISHES — SCHULTZ 65

fins white; basally tlio dorsal, anal, and caudal fins are dark brown;
more or less obscure pale bar across base of caudal fin.

Material exaviincd. — The following three specimens, all collected by
Dr. W. L. Schmitt, form the basis of the foregoing redescription of
this species:

U.S.N.M. No. 88822, 26.5 mm., Salinas, Ecuador, September 15, 1926.
U.S.N.M. No. S8823. 58 mm., Guayaquil. Ecuador, 1026.
U.S.N.M. No. 107142, 62.6 mm., Paita, Peru, October 7, 1926.

The three young specimens listed below, also collected by Dr.
Schmitt, are referred to this species with uncertainty. They appear
to be more robust than the adults.

U.S.N.M. No. 101713, 2 specimens, 14.5 and 15 mm., Cupica Bay, Colombia,
January 26, 1935.

U.S.N.M. No. 101938, 1 specimen, 9.5 mm., Cupica Bay, Colombia, .Tanuary
26, 1935.

Range. — Panama Bay to northern Peru.

COTYLIS NIGRIPINNIS NIGRIPINNIS Peterg
Pl.\te 1, B

Cotiilis nigrijjinnis Peteks, Monatsb. Akad. Wiss. Berlin, 1S59, p. 412, May 8,
ISfX) (Puerto Cal)ello fprol)al)iy Venezuela]).

Oobifsox nigripinnin GCntiieb, Catalogue of the fi.-^hes in the British Museum,
vol. 3, p. 502, 1S61 (Puerto Caliello) ; Trans. Zool. Soc. London, vol. 6, p. 390,
1869 (Puerto Cabello). — Jokdan and Evctmann, Rep. U. S. Comm. Fisli and
Fish, for l.s:).5, App.. p. 4lil. 189(; (Puerto Cabello) ; U. S. Nat. Mus. Bull. 47,
pt. 3, p. 2331. 1898 (Puerto Cabello).— Metzelaab, Bijd. Dierk. Feest. Num.
70th Geboortedag van Dr. Max Weber, pt. 22, p. 140, 1922 (Caracas Bay). —
.loBDAN, E\T,RMANN, and CiJVBK, Kei>. U. S. Comm. Fish, for 1928, pt. 2, p.
488, 19.30 f Puerto Cabello).

Oohieaox slrumosun Cope, Proc. Acad. Nat. Sci. Philadelphia, vol. 22, j). 121, 1870
(Hilton Head, S. C). — Jobdan anil Evekmann, Rep. U. S. Comm. Fish and
Fish, for 1895, App., p. 491, 1896 (Carolina to Florida) ; U. S. Nat. Mus. Bull.
47. pt. 3, p. 2.3;}3, 1898 (Ililioii Uvnd, S. C. ; Indian Illvcr, Fla. ; Titusville).—
Hii.DEBKA.ND and ScnKoEnKii, Bull. U. S. Bur. Fish., vol. 43, pt. 1, p. 3.39, 1928
(Chesapeake Bay). — Jobi>an, Evf^mann. and Ci..\bk, Kep. U. S. Comu). Fish,
for 1J>28, pt. 2, p. 489, 1930 (Maryland to Florida).— Ix)noi.ey and Hiu)E-
BRAND, Sysrematip eatalogue of the fishes of Tortugas, Florida, p. 284, 1941
(TorfuKas. Fla.).

Lrf>adt>iiUHt(r nudun {non Linnaeus) Bi.ocH and Scuneideb, Syslema Ichthyo-
loBiae. p. 2, 1801 (locality ?).

CotyliH nudn {nun LlriiiacuH ) Mi li.ku and 'I'uosc iiKi., ITorai- i( hthyoloKJcae. pt. 3,
p. 17. pi. 3. fig. 2. 1849 [West Indies].

Gohirnox nudiin Brihott de Babnevim.e, Rev. Znol. Soc. Cuv., vol. 9, \>. 141, ISIO
(no locality). — f;ONTHra, Catalogue of the fishes In the British Museum, vol.
3, p. 502, 1861 (West Indies; Island of Cordova) ; Trans. Zool. Soc. London,
vol. 6, p. ,3JK), 1869 ((Pardon).

OnhicHoi ff]/rinuif Jordan and Evkicmann, Rep. U. S. Conun. Fish and Fish, for
1895. p. 491, 1896 (nomcn nudum) ', U. S. Nat. Mus. Bull. 47, pt. .3, p. 2.'m.
1898 (West Indies) (I)a8ed on Oobiesox nudug \non Linnaeus] GUnllier). —

66 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Jordan, Evekmann, and Claek, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 489,
1930 (West Indies).

GoMesox virgatulus Goode and Bean, Proc. U. S. Nat. Mus., vol. 5, p. 236, 1882
(Gulf of Mexico) {nomen nudum). — Jordan and Gilbert, Froc. U. S. Nat.
Mus., vol. 5, p. 298, 1882 (Pensacola, Fla. ) .—Jordan, Proc. U. S. Nat. Mus.
vol. 7, p. 149, 1884 (Egmont). — Jordan and Evermann, Rep. U. S. Comm. Fish
and Fish, for 1895, App., p. 491, 1896 (Pensacola Bay to Charleston, S. C.) ;
U. S. Nat. Mus. Bull. 47, pt. 3 p. 2333, 1898 (Pensacola Bay north to Charles-
ton). — Smith, Fishes of North Carolina, p. 374, 1907 (Beaufort Havbor;
Fort Macon; Charleston).— Jordan, Evermann, and Clark, Rep. U. S. Comm.
Fish, for 1928, pt. 2, p. 489, 1930 (Pensacola Bay to Charleston).

OoMesox sancti-martini Metzelaae, Report on the fishes collected by Dr. J. Boeke
in the Dutch West Indies, 1904-1905, pt. 1, p. 151, fig. 48, 1919 (St. Martin,
Simsonsbay Lagoon). — Jordan, Evermann, and Clark, Rep. U. S. Comm.
Fish, for 1928, pt. 2, p. 490, 1930 (West Inaies).

OoUesox bariattilus Starks, The fishes of the Stanford Expedition to Brazil, p.
73, pi. 14, 1913 (Natal). — Ribeiko, Fauna Brasiliense . . . Peixes . . .
Gobiesocidae, p. 2, 1915 (Lagoa em Natal).

"GoMesox yuma Nlchols=[non] Gobiesox vittatus Metzelaar=[non] Gobiesox
punctulattis Poey," Longley, Carnegie Inst. Washington Year Book No. 34,
p. 284, 1935.

"Gobiesox virgatulus Jordan and Gilbert=G. strumosus Cope," Longley, Carnegie
Inst. Washington Year Book, No. 33, p. 270, 1934.

"Gobiesox barbatulus Starks= Goftiesoa? gyrimis Jordan and Evermann=Go&iesoa?
nigripinnis Peters," Longley, Carnegie Inst. Washington Year Book, No. 34,
p. 284, 1935.

Gobiesox yuma Nichols, Bull. Amer. Mus. Nat. Hist., vol. 37, No. 37, p. 876, fig. 1,
1917 (Sanibel Light, Fla., vrest coast). — ? Beeder, Bull. Bingham Oceanogr.
Coll., vol. 1, art. 1, p. 85, 1927 (Royal Islands, Bahamas). — Jordan, Ever-
mann, and Claek, Rep. U. S. Comm. Pish, for 1928, pt. 2, p. 490, 1930
(Florida).

Remarhs. — Miiller and Troschel's description of Cotylis nuda
(1849, pp. 17-18) leaves little doubt that their species is the same as
the one recognized here as nigripinnis, since small barbels are said
to occur around the mouth and the coloration is brownish with streaks
of dark spots. In addition, fin rays are given as dorsal 12, anal 7.

Wlien the form along the Atlantic coast from Chesapeake Bay to
the east coast of Florida is studied in the minutest detail, it may be
recognized as distinct from nigripinnis, but I have not thoroughly
investigated the variation in the various localities from Maryland to
Brazil. There are several names available for the races or subspecies
that may be recognized.

Dr. S. F. Hildebrand kindly turned over to me the notes made by
Dr. W. H. Longley at Amsterdam on the type of Gobiesox sancti-
martini Metzelaar. I quote:

T. L. [total length] G9 mm. D. 12, A, 8, P. 23-24 including a stub above.
Diameter of eye (orbit) 3.0 mm. Interorbital width 7.0 mm. Nasal cirri
expanded, bilobed, vpithout fringe. Twenty-nine coarse cirri, becoming bulbous
under pressure of the tissue behind them, along front of ventral disk in single

REVISION OF AMERICAN CLINGFISHES — SCHULTZ

series. The fleshy border lateral to them only slightly crenulated before the
anterior ray of the ventral fin. The lower angle of the pectoral moderately
prouiinent, not cxserted. Opercular cleft extending upward to the base of Gth
ray, the fold before the base of the fin complete, continuous with the fleshy
border of the operculum. Anterior teeth little if any flattened, the lateral in
the upper j;iw running in behind tiie front but not as regularly as in some. . .

Table .3. — Counts and measurements made on species of Cotylis, expressed
in hundredths of the standard length

Characters

microtpilut

nigripmnia
nigripinnis

ni(;TipinnU
woodii

U.S.N.M.
No. 88822

U.S.N.M.
No. 88823

U.S.N.M.
No. 107142

U.S.N.M.
No. 87752

F.M.N.H.
No. 41974

Standard length (in milli-
metors)

26.5
41.5
15.8
32.0
8.30

9.44
11.7

7.55
10.9
26.4
35.2

9.44
13.6

10.9
6.04
61.1
73.2
64.9

32.5
21.1
17.0
15.1
30.6
15.8

14.0
24-26
14
10

6-7

41.2
18.3
31.1
9.32

8.62
12.2

5.86
14.7
24.2
34.5

11.2
13.0

11.7
4.14
59.2
70.7
66.9

32.4
22.1
15.0
12.4
26.7
15.5

12.9
27-27
14
10

7-7

62.6
41.0
18.2
35.1
8.79

8.03
13.3

5.59
13.6
2.5.7
34.3

9.60
16.0

13.6
4.80
60.0
70.2

65. 5

33.5
21.6
16.0
12.8
26.4
16.5

12.1
25-26
14
10

6-«

40.6

18.8

39.1

10.1

9.42
15.5

5.50
13.0
24.3
36.5

12.3
15.2

11.7
7.10
64.5
69.6
62.3

25.5
21.7
14.8
12.0
24.3
16.2

13.0
25-26
11

0-0

42.0

22.2

35.0

11.5

10.5
12.8
5.25
12.7
25.8
38.7

12.5
12.3

10.5
9. 25
65.0
74.0
65.0

29.0
20.0
14.3
13.0
20.3
18.3

1.5.5
25-25
11

0-7

Len^h of head

41 2

Greatest depth of body

Greatest width of head

Lenf;th of caudal podunclo

Least depth of caudal pe-

Hnnrln

13.7
36.4
7.58

Length of snout

11 2

Diameter of eye

7 68

Width of intororbital space. ..

Postorbital length of head

Length of disk

9.40
22.8
32 4

Distance from base last dorsal

ray to midcaudal fln base...

Gape or tip of snout to rictus. .

9.70

Distance from rear margin of
disk to anus

11 2

Anus (center) to anal origin...

Snout tip to dorsal origin

Snout to anal origin

10.9
69.1
77 3

Snout to anus

63 7

Length of dorsal fln base (to
base of last ray)

25 8

Length of anal fln base

Longest ray of dorsal fln

Longest ray of anal fln

Longest ray of caudal fln

Longest ray of pectoral fln

Length of third ray from bot-
tom of p*«toral On

16.7
11.2
11.5
20.4
12.4

11.6

Numijer of pectoral fln rays...
Dorsal rays

22-22
10

Anal rays

Number of upper pectoral
rays above upper edge of
attachment of gill mem-
branes

6-0

See table 3 for measurements made on two specimens from Brazil.

I have cxaiiiinod tlie type of Gohirsox ynjiui Nichols and find that it
jjosse.sses the harbcllike .structures around the mouth and in other
respects resembles Cotylis nigripinnis nigHpinnis to which I refer it

68 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

as a synonym. The teeth of the lower jaw at the front have uneven
edges but are not bifid or trifid as in certain other genera. I count
dorsal rays as 12, anal as 9, and pectoral 23.

The following two collections, referred to this species with uncer-
tainty, contain very small specimens that do not show certain charac-
ters fully developed and may represent an undescribed species of
small size :

U.S.N.M, No. 83862, 4 specimens, 10.5 to 11 mm., Trinidad, Albatross, January
SO-February 21, 1884.

U.M.M.Z. No. 131173, 4 specimens, 8.3 to 9.5 mm., Velasco, Tex., Rice Institute,
April 17, 1923.

Material examined. — Maeyland and Virginia: U.S.N.M. Nos.
30400, 30407, 43064, 68829, 67760, 67761, 68391, 74852, 76530, 76531,
77929, 83593, 85087, 85681, 86313, 88583, 88586, 89340, 91210-91239,
92024, 92031, 93759, 93805, 104930, 109846, 122392, 122394, totaling 420
specimens. North Carolina: U.S.N.M. Nos. 4905, 85088, 122395,
122396, total 4 specimens. South Carolina: U.S.N.M. Nos. 26311,
59053, 59061, totaling 9 specimens. Florida: U.S.N.M. Nos. 26611,
30471, 30861 (2 cotypes of virgatulus) , 32760, 34719, 34725, 73250, 85089,
85090, 91456, 92213, 92215, 93716, 93882, 94896, 116933-116935, 125493,
totaling 31 specimens. Alabama: U.S.N.M. No. 73545, 2 specimens.
Mississippi : U. S. N. M. No. 32625, 1 specimen. Louisiana : U.S.N.M.
Nos. 86134, 122393, 124979, totaling 3 specimens; U.M.M.Z. No. 128860,
3 specimens. Texas: U.S.N.M. Nos. 69347, 69348, 118542, totaling 4
specimens; U.M.M.Z. Nos. 111746 and 114471, totaling 15 specimens.
Brazil: U.S.N.M. Nos. 87752 and 87798, totaling 3 specimens.

Range. — Chesapeake Bay to Brazil ; West Indies.

COTYLIS NIGRIPINNIS WOODSI, new subspecies

Holotype. — F.M.N.H. No. 41974, a specimen 33 mm. in standard
length, from Cocos Island at Wafer Bay, collected February 23, 1941.

Description of only known specimen. — Detailed measurements were
made and these are recorded in hundredths of the standard length in
table 3.

Head contained about 2^/^, disk 3, greatest depth of body about 7, in
standard length ; eye equal to bony interorbital space and 1% in fleshy
interorbital space; disk about 1.3 in head; tip of chin to front of disk
about % leng-th of disk ; distance from base of last dorsal ray to mid-
base of caudal fin 2.7 in length of base of dorsal fin; dorsal origin
equidistant between midbase of caudal fin and base of upper pectoral
ray; tips of rays of depressed anal fin reaching a little past a line
through base of caudal fin and depressed dorsal fin not reaching quite
to that line; size and arrangement of papillae around mouth essentially
as described for nigripinnis ; about three pairs of incisorlike teeth at

REVISION OF AMERICAN CLINGFISHES — SCHULTZ 09

front of lower jaw projecting obliquely forward, followed laterally
by one or two somewhat enlarged conical teeth, then posteriorly by a
short row of small conical teeth ; inside of larger outer row of teeth
a few smaller ones at front of lower jaw ; upper jaw with conical teeth,
those at front a little enlarged; none of the teeth with trifid tips; front
of upper jaw inside of outer teeth with a few minute teeth ; interorbital
space flat ; each anterior nostril with a bifid dermal flap on posterior
margin ; shoulder girdle with a fleshy lobe on its lower margin under
gill cover; base of pectoral fin with a fleshy lobe, the posterior and
ventral margins with a free edge that extends to the attachment of the
opercular membrane, both of which are fused opposite the base of the
sixth pectoral fin ray; upper edge of axial dermal flap behind pectoral
fin fused to base of fin opposite the sixth pectoral ray; pelvic fins at-
tached near base of about fourth pectoral fin ray; lower rays of pec-
toral fin nearly as long as middle rays ; margins of disk and pelvic pads
with low flattened papillae; anus a trifle closer to anal origin than to
rear margin of disk.

Coloration. — General coloration pale brownish in alcohol, with five
wide indistinct bars on body, the paler interspaces narrower than
eye ; sides of body with several very narrow pale lines; a dark elongate
spot behind eye and a few narrow pale lines radiating posteriorly from
orbit across gill cover; median fins black with tips of rays white.

Rnnarkft. — This new subspecies is the representative of a similar
form in the Atlantic from Maryland to Brazil herein recognized under
the n;inie 7Uf7ripinnift. From that form iroodsi may be distinguished
by a larger eye and a less deep body, as indicated in the key.

Named voodul in honor of Lt. Loren P. Woods, U. S. N. R., who
tentatively suggested this specimen to be an undescribed species when
he learned that I was studying the American clingfishes. Described
with the permission of the authorities of the Chicago Natural History
Museum.

COTYLIS PINNIGER (Gilbert)

OohtcHox pinnigcr Gn.BERT, Vroc. U. S. Nnt. Mus., vol. ];i, p. W, 1890 (Puerto Re-
fuKld. AriKf'l Islnrid. San Luis GotizaloH I'ay, and La Paz, Gulf of California). —
Ilxr.EfRiN, P.ull. Mns. Hist. Nat. Paris, vol. 7, ii. 1<;2. 1001 (fJiilf of r'alifornin).

Brj/nxrlfirrf X pitinif/rr .Fobdan and Evkbmnnn, I'roc. California Acad. Sci., vol. 0, p.
2.^ pi. .'51. ISrtf,; IN-p. t:. S. Conin*. Fish and Fisli. for m'tr>. App.. p. 401, 1806
(Gulf of California) ; U. S. .Nat. Mns. I'.ull. 17, p. 12.12S. ISOS (Gulf of Cali-
fornia). — JouD.VN, EvFUMANN, and CiJVKK, Hep. U. S. Conini. Fisl). for 1028,
pt. 2, p. 4.S8, 10.30 (Gulf of California). -KnFOJiat, Bull. Binpham Oconnogr.
Coll., vol. 2. art. 3, p. 4H, 10.36 (Puerto Krfupio ; Gonznp) Ba.v).

ReTnarTcH. — The longer base of the dorsal fin is not considered of gen-
eric significance in view of other related species with dorsal fins of
nearly the same length.

70 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Material examined. — Gulf of California: U.S.N.M. Nos. 44377
(type of pinniger), 46694 (4 cotypes of pinniger), 126808 (25 cotypes
of pinniger) , totaling 30 specimens ; F.M.N.H. No. 3338, 19 specimens.

Range. — Gulf of California.

Genus SICYOGASTER Brisout de Barneville

Sicyogaster Brisout de Baenevuxe, Rev. Zool. Soc. Cuv., vol. 9, p. 144, 1846.

(Genotype: Ooiiesox marmoratus Jenyns.)
Caularchus Giix, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 330, 1862.

(Genotype: Caularchus reticulatus=^Lepadogaster rcticulatus Girard.)

SICYOGASTER MARMORATUS (Jenyns)

Oobiesox marmoratus Jenyns, The zoology of the voyage of H. M. S. Beagle,
pt. 4, Fishes, p. 140, pi. 27, figs. 1, la, lb, 1842.— Gunther, Catalogue of the
fishes in the British Museum, vol. 3, p. 504, 1861 (Chile). — ?Abbott, Proc.
Acad. Nat. Sci. Philadelphia, 1899, p. 363 (Peru).— Delfin, Rev. Chilena
Hist. Nat, vol. 3-4, p. 91, 1901 (Algarrobo; Chauaral ; Punta Arenas; Cal-
buco ; Iquique; Isla de Juan Fernandez). — Tortonese, Bol. Mus. Zool. Anat.
Comp. Univ. Torino, vol. 47, p. 206, 1939 (Valparaiso).

Sicyogaster marmoratus Beisout de Baenevuxe, Rev. Zool. Soc. Cuv., vol. 9, p. 144,
1846 (Chile).

Cotylis marmoratus MvIi-lee and Troschel, Horae ichthyologicae, pt. 3, p. 19,
1849 (Chile).

Remarks. — The following notes on the type of Gohiesox marmora-
tus in the British Museum from "Archipelago of Chiloe," made by Dr.
W. H. Longlefy, were kindly turned over to me by Dr. S. F.
Hildebrand :

Two specimens of T. L. [total length] 56 and 64 mm. considerably macerated,
the smaller better preserved. D. 12; A. 10; the last anal ray missing, but its
support still evident. The pectoral both sides with 23 rays including the rudi-
mentary one above. In the larger fish D. 12, A. 11.

In the small fish again I found that the membranous structure at pectoral
base is evident for only half the vertical height of the fin but that in the lower
half, where it is present, it exists as a very evident, freely projecting lobe.

On very careful examination, I find that the opercular cleft extends dorsally
about to the base of the upper pectoral ray.

Material examined. — Peru: U.S.N.M. No. 101706, 1 specimen.
Chiles U.S.N.M. Nos. 77381, 88819-88821, 88824, totaling 6 specimens.
Range. — Peru and Chile.

SICYOGASTER MAEANDRICUS (Girard)

Lepadogaster reticulatus Girard, Proc. Acad. Nat. Sci. Philadelphia, 1854, p.

155 ( San Luis Obispo, Calif . ) (preoccupied).
Lepadogaster maeandricus Girard, Explorations and surveys for a railroad

route from the Mississippi River to the Pacific Ocean, vol. 10, pt. 4, p. 130,

1858 (San Luis Obispo, S. Faralones, Calif.) (new name).
Oobiesox maeandricus GiiNTHER, Catalogue of the fishes in the British Museum,

vol. 3, p. 505, 1861 (Monterey).
Caularchus reticulatus Gill, Proc. Acad. Nat. Sci. Philadelphia, vol. 14, p. 330,

1862.

REVISION OF AMERICAN CLINGFISHES — SCHULTZ 71

Gobicsox rcticulatun Jordan and Joi'Y, Proc. U. S. Nat. Mus., vol. 4, p. 5, 1881
(Monterey and Cniie Flattery). — Joiu)an and GiiJiUix, Proc. U. S. Nat. Mus.,
vol. 4. p. 63. ISSI (Monterey to Pu.uet Sound).— Rosa Smith, Proc. U. S.
Nat. Mu.s., vol. 4. p. 140, 141. iSSl (San Diego).

Caularchus nincaiuiricus Joiu)an and Evkkma.nn. Rep. U. S. Comm. Fish and
Fish, for ISO'), App., p. 491, 1S96 (Vancouver Island to Monterey) ; U. S.
Nat. Mus. Bull. 47, pt. 3, p. 2328, 189S (Vancouver Island to Point Concep-
tion).— Ouitel, Arch. Zool. Exper., vol. 5, No. 5, pp. 625-()39, 1906 (anat-
omy). — EvKEMANN and GoiJ)SB()KOUGH, Bull. U. S. Bur. Fish., vol. 26 p.
336, 1907 (Fort Rupert; Gabriola Island). — Joudan, E\'ei:mann. and Clabk,
Rep. U. S. Comm. Fish, for 1928. pt. 2, p. 4S8. 1930 (Vancouver Island to
Monterey). — Schultz, Keys to the tishes of Washington, Oregon and closely
adjoining regions, ed. 1, p. 197, 1930 (British Columbia to Point Arguello,
Calif.). — ScHULTZ and DeLacey, Journ. Pan-Pacific Res. Inst. (Mid-Pacific
Mag.), July-September, 1936, p. 211, 213 (British Columbia to Point
Arguello, Calif.) (see this reference for additional references). — Wilby,
Copeia, 1936, p. 116 (British Columbia).

Material examined. — British Columbia : U.S.N.M, Nos. 49083,
60.548, 60549, 64022, 82153, 82154, 103563, 103564, 103566, 103567, 120446,
120447. 12G811. totalin*!; 64 specimens. Washington: U.S.N.M. Nos.
23405, 27329, 3S334, 42049, 83208, 83964, 103565, totaling 32 specimens.
Ori:gon: U.S.N.M. No. 91974, 1 specimen. California: U.S.N.M.
Nos. 516 (type of reticulatus^maeandricus) , 101382, 101383, 101388,
totaling 4 specimens.

Range. — Queen Charlotte Islands to San Diego, Calif. ; Puget Sound.

ARCOS, new genus

Genotype. — Gobiesox erythrops Gilbert.

This genus is characterized by the groove along the anterior or upper
margin of the preniiixillary which arches in a convex manner over the
tip of the snout; the orbits are larger than in any other genus of Amer-
ican clingfishes, their diameter much greater than the least width of the
bony interorbital. In addition, the axial dermal flap bcliind the pec-
toral lin has its dorsal edge attached mucii above the niidbase of pec-
toral ; the incisorlike teeth at front of lower jaw have smooth tips, and
lhe.se teeth project forward horizontally and do not oppose the teeth
at front of upper jaw, whicli are nearly conical; there are no papillae
around the mouth, although the usual lobelike ridges occur on lower
jaw and chin; gill membranes are joined at upper edge of pectoral fin
ba.se or appear to be opposite base of first pecloral fin ray; the free
posterior margin of fleshy pad on outer surface of pectoral base is con-
fined to the lower half of that fin and not iibove the thirteenth ray from
the top. Other characters are tlio.se of the genotype.

Names Arcos in reference to the arched groove on the tip of the
snout.

72 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

ARCOS POECILOPHTHALMUS (Jenyns)

Gohiesox poecilophthalmus Jenyns, The zoology of the voyage of H. M. S. Beagle,
pt. 4, Fishes, p. 141, pi. 27, fig. 2, 2a 2b, 1842 (Chatham Island).— GtJNTHER,
Catalogue of the fishes in the British Museum, vol. 3, p. 503, 1861 (Chatham
Island). — Jordan and Eveemann, Rep. U. S. Comm. Fish and Fish, for 1895,
App., p. 491, 1896 (Chatham Island) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2335,
1898 (Chatham Island).

Cotylis poecilophthalnius Mtjller and Teoschel, Horae ichthyologicae, pt. 3, p. 19,
1849 (Galapagos).

Tomicodon poecilopMMlmos Bbisout de Barne\ille, Rev. Zool. Soc. Cuv., vol. 9,
p. 144, 1846.

GoUesox erythrops Jordan and Gilbert, Proc. U. S. Nat. Mus., vol. 4, p. 360, 1882
(Mazatlan). — Jordan and Evermann, Rep. U. S. Comm. Fish and Fish, for
1895, App., p. 491, 1896 (Mazatlan ; Tres Marias Island) ; U. S. Nat. Mus. Bull.
47, pt. 3, p. 2336, 1898 (Mazatlan) . — Jordan, Evermann, and Clark, Rep. U. S.
Comm. Fish, for 1928, pt. 2, p. 490, 1930 (Mazatlan; Tres Marias Island).

GoUesox paradiseus Herre, Publ. Field Mus. Nat. Hist., zool. ser., vol. 18, p. 432,
1935 (Eden Island ; South Seymour Island) ; vol. 21, p. 393, fig. 36, 1936 (Eden
Island; South Seymour Island).

Remarks. — This species is recognizable by its very large eyes and
narrow interorbital space. It is a small species and usually red in
color.

Dr. S. F. Hildebrand kindly turned over the following note by Dr.
W. H. Longley on the type of Gohiesox 'poecilophthalmus from Chat-
ham Island :

T. L. [total length] 45 mm. D. 8, A. 7, P. 21+ rod [or 22 rays].

Material examined. — Mazatlan: U.S.N.M. No. 30885 (type of
erythrops). Panama Bay (Secas Islands) : U.S.N.M. No. 101708, 5
specimens. Galapagos Islands: U.S.N.M. No. 65516, 1 specimen;
F.M.N.H. Nos. 17404 and 17405 (type and paratype of paradiseus) .

Range. — Mazatlan to Panama and Galapagos Islands.

ARCOS MACROPHTHALMUS (Giinther)

GoMesox macropMhalmus Gunthee, Catalogue of the fishes in the British
Museum, vol. 3, p. 502, 1861 (habitat unknown) [probably West Indies]. —
Jordan and Evermann, U. S. Nat. Mus. Bull. 47, pt. 3, p. 2335, 1898 (St.
Thomas). — Metzelaar, Bijd. Dierk. Feest. Num. 70th Geboortedag van Dr. Max
Weber, pt. 22, p. 140, 1922 (Caracas Bay).— Beebe and Tee- Van, Zoologica,
vol. 10, No. 1, p. 251, fig., 1928 (Lamentin Reef, Port-au-Prince Bay, Haiti). —
Jordan, Evermann, and Clark, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 489,
1930 (probably West Indies).

Gohiesox cerasinus Cope, Trans. Amer. Philos. Soc, vol. 14, p. 413, 1871 (St. Mar-
tins, West Indies).— Jordan and Evermann, Rep. U. S. Comm. Fish and Fish,
for 1895, App., p. 492, 1896 (St. Martins) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2336,
1898 (St. Martins). — Fowler, Proc. Acad. Nat. vSci. Philadelphia, vol. 71, p.
143, 149, 1919 (St. Martins; Jamaica).—? Metzelaae, Bijd. Dierk. Feest.
Num. 70th Geboortedag van Dr. Max Weber, pt. 22, p. 140, 1922 (Caracas
Bay). — Jordan, Eveemann, and Clark, Rep. U. S. Comm. Fish, for 1928, pt.
2, p. 489, 1930 (St. Martins, West Indies).

REVISION OF AMERICAN CLINGFISHES — SCHTJLTZ 73

Gobicsox tudes Bvtujmann and Maksh. Bull. U. S. Fish Couun., vol. 20, pt. 1, p.

305, 1900 (Culebru. Puerto Rico).
Sicyancs yumurina Rivebo, rroc. Boston Soc. Nat. Illst., vol. 41, No. 4, p. 74,

1936 (Matanza.s, reef at entrance of the Bay).
Ouhiesox androi^icHxis Rcsen, Lunds Univ. Arf^-Skr., now .sor. (Afd. Math. Nat.),

vol. 7. No. 5, p. 6."), 1911 (Ma.'^tic Point, Andros; Nas.sau. Bahamas). — Jokdan,

EvEKMANN. and Clabk, Rep. U. S. Comm. Fi.<;h. for 1928, pt. 2, p. 4S9, 1930

(Bahamas).
Gohienox androHiensis Rosen=[not] Oobiesox riihifjinosus (I'oey), Longley,

Carnegie Inst. Washington, Year Book, No. 34, p. 284, 1035.
Oobicsox ccphalus Metzei^vak, Bijd. Dierk. Feest. Num. 70th Geboortedag van Dr.

Max Weber, pt. 22, p. 139, 1922 (Caracas Bay) (see comment below).
Gobicsox ma:'rophthaU}ius Jordan and E\'ermann, Rep. U. S. Comra. Fish and

Fish, for 1895, App., p. 492, 1896 (West Indies).

Re77ia7'A-s. — Tliis species has the largest eyes of any American form
in the Athintic and is red in color when alive. The eyes are much
wider than the narrow int^rorbital space.

Tlirough the kindness of Dr. Thomas Barbour I have examined a
paratype of Sicyases yumurina Kivero and refer it to this species.
Some traces of the red color still remain on this specimen.

Dr. S. F. Hildobrand very kindly turned over to me the following
wuiQs, by Dr. W. H. Longley made on the type of Gohiesox niaci'oph-
thalmus Giinther in the British Museum:

T. L. [total length] .'4 mm. D. 8; A. 7. First ray in each tin very slightly
tiiamentous. I'. 22 and a short, vestigial uijper 23rd. Same on both sides. The
outline of the fin rounded. A strong subopercular spine with a deep groove on
Its ventral surface and reaching beyond the base of any of the pectoral rays.
The membranous sac at the ba.se of the pectoral extends upward only to the
base of the eighth ray counting up from the ventral margin, but is a very evident
structure. The opercular cleft is of the full width of the iMX-toral base and
extend.s up to the level of the upper margin of the dwarf ray, which is quite a
sizable stub one-third the length of the secctnd.

Eye 5.0 mm.=snout; bony interorbital — 3.0 nim.=preorbital width . . .

The dorsal origin midway between tip of caudal and posteri(»r margin of the
I)upil.

Di'. Uddf'brand al.M) turned over to nie the following notes made by
Dr. Longley in the Berlin Museum, on the probable types of Gohiesox
an//ronirnftis collected by Rosen in the Bahamas:

Spec. 1. T. L. [total length) 2^5 mm. 1 ►. 7. A. 6, P. 24-24. the outer ray short.
Two or rhn^ pairs of teeth above slightly flattened. Three pairs below more
flattened, the anterior distinctly etdarged. The bnrder of the lower inci.sors only
slightly crernilated, more nearly tnuicate fhan on No. 3. The branchial deft ex-
tending up to the base «»f th(! upper p<'cloral ray. No groove on the siibopercidar
spine. The nasal cirrus is a flap half flic diameter of the narlal orifice In width.
On one side It ends In two filaments . . ,

Spec. 2. T. L. 25 nmi. I). 8, A. 7. Gill apparatus as above . . .

Sp«'C. 3. T. L. 22 mm. D. 0. A 7, P. 22-22. Gill apparatus as above . . .
Anterior face of lower Incisors slightly fluted, the fre<? border of the teeth almost
bicuspid.

74 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Dr. Hildebrand furnished me the following notes made by Dr.
Longley in the Museum at Amsterdam on specimens reported upon by
Metzelaar (1922) from Curasao (Caracas Bay) :

Oobiesox macrophthalmus . . . much fringed nasal cirri, the wide opercular
cleft, the incomplete fold behind it with isolated lower lobe.

T. L. [total length] 70 mm. D. 8, A. 7, P. 24-24 including stub.

Gohiesox cephalu>s ... Is the same as last [macrophthalmus] . . . T. L. 20
mm. D. 8, A. 7, P. 23-23, stub included . . . The gill cleft extends entire width
of base of pectoral fin. I get no fold at all along the fin base. The anterior
teeth of the lower jaw are enlarged, the middle much flattened and larger than
the next pair.

Material examined. — Bahama Islands: U.S.N.M. Nos. 38386 and
53220, totaling 2 specimens. Jamaica : U.S.N.M. No. 78142, 1 speci-
men. St. Thomas : U.S.N.M. Nos. 78157 and 78158, totaling 2 speci-
mens. San Juan Island : U.S.N.M. No. 117423, 4 specimens. Virgin
Islands : U.S.N.M. No. 117412, 3 specimens. Martinique : U.S.N.M.
No. 117452, 2 specimens. Puerto Kico: U.S.N.M. No. 126181, 1
specimen.

Range. — Bahama Islands and West Indies.

Genus GOBIESOX Lacepede

Oohiesox Lacepede, Histoire naturelle des poissons, vol. 2, p. 595, fig., 1800.

(Genotype: OoMesox cephalus Lacepede.)
Megaphalus Rafinesque, Analyse de la nature, p. 86, 1815. (Genotype: Oobiesox

cephalus.) (Substitute name for Oobiesox.)

GOBIESOX CEPHALUS Lacepede

Oobiesox cephalus Lacepede, Histoire naturelle des poissons, vol. 2, pp. 595, 596,
fig., 1800 (fresh-water rivers of South America). — Brisout de Barneyille,
Rev. Zool. Soc. Cuv., vol. 9, p. 145, 1846 (Martinique). — Gxjnthee, Catalogue
of the fishes in the British Museum, vol. 3, pp. 499, 566, 1861 (Caribbean
Sea; St. Domingo; West Indies). — Jordan and Eveemann, Rep. U. S. Comm.
Fish and Fish, for 1895, App., p. 491, 1896 (West Indies) ; U. S. Nat. Mus.
Bull. 47, pt. 8, p. 2332, 1898 (Caribbean Sea ) .— GinxEL, Arch. Zool. Exper.,
vol. 5, No. 5, pp. 640-645, 1906 (anatomy). — Blosser, Ann. Carnegie Mus.,
vol. 6, p. 300, 1909 (St. Croix).— Fowler, Proc. Acad. Nat. Sci. Philadelphia,
vol. 71, p. 143, 1919 (St. Martins, West Indies).— Jordan, Evermann, and
CL-^RK, Rep. U. S. Comm. Fish, for 1928, pt. 2, p. 489, 1930 (West Indies).—
?RiVEEO, Proc. Boston Soc. Nat. Hist, vol. 41, No. 4, p. 73, 1936 (Habana).

Oobiesox tudes Richardson, Zoology of the voyage of H. M. S. Sulphur, Ichthyol-
ogy, vol. 1, p. 103, pi. 46, figs. 1-3, 1844 (locality unknown ) .—Jordan and
Eveemann, U. S. Nat. Mus. Bull. 47, pt. 3, p. 2333, 1898 (West Indies).—
Joedan, Eveemann, and Clabk, Rep. U. S. Comm. Fish, for 1928, pt. 2,
p. 489, 1930 (probably West Indies).

Oobiesox tudes 'Richardson= Oobiesox cephalus Lacepfede, Longlet, Carnegie
Inst. Washington Year Book, No. 34, p. 284, 1935.

Cotylis stannii MtJiXER and Tboschel, Horae ichthyologicae, pt. 3, p. 18, pi. 3,
fig. 3, 1849 (Brazil).

Oobiesox costaricensis Meek, Publ. Field Mus. Nat. Hist., zool. ser., vol. 10, No.
7, p. 74, 1912 (Zent River, Atlantic drainage, Costa Rica). — Behre, Ann.

REVISION OF AMERICAN CLINGFISHES — SCHULTZ 75

Carnegie Mus., vol. IS, p. 314, 192S (tributary to Rio Cricatuola near Con-
quautu). — JOKOAN, Evkkmann, and Ci-\kk, Kep. U. S. Couim. Fish, for 1U2S,
pt. 2, p. 4S9, 1930 (Costa Kica).
Gobiesox ramsdeni RnxEo, Proc. Boston Soc. Nat. Hist., vol. 41, No. 4, p. 73,
193G (Rio Toa. "El Palenque" Yateras, GuautAnanio, in Orionte Province,
Cuba ) .

Remarks. — Dr. S. F. Hildebrand kindly turned over to me the fol-
lowintr note made by Dr. W. H. Longley in the Paris Museum of
Natural History on a specimen, No. 513-1:, of Gobiesox cephalus Lace-
pede:

T. L. [total length] 9.") mm., D. S, A. G, P. 21 and a stub on outer sitle. If any
are Lacei)ttde's specimens, this must be it, others are all too late.

Contrary to most references in the literature, Lacepede did not report
G. ccphalus from the ''Caribbean Sea" but from fresh-water rivers of
South America. Since Lacepede mentions fresh water once and rivers
twice in his dc-^cription as the habitat of this species, I assume he did
not make a mistake in the locality where the species occurred.

1 have before me a series of specimens of Gobiesox from fresh-water
streams of northern South America and Central America on the At-
lantic side, and these agree in most characteristics throughout the area
represented. Noteworthy is the arrangement of the teeth. On the
lower jaw anteriorly the teeth of the outer row are somewhat enlarged,
short, narrow and incisorlike, not crowded or projecting forward at the
symphysis; front of upper jaw with conical teeth; lateral teeth in
both jaws conical ; sometimes with one or two enlarged caninelike teeth
at front sides of lower or upper jaws; upper jaw with an inner patch
of small conical teeth ; origin of dorsal fin usually equidistant between
midbase of caudal fin and tips of pectoral fin rays; anus usually equi-
distant between anal origin and rear margin of disk or a little closer to
anal origin ; anal origin behind middle of bases of dorsal fin rays or
under the liftli or sixth ; anus slightly in front of a vertical line through
dorsal origin; head 2.2 to 2.G; disk 2.G to 3.2; depth 4 to 5.5, all in
standard length; interorbital equals snout.

There is a black blotch near base of dorsal fin on first rays that ap-
pears to occur constantly on the specimens exiunined.

William C. Schroeder, Mu.seum of Comparative Zoology, kindly
checked the type of Gobiesox raiiisdeni (M. C. Z. No. 34152) and made
the following observations:

Posterolateral teeth c>f lower jaw more rnniiiclike and not smaller than front
tf^eth; no papilla on upiH'r or lower jaws (unless I overlooki-d this) ; length of
disk e^juals disk to midbase of anal; anus closer to anal origin than to rear
ninrpln of disk by an i'ye's diameter. Standard length 107 [mm.]; head 43;
length of disk 30; depth of body 25; D. 8; A. 5 or 6; I'. 20; eye .^; interorbital 10.

The foregoing counts may be considered as correcting those given in
the original description of ramsdeni.

76 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Matenal examined. — ^West Indies: U.S.N.M. Nos. 26479, 29848,
93793, totaling 11 specimens. Costa Rica : U.S.N.M. No. 74246 (para-
type of costarwensis) ; F. M. N. H. Nos. 7677 and 7813 (type and 2
paratypes of costaricerisis) , 7814, 7815, totaling 6 specimens. Vene-
zuela (Macuto): U.S.N.M. Nos. 93815-93817, 93820-93822, 93827,
totaling 16 specimens. Colombia (Eio Dagua) : U.S.N.M. No. 120217,
1 specimen.

Range. — ^West Indies; Costa Rica to Brazil.

GOBIESOX FULVUS Meek

Gobiesox fulvus Meek, Publ. Field Columbian Mus., zool. ser., vol. 7, No. 5, p. 149,
1907 (Cocos Island).

Remarks. — The following specimens were studied: Cocos Island:
F. M. N. H. No. 6035 (type of fulvus) ; U.S.N.M. No. 91832, 1 speci-
men ; U. M. M. Z. Nos. 131512 and 131513, 2 specimens.

Range. — Cocos Island.

GOBIESOX PUNCTULATUS (Poey)

Sicyases punctulatus Poet, Enumeratio piscium Cubensium, p. 124, 1876
(Cuba). — Jordan, Evekmann, and Clark, Rep. U. S. Comm. Fish, for 1928, pt.
2, p. 490, 1930 (Cuba).

Gobiesox punctulatus Jordan and E\'ekmann, Rep. U. S. Comm. Fish and Fish, for
1895, App., p. 492, 1896 (Cuba) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2338, 1898
(Cuba). — Metzeiaak, Bijd. Dierk. Feest.-Num. 70th Geboortedag van Dr.
Max Weber, pt. 22, p. 140, 1922 (Caracas Bay).

Gobiesox Jiaeres Jordan and Bollman, Proc. U. S. Nat. Mus., vol. 11, p. 552, 1889
(Green Turtle Cay, Bahamas). — Jordan and Evebmann, Rep. U. S. Comm.
Fish and Fish, for 1895, App., p. 492, 1896 (Green Turtle Cay, Bahamas) ;
U. S. Nat. Mus. Bull. 47, pt. 3, p. 2337, 1898 (Green Turtle Cay, Bahamas).

Gobiesox nudiis Metzelaar, Report on the fishes collected by Dr. J. Boeke in the
Dutch West Indies 1904-1905, pt. 1, p. 151, 1919 (Curasao and Bonaire).

Gobiesox l[h]aeres Jordan and Bollman=Gf. punctulatus (Poey), Longley, Car-
negie Inst. Washington, Year Book, No. 33, p. 271, 1934.

Gobiesox vittatus Mepzexaae, Bijd. Dierk. Feest.-Num. 70th Geboortedag van
Dr. Max Weber, pt. 22, p. 140, fig. 3, 1922 (Caracas Bay ).— Jordan, Evekmann,
and Clark, Rept. U. S. Comm. Fish, for 1928, pt. 2, p. 489, 1930 (Curagao).

Sicyases haeres Jordan, Eveemann, and Clark, Rep. U. S. Comm. Fish, for 1928,
pt. 2, p. 490', 1930 (Green Turtle Cay, Bahamas).

Remarks. — This is a small species and may be recognized by its
relatively short, thick and deep body giving it the appearance of
robustness.

Dr. S. F. Hildebrand kindly turned over to me the notes made by
Dr. W. H. Longley in the Museum at Amsterdam on the type of
Gobiesox vittatus Metzelaar, which follow :

T. L. [total length] 29.0 mm., D. 11, A. 7. P. 1^20, orbit 2.0 mm. Interorbital
2.0 mm., anterior margin of sucking disk crenulated, the units not stalked. The
nasal cirrus small, expanded, doubly pointed. Opercular cleft extending up to

REVISION OF AMERICAN CLINGFISHES — SCHULTZ 77

base of 5th pectoral ray, tin' fold behind it complete, with a distinct lobe below.
The origin of the dorsal midway between tip of snout and of tip of caudal. The
anterior teeth above simple, sharp-pointed, circular in cross-section, I think.
Three pairs anterior teeth below flattened, not truncate, but leaf-shaped or even
a little more spatulate.

Body pretty uniformly covered with dark chromatophore.s at an average distance
from one another of twice their diameter . . .

Material examined. — Bahama Islands: U.S.N.M. No. 41733 (type
of hacres). Cuba : U.S.N.M. No. 37.->31, 5 specimens. Tkxas : U.S.N.M.
No, 1219G2, 1 specimen. British Honduras: U.S.N.M. No. 91816,
1 specimen. Locality unknown: U.S.N.M. No. 34442, 1 specimen.

Range. — Bahama Islands and West Indies; Texas and British
Honduras.

GOBIESOX ADUSTUS Jordan and Gilbert

GobiesoT adustus Jobi>.\n and Gilbert, Proc. U. S. Nat. Mus., vol. 4, p. 360, 1882
(Mazatlfin) ; vol. .">, p. 627, 1883 (Central America). — .Joudan and Evebmann,
Kep. U. S. Comm. Fish and t^sh. for 1805, App., p. 491, 1896 (Mazatian);
U. S. Nat. Mus. Bull. 47, pt. 3, p. 2334, 1898 (Mazatlfin; Pacific coa.st of
Mexico. — JoKUAN, Evermann, and Clakk, Rep. U. S. Comm. Fish, for 1928,
pt. 2, p. 489, 1930 (Pacific coast of Mexico).

Material examined. — U.S.N.M. No. 29249 (3 types of adustus).^
Mazatlan, C. H. Gilbert.
Range. — Mazatlan, Mexico.

GOBIESOX FUNEBRIS Gilbert

Gohuxox lunehrxH Giijiert, Proc. U. S. Nat. Mu.s., vol. 13, p. 95, 1890 (Puerto
Kefugio, Angel Island, and La Paz, Gulf of California). — Jordan and
E\xbmann, Rep. U. S. Comm. Fish and Fish, for 1895, App., p. 491, 1896
(Gulf of California) ; U. S. Nat. Mus. Bull. 47, pt. 3, p. 2334, 1898 (Angel
Island and La Paz, Gulf of California). — Jordan. Evermann, and Clark,
Rep. U. S, Comm. FisJi. for 1928, pt. 2, p. 4.S9, 1930 (Gulf of California).—
Bredrr, Bull. BiiiKham Oceanogr. Coll., vol. 2, art. 3. p. 48, 1936 (Puerto
Refupio) .

Rem/irkf(. — This species may be recofjnized by its small disk, the at-
tachment of the gill membranes opposite the bases of the fifth or
sixth pectoral fin rays, and the poor development of the fleshy pad on
otitor pectoral fin base, along with its narrower head.

Material examinrd. — (julf of California: U.S.N.M. Nos. 44378
(type of funebris). 119720, 124955 (2 cotypes of funehris), totaling 4
specimens; F.M.N.H. No. 8997, 2 specimens.

Range. — Gulf of California.

u I eovmNiii NT miNTiNs orricti t*4«

PROCEEDINGS OF THE UNITED STATES NATIONAL MUESUM

issued hlik^ V'gl^l hy the

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM

Vol. % Waihington: 1945 No. 3188

NEW BEETLES OF THE FAMILY EUCNEMIDIDAE FROM
CENTllAL AMERICA AND THE WEST INDIES

By W. S. Fisher

During the process of rearranging the American species of the
family Eucnemididao (Coleoptera) in the United States National
Mu>eum, a number of new genera and species were found. These are
described herein. According to Fleutiaux (Rev. Franc. Ent., voL 2,
p. 1, 1935), the name Eucnemididae sliould be used for this family in-
stead of Melasidae.

Genus TEMMLLUS Bonvouloir

TEMNILLUS ASITRICOLUS, new species

Oval-oblong, dark reddish brown, the antenna sliglitly paler, opaque,
glabrous.

Head in-egiilarly convex, with an irregular, deep, transverse depres-
sion on vertex and a very deep, irregular depression between antennae,
coarsely, deej)ly, densely punctate, the intervals vaguely granulose;
clypeus short, wide, anterior margin strongly, obtusely toothed at
middle, broadly, arciiately cniarginaU' on each side. Antenna not
exieiidiiig to base of j)roiiotum, comjiact ; first segment large, irregular,
slightly flattened, as long as following five segments united, with a
sharp tooth on underside at apex; segments -2 to 10 wider than long,
each with two round (lej)ressions on npi)er surface; segment 11
obliquely truncate at apex.

PionotMHi distinctly widei- than long, strongly conv«'.\, with a inir-
Tow, longit ndinal, median gi-oove extending fi'oin base to apex and a
small, deeper depression at middle of groove; sides nearly parallel,
slighth' sinuate along basal two-thirds, bioadly ronnded anterinily ;
base truncate, posterior angles strongly piojecting backward ; surface
r.'jdiijii tr. ~,j

80 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

coarsely, asperately punctate at middle, ocellate-punctate toward sides.
Scutellum square, coarsely j)unctate at middle, broadly rounded at
apex.

Elytra very strongly convex, strongly striate, the striae deeper near
apex ; sides parallel from bases to apical third, then arcuately narrowed
to apices, which are conjointly obtusely angulated; surface densely,
finely rugose and finely punctate, the striae coarsely, deeply punctate.

Abdomen beneath coarsely, densely, uniformly punctate, last ster-
nite with two round, deep depressions; prosternal process narrow,
coarsely punctate, rounded at apex, sides nearly parallel; posterior
coxae triangular.

Length 9.5 mm., width 3.25 mm.

Ty])e locality. — Verdant Vale, Arima, Trinidad, British West In-
dies.

T'y^e.—U.S.N.M. No. 57163.

Remarks. — Described from a single specimen collected at the type
locality.

This species resembles Temnillus Uprieuri (Guerin) but differs
from that species in being more elongate and in having the head very
deeply depressed between the antennae and on the vertex, with the
surface more finely punctured, the anterior margin of the clypeus
strongly toothed at the middle, the pronotum longitudinally grooved
with a distinct depression at the middle, and the disk asperately punc-
tured, the elytra strongly striate with the intervals finely rugose and
the scutellum and prosternal process coarsely punctured.

NEODIAPODIUS, new genus

Body elongate, subcylindrical, slightly attenuate posteriorly. Head
large, moderately convex, boardly, transversely flattened behind clyp-
eus; labrum invisible; eyes partially covered by prothorax; clypeus
moderately contracted at base. Antenna short, extending slightly
beyond base of pronotum, slightly serrate in male, strongly serrate
in female ; second segment very small ; third segment slightly shorter
than first ; segments 4 to 10 slightly shorter than third and subequal in
length to one another; segment 11 narrowly oblong, acute at apex.
Palpi abnormal, last segment of labial and maxillary palpi in male
broadly rounded at base, nearly four times as long as wide at base,
one-third as wide at apex as at base, with the outer margin slightly
concave and inner margin slightly convex, in the female short, broadly
oblong, twice as long as wide at middle, with outer margin obliquely
trmicate on apical half and inner margin strongly convex, the outer
surface with a round depression at middle. Pronotum as long as wide,
rounded and slightly sinuate in front, but not carinate ; lateral mar-
gins distinct, entire; posterior angles strongly projecting backward.

NEW EUCNEMIDID BEETLES — FISHER 81

Sciitelluni elongute-tritinguliir, gibbose autorioily, .sli<i:hlly emarginute
at apex. Elytra strongly convex, attenuate posteriorly, distinctly stri-
ate. Propleural triangle as long as ^vide at base; antennal groove
along lateral margin shallow, not very wide, smooth along outer mar-
gin, not margined internally. Marginal carina and prosternal suture
converging anteriorly, the latter closed, carinate and strongly ele-
vated anteriorly. Metasternal einmeron visible. Metasternal epi-
tternum broad, sides parallel. Posterior coxae broad, sides parallel.
Abdomen convex; last sternite slightly gibbose toward apex, which
is produced into a short, broad, truncate projection. Femora com-
pressed. Tibiae subcylindrical. Tarsi shorter than tibiae, without
lamellae; first segment slightly shorter than following segments
united; fourth segment hollowed out on dorsal surface for receiving
following segment, truncate at apex; tarsal claws robust, cleft near
apices.

Genotype. — Neodiapodms buscki new species.

This genus is allied to Diapodius Bonvouloir, but it differs from the
description given for that genus in having the pronotum not longer
than wide, the propleural triangle as long as wide at the bottom, the
metasternal epimeron visible, the clypeus sinuate in front, the meta-
sternal episternum broad with the sides parallel, the posterior tarsi
shorter tlian the tibiae, the first segment of the posterior tarsus slightly
shorter than the following segments united, the tarsal claws cleft near
the tips, and the male antenna only slightly serrate.

NEODIAPODIUS BUSCKI. new species

MaJr. — Elongate, strongly convex, uniformly reddish brown, with
the legs and palpi slightly paler, subopaque. rather densely clothed
with very short, recumbent, whitish hairs.

Head coarsely, confluently, rugosely punctate, without a longitudinal
■ iirina; interocuiar carinae slightly elevated, not extending beyond
antonnal sockets; clypeus at base twice as wide as distance between it
and eye, in front broadly rounded at middle and sinuate on each side,
surface longitudinally carinate at middle.

Pronotum stroi^gly convex anteriorly, depressed along base, witli a
more or les.s distinct longitudinal, median depression extending from
base to near apex; sides parallel along basal two-thirds, arcuatcly
narrowed near aj>^x; surface coarsely, confluently. rugoselv punctate.

Elytra with sides parallel to middle, then gradually narrowed to the
lips, which are slightly separated and acute; surface rather deei)ly.
longitudinally striate; intervals slightly convex, densely, finely granu-
losa' basally, becoming finely, sparsely punctate toward apices.

Body beneath densely punctate on abdomen, more coarsely and
deeply on i)rostei nuni, mesosternum, and melasternum; last abdominal
sternite coarsely granulosc toward apex.

82 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se

Length 14 mm., width 4.5 mm.

Female.— T>\S.Qvs from the male in being larger, and in having the
abdomen more densely clothed at the middle with more conspicuous,
longer, erect hairs, and the last sternite more obliquely narrowed toward
the apex, the tarsal claws cleft farther from the tips with the inner
tooth of each claw much shorter than the outer one, and the palpi and
antennae of different shapes.

Length 17-20 mm., width 5.5-6 nun.

Type locality --Lia, Chorrera, Panama.

Type, allotype, and paratypes. — U.S.N.M. No. 57164.

Remarks. — Described from five specimens (1 male type) collected
at the type locality by August Busck, the male type on May 14, 1912,
and four females on May 17, 1912.

Genus DROMAEOLUS Kiesenwetter

DROMAEOLUS PULCHER, new species

Oblong, slightly convex above, moderately shining, dark brown,
elytra and abdomen more reddish, legs yellowish brown, the elytra
ornamented with whitish pubescent designs.

Head slightly convex, vaguely, transversely depressed in front of
clypeus, with a vague, short, longitudinal, smooth, median carina,
finely, confluently punctate, sparsely clothed with moderately long,
semierect, whitish hairs; interocular carina very strongly elevated,
not interrupted and broadly rounded at middle ; clypeus very narrow
at base, one-half as wide as distance to eye, broadly sinuate at apex,
broadly depressed near anterior margin and with a strongly elevated,
short, longitudinal carina at base. Antenna rather robust, densely
pubescent, longitudinally carinate, not distinctly serrate; third seg-
ment vaguely longer than second; segments 3 to 10 as wide as long
and subequal in length to one another; segment 11 oblong, acute
at apex.

Pronotum slightly shorter at middle than wide at base, strongly
convex, deflexed along base; sides arcuately converging from bases
to apices; posterior angles strongly projecting backward along sides
of elytra; surface densely, finely, uniformly punctate, densely
clothed along sides and base with long, recumbent, whitish hairs,
hairs on median part not conspicuous.

Elytra slightly convex ; sides converging from bases to tips, which
are conjointly broadly rounded; surface not striate, densely, finely
punctate, densely clothed with long, semierect, blackish hairs, and
each elytron ornamented with long, recumbent, whitish hairs as fol-
lows : A narrow fascia along base broadly expanded at humeral angle,
a narrow band along sutural margin connected to a broad fascia at
middle, and a broad fascia at apex.

NEW EUCNEMIDID BEETLES — FISHER 83

Body beneath finely, densely punctate, rather densely clotlied %vith
moderately long, recuini)ent, whitish haii-s; antennal grooves mar-
ginal, narrow, equal in width for their entire length; prosternal su-
ture not deeply grooved ; posterior coxae strongly expanded internally.

Length 7.5 mm., width li.o mm.

Type local it y. — Portobelo, Panama.

fypc.—V.SS.^l. No. :.71G5.

ReiJiarkM. — Described from a single specimen reared from a pupa
collected in the bark of an unrecognized tree at the type locality,
^rarch 1911, by E. A. Schwarz.

This species resembles Dromaeolus ornatidu-s Horn but differs from
that species in being larger, in having the elytra and abdomen reddish
brown and the clypeus carinate at the base, and in not having the white
j)ubescence extending along the lateral margins of the elytra or the
elytra striate.

DROMAEOLUS PANAMENSIS, new species

Oblong, moderately convex, slightly shining, dark brown, the legs
and antenna yellowish brown, rather densely, uniformly clothed
above with short, semierect, brownish hairs, and beneath with short,
recumbent, yellowish hairs.

Head nearly flat, densely, coarsely ocellate-punctate, slightly' de-
pressed near interocular carina, which is strongly elevated, arcuate,
and not interrupted at middle; clypeus very narrow at base, one-
third as wide as distance to eye, not carinate, broadly rounded at
apex, the surface coarsely, confluently ocellate-punctate. Antenna
rather robust, densely pubescent, vaguely, longitudinally carinate]
compact, not serrate; third segment slightly longer than second; seg-
ments 3 to 10 slightly longer than wide and subequal in length to
one another; segment 11 oblong, subacute at apex.

Pronotum distinctly wider than long, strongly convex, deflexed
along ba.se, without distinct depressions or carinae; sides arcuately
converging from ))ases to apices; surface; densely, coarsely ocellate-
punctate.

Klytra moderately convex; sides converging from bases to tips,
which are conjointly broadly rounded; surface finely sti'iate, intervals
flat, densely, finely rugose basally, finely, sparsely, punctate toward
apices.

I'ody beneath finely, densely punctate on abdomen, more coarsely
ocellate-punctate on j)rosternum, mesostennnn. and metasternum ;
nntennal grooves marginal, narrow, (h\\\'a\ in width for their entire
length; j)rosternal suture nr)t deeply grooved; pi-oj)leural triangle
with iK>sterior margin shorter than inner nnirgin; posterior coxae
.strongly expanded internally.

Length 3.4 mm., width 1.3 mm.

Tyi>r locality. — Cabima, Pnriania.

84 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Type.—U.S.^M. No. 57166.

Remarks. — Described from a single specimen collected at the type
locality on May 28, 1911, by August Busck.

This species is allied to Droinaeoliu moerens Horn but differs from
the description given for that species in being dark brown, with the
legs and the antennae yellowish brown, and in having the third seg-
ment of the antenna subequal in length to the fourth, the elytra rugose
at the bases, and the pronotum distinctly wider than long.

Genus FORNAX Laporte

FORNAX POEYI, new species

Elongate, subcylindrical, strongly convex, slightly narrowed
posteriorly, moderately shining, reddish brown, anterior margin of
pronotum more reddish, and legs and antennae brownish yellow,
rather densely, uniformly clothed with short, recumbent, yellowish
hairs.

Head strongly convex, without depressions or carina, coarsely,
confluently ocellate-punctate, and more or less granulose ; interocular
carina strongly elevated, not extending on base of clypeus; clypeus
at base slightly wider than distance to eye. Antenna slender, ex-
tending slightly beyond base of elytra, not distinctly serrate ; segment
2 subequal in length to segment 4; segments 4 and 5 united longer
than 6; the following segments subequal in length to one another;
segment 11 oblong narrowly rounded at apex.

Pronotum slightly wider than long ; sides nearly parallel from base
to apex; posterior angles acute; disk moderately convex, without
depressions or median carina ; surface rugose and finely, confluently
ocellate-punctate.

Elytra moderately convex; sides parallel to behind middle, then
gradually, arcuately narrowed to tips, which are conjointly narrowly
rounded; surface finely striate, finely, densely granulose basally, be-
coming more sparsely, finely punctate toward apices; epipleura
punctate, not grooved.

Body beneath rather densely punctate, finely on abdomen, coarsely
on prosternum, and rugosely on metasternum ; last abdominal sternite
coarsely granulose toward apex; antennal groove narrowed by eye,
deep, not half as wide as propleural triangle, not wider in front, the
inner margin not very sharply defined; prosternal process flat, sides
nearly parallel to behind coxae, acute at apex; first segment of
posterior tarsus subequal in length to the following segments united.

Length 4.5 mm., width 1.25 mm,

T]/pe locality. — Cayamas, Cuba.

Type.— U.S.^.M. No. 57167.

NEW EUCiSTEMIDID BEETLES — FISHER 85

Rt7narks. — Described from u sin<»:le speciiutMi collected at tlie type
locality, January 24, by E. A. Schwarz.

This species is allied to Fornax hdd'nis (Melsheimer) l)ut (Hirers
from that species in bein«j: smaller and in havin<ij the head and pronotum
finely, continently punctured, and the inner mar*rin of the antennal
suture on the prosternum not very sharply defined.

FORNAX VALFRIO, new species

Oblong, subcylindrical. moderately convex, slightly narrowed pos-
teriorly, uniformly dark, reddish brown, the legs and antennae slightly
paler, densely clothed above with moderat^ely long, recumbent and
semierect, brownish-yellow pubescence, which does not conceal the
surface, beneath with shorter and less conspicuous pubescence.

Head convex, without depressions or longitudinal carina, finely, con-
Ihienlly punctate, finely rugose; interocular carinae slightly elevated,
not extending on base of clypeus, the latter at base slightly wider than
distance to eye. Antenna slender, extending to posterior coxa, not
serrate; segments 4 to 10 similar, becoming gradually longer toward
t ip of antenna ; segment '2 very small, shorter than 4; segment 3 longer
than 4.

Pronotum wider than long; sides parallel posteriorly, arcuately
narrowed anteriorly; posterior angles subacute; disk strongly convex
anteriorly, transversely, obliquely depressed behind middle, with a
short, smooth line in front of scutellum, and a distinct pitlike depres-
sion along base on each side at outer angle of scutellum ; surface finely,
densely granulose. finely, obsoletely I'ugose anteriorly and towaid
Literal margins.

Elytra moderately convex; sides ])arallel to apical third, then
slightly narrowed to the tips, which are conjointly broadly rounded,
distinctly striate; intervals flat, finely, densnly granulose basally, be-
coming punctate po.steriorly ; epipleurae smooth, grooved their entire
length.

Body beneath finely, densely punctate, more coarsely on prosterniini :
antennal groove narrow'ed by the eye, deep, not half as wide as pro-
I)leural triangle, not wider in front, innei- margin sliar|)ly <lelined;
j»ro>t('rnai pi-ocess tleflexed and abruptly narrowed i)ehind anterior
coxae, acute at apex; last abdominal sternite longilu'linally depressed
along sides, longitudinally conipressetl ])ostei'iorIy, scabrous and acute
at apex: first segment of posterior tarsus >Mbe(iual in length to the
following .segments united.

Length Il-IT) mm., width .'{.r>-4.r) mm.

Tijpr locdlitu. — Chitaria, Cost a Ricii.

Type and paratijpcs.—\] .f^.^ M. No. .57108.

Rrrn/irl's. — Described fi-om thi-cc sp(>cime)is (1 typ(>) collected nt
the type locality. December 4, HOO, by M. Valerio.

86 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

This species resembles Fornax mendax Bonvouloir and Fornax
ohi'utus Guerin, but it differs from both these species in having the
pronotum transversely, obliquely depressed behind the middle, with a
distinct pitlike depression along base on each side at outer angle of the
scutellum, and with the surface finely granulose, and the elytra finely
granulose basally, and it also differs from ohrutus Guerin in having the
prosternal process deflexed behind the anterior coxae, and abruptly
narrower to the tip, which is acute.

Genus PLESIOFORNAX Bonvouloir

PLESIOFORNAX NIGRINUS, new species

Narrowly elongate, subcylindrical, strongly shining, black, with a
faint bluish or greenish tinge, and legs brownish black, sparsely
clothed with short, recumbent, inconspicuous hairs.

Head strongly convex, without distinct depressions or carina, rather
coarsely, densely punctate ; interocular carina not elevated or extend-
ing along base of clypeus, which is strongly contracted at base, its
width there being less than half the distance to eye, and broadly, sin-
uately rounded in front. Antenna slender, more robust toward tip,
extending to basal fourth of elytra, basal segments slightly longitu-
dinally carinate; segment 2 as long as segment 5; segment 3 much
longer than either 2 or 4 ; segment 4 longer than wide, more than half
as long as 5, the following segments becoming gradually longer.

Pronotum as long as wide, strongly convex; sides parallel along
basal two-thirds, feebly, arcuately narrowed along apical third ; sur-
face very sparsely, finely punctate, with a median, longitudinal de-
pression extending from base to middle of pronotum, the depression
becoming broader and deeper posteriorly.

Elytra strongly convex; sides parallel from base to apical third,
then arcuately narrowed to tips, which are conjointly broadly
rounded ; surface rather coarsely, not very densely, irregularly punc-
tate, not striate.

Abdomen finely, densely punctate, more confluently toward apex
of last sternite, which is obtusely angulated. Presternum sparsely
punctate on median part, densely on propleural triangle, which is
distinctly longer than wide at base ; antennal groove deep, smooth, and
shining in its entire length; prosternal process broad, flat, without
depressions, sides obliquely converging to apex, which is broadly
rounded. Posterior tarsus with first segment as long as following
segments united ; tarsal claws long, slender, slightly swollen near base,
but not toothed.

Length 11-13 mm., width 2.5-2.8 mm.

Type locality. — Rio Hondo, Costa Rica.

Type and paratypes.—V.S.^M. No. 57169.

NEW EUCNEMIDID BEETLES — FISHER 87

Remarks. — Described from three specimens (1 type). Type and
one paratype collected at the type locality by J. Pittier, and one para-
type collected at San Jose, Costa Rica, 1,000 to 1,200 meters, January
15, 1934, by Ferdinand Nevermann.

This species is allied to Ple^iofornnx loTigicorms Horn but differs
from the description given for that species in being shining black with
a bluish or greenish tinge and in having the antenna extending only
(o the basal fourth of the elytra, the pronotum as long as wide, with
the sides parallel along the basal two-thirds, the proi)leural triangle
densely punctate, and the first segment of the posterior tarsus as long
as the following segments united.

Genus FARSUS Jacquelin Du Val

FARSUS CONVEXUS, new species

Cylindrical, robust, strongly convex, scarcely narrowed posteriorly,
subopaqup. uniformly reddish brown, sparsely, uniformly clothed witli
-hort, semierect, yellowish hairs, which are more recumbent on under-
side of body.

Head moderately convex, vaguely depressed behind clypeus, coarsely,
confluently ocellate-punctate ; interocular carina vaguely elevated', not
extending along base of clypeus; clypeus strongly contracted at base,
which is nearly as wide as distance to eye, broadly sinuate or rounded
in front. Antenna rather robust, extending slightly beyond base of
elytra; segment 2 as long as 4, which is one-half as long as 3, the fol-
lowing segments subequal in length to one another except 11, which is
longer and oblong; segments 4 to 10 compact, wider than long.

Pronotum sliglitly wider than long, strongly convex anteriorly, ob-
liquely deflexed along base; sides parallel to near apex, then arcuately
rounded ; anterior margin broadly rounded, distinctly carinate, the
cai'ina curving backward on each side near apical angles; base broadly
emarginate in fiont of scutellum; surface coarsely, densely ocellate-
punctate at middle, scabrous toward sides.

Elytra strongly convex, vaguely striate, the intervals rather densely,
irregularly punctate; sides parallel to near apices, which are conjointly
broadly rounded.

Body beneath densely, coarsely ocellate-punctate; abdomen strongly
convex, last sternite broadly rounded at apex. Prosterninn with
lateral margin strongly elevated posteriorly, obsolete toward apex;
supplementary carina sinuate, strongly elevated anteriorly, extending
along j)rosternal suture to middle of proplenral triangle. Femora ro-
i;usf. P<;sterior coxae strongly, angularly expanded internally.

Length 4-6 mm., width 1.25-2 mm.

Type lorol'ity. — Tabernilla, Caiutl Zone.

Type and jmratyprn. — IJ.S.N.M. No. 57170.

88 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Remarhs. — Described from four specimens (1 type). The type and
two paratypes collected at the type locality, May 14-17, 1907, by August
Busck, and one paratype taken from an arc-light globe at Ancon,
Canal Zone, during April 1911. Under this species is also included a
specimen collected at General Ballivian, Salta Province, Argentina,
during 1927-28, by G. L. Harrington. This specimen differs from the
type only in having the supplementary carina on the prosternum
vaguely indicated posteriorly and joined to the lateral carina near the
base.

This species is allied to Farsus oblitus Horn but differs from that
species in being more robust, more convex, and scarcely narrowed
posteriorly and in having the antenna shorter, with the segment 2 as
long as segment 4, and the following segments wider than long, the
pronotum coarsely ocellate-punctate at the middle, the sides of the
elytra parallel to near the apices, with the surface irregularly punc-
tate, the femora robust, the posterior coxae strongly, angularly ex-
panded internally, and the supplementary carina on the prosternum
strongly elevated anteriorly and extending only to the middle of the
propleural triangle.

^ Genus ARRHIPIS Dejean

ARRHIPIS CUBANUS, new species

Elongate, subcylindrical, moderately convex, rather strongly shin-
ing, reddish brown, the antenna and legs yellowish brown, rather
densely, uniformly clothed with short, semierect, whitish hairs.

Head slightly convex, without depressions or carina, coarsely,
deeply, confluently punctate; interocular carina slightly elevated,
extending around inner margin of antennal fossa, but not along
base of clypeus ; clypeus short, at base twice as wide as distance to eye,
feebly, broadly rounded in front. Antenna moniliform, slightly thick-
ened toward apex, extending to base of pronotum ; segment 3 as long
as following two segments united ; segments 4 to 10 as wide as long,
subequal in length to one another; segment 11 oblong, feebly incised
on inner margin, narrowly rounded at apex.

Pronotum as long as wide, widest near apex ; sides nearly parallel ;
anterior margin broadly rounded, the carina extending nearly to lat-
eral carinae; disk slightly flattened at middle, transversely deflexed
along base ; surface densely, coarsely ocellate-punctate at middle, more
densely punctate and rugose toward sides.

Elytra moderately convex; sides parallel from bases to behind
middle, then arcuately narrowed to the tips, which are conjointly
broadly rounded and terminating into a short spine; surface feebly
striate, finely rugose, rather densely, coarsely punctate basally, becom-
ing coarsely asperate toward apices.

NEW EUCNEMIDID BEETLES — FISHER 89

Abdomen beneath finely, sparsely punctate; last sternite coarsely,
densely punctate at apex, with a transversely oval, densely granulose
area. Prosternuni, mesosternuni, and nietasternum rather densely,
coarsely ocellate-punctate; antennal groove absent but replaced ante-
riorly by a smooth, shining space. Metasternal episternum narrow,
sides parallel. Posterior coxae large, strongly, triangularly expanded
internally.

Length 4.25 mm., width 1.25 mm.

Type locaTit}/. — Cayamas, Cuba.

7V;>e.— U.S.N.M. No. 57171.

RcmwA'-f. — Described from a single male collected at the type local-
ity, March 6, by E. A. Schwarz.

This species is allied to Arrhipis lanieri Guerin but differs from that
species in being smaller and of a pale reddisli-brown color, and in hav-
ing the hairs on the elytra shorter, denser, and more uniformly dis-
tributed, the outer segments of the antenna compact and as wide as
long, and the prosternum flattened in front with the prosternal proc-
ess triangular between the anterior coxae and knife-shaped behind the
coxae.

ARRHIPIS INSULARIS, new species

Elongate, subcylindrical, moderately convex, strongly shining, uni-
formly yellowish brown, antennae and legs slightly paler, sparsely
clothed witli very short, recumbent, inconspicuous hairs.

Head slightly convex, without depressions or carina, coarsely,
rather densely, but not deeply, ocellate-punctate; interocular carina
vaguely elevated, extending around inner margin of antennal fossa;
clypeus shoit, at base twice as wide as distance to eye, broadly rounded
in front. Antenna moniliform, slightly thickened toward apex, ex-
tending slightly beyond base of pronotum; segment 3 nearly as long
as following two segments united ; segments 4 to 10 vaguely wider than
long, suboqual in length to one another; segment 11 oblong, narrowly
rounded at apex.

Pronf)tum slightly wider than long, widest near apex; sides nearly
))aiallel; anterior margin broadly rounded, the carina extending
nearly to lateral margins; disk convex, not flattened at middle,
arcuately dcflexed along base; surface coarsely, rather densely ocel-
late-punctate at middle, more conflucntly punctate toward si(h's. inter-
vals vaguely granulose.

Elytra modei'ately con\c\; sides j);ii!iIK'l fioni bases to behind
middle, then arcuatx^ly narrowed to the tips, which are conjointly
broadly rounded nnd terminating into 'i short spine; surface not
striate, coarsely, rather d<'nsely jiunctate, slightly asperate toward
apices, the intervals vaguely granulose.

Abdomen beneath sparsely, very coarsely puncUite; last sternite
with a transversely oval, densely granulose area. Prf)stermini. meso-

90 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

sternum, and metasternum rather densely, coarsely punctate ; antennal
groove absent but replaced by a smooth, shining space. Metasternal
episternum very narrow, wider behind than in front. Posterior coxae
not strongly expanded internally.

Length 3 mm., width 0.9 mm.

Type locality. — Cayamas, Cuba.

Ty^e.— U.S.N.M. No. 67172.

Remarks. — Described from a single male collected at the type lo-
cality, March 15, by E. A. Schwarz.

This species is allied to Ai-rhifis cubcmus Fisher but differs from
that species in being yellowish brown, with very short, inconspicuous
hairs, and in having the pronotum convex at the middle and not
flattened, the outer segments of the antenna vaguely wider than long,
the metasternal episternum very narrow, and wider behind than in
front, the prosternal process triangular behind the anterior coxae, the
pronotum arcuately deflexed along the base, and in not having the
elytra striate.

Genus DIRHAGUS Laporte

DIRHAGUS ALBOFASCIATUS, new species

Elongate, subcylindrical, moderately shining, reddish brown, except
legs, which are yellowish brown, the pronotum and elytra ornamented
with whitish pubescence.

Head strongly convex, vaguely depressed near clypeus, without
longitudinal carina, densely, finely granulose, rather densely clothed
with short, erect, inconspicuous, whitish hairs; interocular carina
elevated, broadly interrupted at middle; clypeus at base one-half as
wide as distance to eye, truncate at apex; eyes large, not incised
beneath. Antenna about two-thirds as long as body, strongly pecti-
nate from segment 4 in male, acutely serrate from segment 3, except
segment 11, which is narrowly elongate, in female; segment 3 much
longer than 2.

Pronotum distinctly wider than long, strongly convex, a short,
longitudinal, antescutellar carina with a slight depression on each
side ; sides parallel ; anterior margin arcuately rounded, the anterior
supplementary carina strongly elevated, short, not extending to middle
of pronotum ; surface densely, finely ocellate-punctate, sparsely clothed
at middle with short, erect, inconspicuous hairs, and at sides and
toward base with long, recumbent, whitish hairs.

Elytra moderately convex ; sides slightly converging from bases to
tips, which are very broadly conjointly rounded; surface vaguely
striate along sutural margins near apices, densely, coarsely, rugosely
punctate, sparsely clothed with short, semierect, inconspicuous hairs,
and irregularly clothed with longer, recumbent whitish hairs along

NEW EUCNEMIDID BEETLES — FISHER 91

buses and skies uiul forinin<r a more or less distinct broad, transverse
fascia behind middle and a broad spot at apices.

Body beneath coai*sely, densely punctate, sparsely clothed with
short, recumbent, whitish hairs; juxtasutural jjroove broad, smooth,
sharply limited externally; lateral marj^inal carina sinuate, obsolete
posteriorly and curving toward posterior supplenlentary carina, which
i< stronuly elevated posteriorly, becoming: obsolete in front of middle;
metasternal episternum narrow, wider behind than in front; posterior
coxae slightly expanded internally.

Length 3 mm., width 1 mm.

Type locality. — Felton, Cuba.

Type^ allotype., aiid paratypes. — U.S.N.M. No. 57173.

Remnrl-a. — Described from five specimens (1 male typo). The type
and allotype were collected at the type locality by W. M. Mann; two
paratypes were collected at Baracoa, Cuba, during August 1901. by
August Busck; and one paratype was collected at Higueral, Domini-
can Republic, during February 1916, by E. G. Smyth.

This species is allied to Dirhagus pectinatus LeConte but differs
from that species in being smaller and in having the pubescence on the
pr(;notum and elytra longer and ornamented with more or less dis-
tinct white pubescent designs, the juxtasutural groove smooth, with
the sides parallel, the posterior supplementary carina extending to
the middle of the pronotum, the antenna in the male pectinate from
the fourth segment, and in not having the head longitudinally carinate
on the occiput.

HYLOTASTELLA, new genus

Body elongate, parallel. Head large, strongly convex; labrum in-
visible; eyes partially covered by prot borax; clypeus strongly con-
tracted at base. Antenna long, robust, cylindrical, segments compact ;
second segment very small, the following segment much longer than
wide. Pronottmi wider than long, lateral margins distinct, entire;
anterior margin simple, not carinate. Scutellum subtriangidar,
longer than wide, narrowly truncate at apex. Propleural triangle
longer than wide, without antcnnal grooves. Marginal carina and
prosternal suture of pro.sternum converging anteriorly, prostenial su-
ture closed, distinct for entire length. Motastermun and abdomen
without tai-sal grooves. Metastenial cpimcron completely covcicd.
Metasternal episternum nairow. sides jiaiallel. \j\\^\ abdoininal ter-
gite not extending over last alxlomiiial sternite. which is nai'rowlv
rounded at ajn-x. Posterior coxa smldenly exjiaiided internally, very
narrow externally. Logs slendei-. Tai>i withoni laniellae; segment
1 of posterior pair as long as following segments unile(l; segment 4
hollowed out on doi-sal surface for receiving following segment, trun
cate at apex.

92 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Genotype. — HylotasteUa schwarsi, new species.

This genus resembles a small species of Hylotastes Bonvouloir, but
it differs from that genus in being gradually narrowed posteriorly
and in having the antenna cylindrical and four-fifths as long as the
body, the clypeus broadly rounded in front, the posterior coxa
abruptly expanded internally, the propleural triangle longer than
wide, the tips of the elytra conjointly broadly rounded, and the last
abdominal tergite not extending over the last abdominal sternite.

HYLOTASTELLA SCHWARZI, new species

Narrowly elongate, moderately convex, dark brown, slightly paler
beneath, with the humeral angles of elytra and prosternum yellowish ;
body above subopaque, beneath slightly shining ; head and pronotmn
densely clothed with long, recumbent, golden-yellow hairs, the hairs
shorter and less conspicuous on elytra and underside of body.

Head strongly convex, without depressions or longitudinal carina,
finely, confluently ocellate-punctate ; interocular carina strongly ele-
vated, interrupted at middle ; clypeus at base one-half as wide as dis-
tance between it and eye, broadly, sinuately rounded at apex. An-
tenna four-fifths as long as body ; segments 3 to 5 subequal in length
to one another, the following segments becoming gradually longer
to tip of antenna.

Pronotum distinctly wider than long; sides parallel posteriorly,
arcuately narrowed anteriorly; disk strongly convex, transversely,
narrowly depressed along base, with a distinct round depression on
each side in front of middle; surface finely, confluently ocellate-
punctate similarly as on head.

Elytra with sides parallel to apical fourth, then arcuately narrowed
to the tips, which are conjointly broadly rounded; surface finely, dis-
tinctly striate, the intervals finely, densely granulose.

Body beneath very finely, densely punctate ; last abdominal sternite
slightly gibbose and coarsely granulose toward apex.

Length 5.75-7 mm., width 1.5-2 mm.

Type locality. — Portobelo, Panama.

Type and paraiypes. — U.S.N.M. No. 57174.

Remarks. — Described from three specimens (1 type) collected at
the type locality, the type and one paratype collected March 1, 1911,
by E. A. Schwarz, and one paratype collected February 20, 1911, by
August Busck.

Genus NEMATODES Berthold

NEMATODES EXIGUUS, new species

Elongate, subcylindrical, moderately convex, strongly attenuate
posteriorly, slightly shining, dark brown, nearly black, except legs
and antennae, which are reddish brown, rather densely, uniformly

NEW EUCNEMIDID BEETLES FISHER 93

clothed with short, recumbent, yollowisli hairs, which arc finer on
underside of body.

Head rather strongly convex, vaguely depressed near clypcus,
coarsely, densely ocellate-punctate; intcrocular carina scarcely ele-
vated, not extending along inner margin of antennal depression;
clypeus slightly narrower at basa than distance to eye, strongly sinuate
at apex. Antenna robust, slightly expanded toward apex, extending
to base of elytra; segment '2 slightly longer than 4; segment 3 longer
than 6 and subequal in length to 4 and 5 united ; segments 7 to 9 slightly
longer than wide : segment 11 oblong, acute at apex.

Pronotum slightly wider than long, widest near middle; sides
vaguely narrowed posteriorly, arcuately narrow^ed anteriorly; pos-
terior angles acute, not carinate or divergent; disk convex, with a
vague, round dei/i-ession on each side in front of middle, sometimes
without depressions, rarely with a longitudinal median, smooth space;
surface densely, finely punctate on median part, finely rugose toward
sides.

Elytra moderately convex; sides nearly parallel along basal half,
strongly attenuate po^ttn-iorly to the tips, which are rather broadly,
conjointly rounded; surface vaguely striate, the sutural stria more
distinct, intervals flat, densely, finely punctate basally, the punctures
becoming sparser toward apices.

Body beneath finely, densely punctate, the punctures coarser on pro-
sternum ; antennal groove broad, shallow posteriorly, deeper anteriorly,
not limited internally by a carina, but sometimes with a short carina
at outer margin near eye.

Length 3.5-G mm., width 0.8-1.75 mm.

Type locdJity. — Cayamas, Cuba.

Type and paratypca. — U.8.N.M. No. 57175.

Remcn'ks. — Described from seven specimens (1 type) collected at the
type locality during PVbruary and March, by E. A. Schwarz.

This species is allied to Nnnatodes alropon (Say) but dilFcrs from
that s|)ecies in being smaller and in having the pronotum widci- Ihan
lung, with the surface densely, finely punclate at the middle, but witli-
out a longitudinal, median depressed line, and flic elylia vaguely
striate.

0. 1. soviNNiiiNr miNriNa orricc > i<49

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM

Vol. % Wwhington: 1945 No. 3189

NEW LANTERNFLTES (FULGOEOIDEA) FROM
SOUTH AMERICA

By R. G. Fennah

Descriptioxs are ^iven herein of seven new species of Fulgoroidea
in tlie cixiid genus Pintalia Stal, of a new genus and species in the
otiocerine Derbidae, and of two new species of Ateson Metcalf
(Achilidae). All the specimens mentioned, including the types, are
in the United States National Museum.

Family CIXHDAE

Genus PINTALIA Stil

Pintalia St.\i.. Svcii.ska Vet.-Akad. IImikII., vol. 3, No. 6, p. 4, 1862. (Genotjrpe,
P. lateralis St&l, dt'sigiiated by Muir, Pnn-I'acific Ent., vol. 1, pp. 103, 100,
1025.)

Tliis rapidly expanding genus is very compact. The characters by
which species may be distinguished include the shape of the carinae
of the vertex and of the margins of the frons at its base, that of the
anal segment of the male, of the pygofer, aodeagus, and genital styles,
and the color pattern on the tegmina. The natural relationship of
the included species cannot be as.sessed with precision until there is
evidence to indicate the order of seniority of the variables used in their
separation. On current evidence it would .seem that the .shape of the
male anal .segment and of the genital styles is .slightly inore stable than
the other characters mentioned, and tiie writer tentatively grouj)S as
follows those species for which the relevant data are availal)le:

620240 — 45

96 PROCEEDINGS OF THE NATIONAL MUSEUM vol. w

Anal segment short, lateroapical lobes narrow, decurved through 130°, genital
styles short, subangulate propna group

Anal segment fairly short, not or scarcely deflexed distally, genital styles expanded
and spatulate distally ecuadoriensis group

Anal segment long, distally deflexed as far as, but not exceeding, 45°, genital
styles long, narrow in ventral view albolineata group

Anal segment asymmetrical, often twisted into a single vertical lamina distally,
genital styles short obtorta group

Anal segment as above, genital styles long infuscata group

Anal segment distally deflexed as far as 45°, genital styles short, clublike.

oiTiata group

Anal segment shortly deflexed through 45°, styles short, angulate, pointed in ven-
tral view huigrensis group

Anal segment deflexed through 90° or slightly more, genital styles distally ex-
panded, truncate or concave on apical margin Mcaudata group

Anal segment as above, very asymmetrical, genital styles rounded at apical
margin vomerifera group

The propria group includes only this species; the second group
includes ecuadoriensis Muir and quadrimaculata (described below) ;
albolineata Muir, by contrast, typifies a large group including hnm-
nivenosa Muir, latinotata Muir, longispinis Muir, quadrispinosa Muir,
fuscomaculata Muir, hlairmontensis Muir, tumatumariensis Muir,
fuscipennis Muir, alhomarginata Muir, ohscurata Muir, and two
species, raamiorata and ohliquivitta^ described below; the ohtorta
group, including obtorta Muir, pulchella Muir, and fuscomarginata
Muir, would appear to be quite closely related to the infuscata group
{infuscata Muir, angustinofata Muir, and falcata (described below) )
if judged by the shape of the anal segment; omata Muir is grouped
with altamasonica Muir and discrepans Muir, huigrensis Muir is
somewhat doubtfully associated with furcata Muir, while bicaudata
Muir, maculipennis Muir, and curvivitta and daedala (both described
below) form a fairly closely knit group ; vomerifera (described below)
is placed apart, though it shares characters with the preceding group
and with the obtorta group, on the ground that the shape of the genital
styles and the general structure of the aedeagus differ considerably
from those of all the species so far considered.

PINTALIA QUADRIMACULATA, new species
PlATE 2, FiGUBES 1, 7-10

Male. — Length, 4.0 mm. ; tegmen, 5.9 mm.

. Vertex with anterior margin and transverse carina straight, lateral
margins of f rons not thickened in basal portion.

Fuscous ; genae, lateral fields of pronotum, sternum, and legs testa-
ceous. Tegmina testaceous, posterior edge of costal cell, base of cell
Sc, basal four-fifths of cell R, cell M, and the larger part of cells Cu^a
and Cuib deeply inf uscate, almost piceous, a white round spot at middle

NEW LANTERNFLIES FROM SOUTH AMERICA — FENNAH 97

of common stalk of M, a similar spot at fork of Cui, a third on Cuia
and a fourth on Cuib, a hyaline area parallel to nodal line and just
basad of it, a fuscous band overlying nodal line and a similar band of
equal width and more clearh' defined obliquely traversing apical cells
acro:ss middle from apex of K to M3, apical margin narrowly fuscous
to M3 then broadly fuscous posteriorly but infuscate area interrupted
by a large testaceous spot distad of apex of clavus.

Anal segment of male short, not deflexed; lateroapical angles only
slightly prominent, very slightly produced ventrally. Pygofer broad
■with each lateral angle produced in a short narrow lobe, distally
rounded. Aedeagus tubular, straight, with three spines arising at
approximately same level at base of flagellum, the middle spine por-
rect, the other two strongly curved. Genital styles in profile angulato
at middle with a pointed eminence on dorsal border, spatulate distally
with greatest width two-thirds from base.

Ti/pe.—V.S.^M. No. 57092.

Described from one male specimen taken at Tumupasa, Bolivia, by
W. M. Mann (December 1921). The dark basal field of the tegmina
and its ornamentation of four cretaceous spots readily distinguish
this species.

PINTALIA MARMORATA, new species

Plate 2, Fiqubes 5, 20 ; Plate 3, FiGtTBES 30-32

Male. — Length, 4.1 mm. ; tegmen, 6.0 mm.

Vertex with anterior margin and transverse carina straight, lateral
margins of frons very slightly thickened basally.

Testaceous; frons, genae, lateral fields of pronotum, and mesonotum
fuscous. Tegmina ivory-hyaline, marbled dull brown in small irregu-
lar spots, with an irregular fascia across apical cells subparallel with
apical margin; veins dull brown. Wings hyaline, veins fuscous.

Anal segment of male moderately long, deflexed through 45° distad
of anal foramen but not much produced. Pygofor broad, lateral
angles large, with dorsal margin sliglitly concave, ventral margin con-
vex. Aedeagus long, tubular, slightly curved dorsad, with three un-
equal spines at base of flagellum, one being much longer than the other
two, flagellum distally expanded, its dorsal margin contiiuied in a
slender curved spine, apex of flagellum somewhat pointed. Genital
styles strongly angulaf o, with a broad-basod, pointed eminence dorsally
near angle, gradually ex[)anded distally, apical margin obli(iuely
truncate.

Ti/pr.—V.S.yiM. No. 57093.

Described from a single male specimen collected at light on deck
in harbor, Puerta Bolfvar, Ecuador, by M. Kisliuk and C. E. Coolcy
(April 9, 19:'.2). This sj^ecies is distinguished by the pattern on tho
tegmina and by the shape of the anal segment, pygofer, and genitalia.

98 PROCEEDINGS OF THE NATIONAL MUSEUM vol. B6

PINT ALIA OBLIQUIVITTA, new species
Plate 2, Figuees 3, 16-19

Male.—Length., 4.0 mm. ; tegmen, 6.0 mm.

Vertex with anterior margin and transverse carina straight, lateral
margins of frons prominently raised but not thickened basally.

Fuscous; sternum testaceous, legs pale fuscous. Tegmina ivory-
hyaline, a broad oblique band from base of stigma to apex of clavus,
three transverse narrow bars in costal cell, the basal bar extended in a
narrow broken fascia to claval suture, a suffusion along nodal line
and two S-shaped areas in apical cells of E, and M, a pale suffusion
over distal part of apical cells, cell Cuib except for a round spot, and
two broad bands between claval suture and first claval vein yellowish
brown. Wings hyaline, veins testaceous.

Anal segment of male moderately long, distally deflexed through
60°, deflexed part not nearly so long as horizontal part. Pygofer
broad, lateral angle about 60°. Aedeagus tubular, a spine one-third
from base on left side directed ventrally and anteriorly, a second spine
on left side at base of flagellum directed dorsally and caudad, a stout
curved spine at middle of dorsal surface directed dorsally and caudad,
a longer slender spine on right side at base of flagellum directed ven-
trally and anteriorly, flagellum with two minute adpressed spines
on right side near middle, and a longer somewhat oblique spine at
apex. Genital styles angulate near base, with a short broad spine
dorsally near angle, distally narrow, slightly expanding caudad,
rounded and setigerous at apex.

Type.—V.S.l^M. No. 57094.

Described from one male taken at Santo Domingo, southeastern
Peru, labeled "collection Kosenberg." This species is distinguished
by the genitalia and by the tegminal pattern.

PINTALIA FALCATA, new species

Plate 2, Figures 4, 22 ; Plate 3, Figures 27-29

Male. — ^Length, 4.3 mm. ; tegmen, 6.1 mm.

Vertex with anterior margin and transverse carina straight, lateral
margins of frons not thickened basally.

Fuscous ; tegmina ivory-hyaline, extreme base of tegmina, a broken
fascia from costa to junction of claval veins, a second fascia from
costa just basad of stigma to commissural margin distad of apex of
clavus, nodal line, apex of each apical vein, and a band from third
apical cell of M to middle of cell Cuia yellowish brown. Wings
slightly infuscate, veins dark.

Anal segment of male angulately deflexed beyond anal foramen,
twisted into a horizontal lobe distally. Pygofer with lateral angles
broadly rounded. Aedeagus tubular, a long straight spine on left

NEW LANTERNFLIES FROM SOUTH AMERICA — FENNAH 99

side at base of fl:i^olIum, a lon<^ curved spine on right side at same
level, both directed anteriorly; llagelluni with a curved tubular mem-
branous lobo arising at middle and curving to left, a pair of broad
unequal spines distally. Genital styles weakly angulate at middle,
distal portion somewhat sinuately expanded, a row of minute denticles
on inner border distally.

Type and parafr/pe.—V.S.^M. No. 57095.

Described from one male and one female collected in French Guiana
by W. M. Schaus.

PINTALIA CURVrVITTA. new species

Plate 2, Figures 2, 11-15

Malr. — Length, 3.9 nmi,; tegmen, 5.9 mm.

Vertex with anterior margin and transverse carina straight, lateral
margins of frons not thickened basally.

Fuscous; sternum and legs testaceous. Teginina ivory-hyaline,
sparsely marked with yellowish brown on membrane, a broad approxi-
mately semicircular band of the same color widely enclosing stigma
and reaching posteriorly to Cuia; a band across apical cells of M, apex
of Cuib, a spot distad of apex of clavus, and a suffusion at middle
of claval suture yellowish brown. "Wings hyaline, veins testaceous.

Anal segment of male deflexed through 100°, symmetrical, deflexed
part almost as long as basal part. P3'gofer with lateral angles pre-
duced into a short narrow lobe, markedly sinuate at apex. Aedcagus
tubular, a short stout spine on dorsal surface one quarter from base,
directed dorsally and unti-riorly, a second spine of similar shape,
though slenderer, dorsally at middle, and a third spine, curved dorsad
and caudad, at base of flagollum, a short slender spine at middle on
left side directed ventrad and forward, a longer sinuate spine sub-
parallel to it at base of flagellum, flagellum devoid of ornamentation.
Genital styles angulatcly bent at middle, expanded distally, apical
margin very shallowdy excavate.

rv/?e.— U.S.N.M. No. 57096.

D('-<ri!)cd from a single male taken at Rurrenabaque, Bolivia, by
W. M. Mann (November 1921). This species is distinguished by tho
shaiK' of the anal segment, of the lateral angles of the pygofer and of
the genitalia, and by the pattern on tho tegmina.

PINTALIA DAEDALA, new iipcrica
Pi>ATE 2, FioxjRKS n, 21 ; Pi.Ai>: .3, Fi<;ure8 33-35

Male. — length, 5.0 mm.; tegmen, 0.0 mm.

Vertex with anterior margin straight or slightly curved, transverse
carina slightly arcuate on each side of middle line, lateral margins of
frons thickened basally.

100 PROCEEDINGS OF THE NATIONAL MUSEUM vol. M

Anterior portion of vertex, basal three-quarters of f rons, distal por-
tion of clypeus, sides of clypeus and genae before eyes, pronotum
medially, mesonotum, mesopleurites, and fore and middle legs piceous-
brown ; posterior portion of vertex, f rons in apical quarter and clypeus
at base, genae above eyes and around antennae, lateral fields of prono-
tum and basal half of procoxae ivory to pallid, hind legs and sternum
pallid testaceous, abdomen dull fuscous. Tegmina hyaline, basal two-
thirds of clavus yellowish, postcubital (first claval) vein dark in middle,
a yellowish-brown fascia from two bars at middle of costal cell to com-
missural margin at level of union of claval veins, a very broad fuscous
band transversely across tegmina, enclosing stigma anteriorly and of
subequal width throughout, a narrow wedge-shaped band from apical
veins of R to penultimate apical cell of M, apical margin expandingly
fuscous from Rs, two opalescent-hyaline spots between stigma and apex
of clavus between Rs and M and Mo and M3+4, respectively, the veins
at Mf , M-Cu, and Cu distad of apex of clavus yellow. Wings smoky,
veins testaceous.

Anal segment of male deflexed distally through 80°, deflexed portion
fully as long as basal, in profile slightly dilated distally, a semi-cir-
cular excavation on apical margin at middle. Pygofer with lateral
angles shortly produced in a small rounded lobe. Aedeagus tubular,
a short, broad-based horizontal spine at middle on right side, and a
slenderer spine directed caudad at same level on left side, two long'
spines at base of flagellum on left side, one curved dorsad, the other
sinuately ventrocaudad, flagellum tubular, somewhat tumid at base.

Type.—V.S.'NM. No. 57097.

Described from one male taken near Bailos, Ecuador, by S. W.
Frost (February 20, 1937) . This species is distinguished by the shape
of the anal segment of the male and of the genitalia and by the pattern
on the tegmina.

PINTALL4 VOMERIFERA, new species

Plate 3, Figures 36-38

Male. — ^Length, 4.7 mm. ; tegmen, 5.5 mm.

Vertex with apical margin and transverse carina straight, lateral
margins of f rons not thickened basally.

Testaceous- fuscous ; tegmina ivory-hyaline, three brown spots in
costal cell, the basal spot extending in a broken fascia faintly across to
junction of claval veins, a brown spot over R-M and M-Cu, membrane
slightly and unevenly suffused fuscous, vein Cuib pallid, a pale spot in
middle of cell Cuia. Wings very slightly smoky, veins testaceous.

Anal segment of male deflexed distally through 95°, right side
dilated, apical margin very oblique, making apex acutely pointed.
Pygofer with lateral angles not produced, or, if so, very obtusely.

NEW LANTERNFLIES FROM SOUTH AMERICA — FENNAH 101

Aedea^^is tubular, a stout spine in middle line ventrally curved pos-
teriorly, a deep median keellike lobe ventrally in distal half, a long
bladelike process arising at base of flagellum on right directed ventrally
and forwanl, two unequal spines at base of flagellum on distal side
directed dorsally, flagellum with two scroll-like folds. Genital styles
weakly angulate at middle, much dilated distally, outer margins
strongly convex, inner margin straight or nearly so.

Typc—V.S.l^M. No. 57098.

Described from one male taken at Banos, Ecuador, by S. W. Frost
(February 20, 1937). This species is distinguished by the shape of the
anal segment and of the genitalia and by the pattern on the tegmina.

Family DERBIDAE

Tribe OTIOCERINI
IQUITOSA, new genus

Vertex longer than wide across base (3 :1) , disk deeply sunken, lateral
margins thickened and pustulate distally, approximated at apex, pos-
terior margin concave; frons linear, clypeus medially carinate; head
compressed, in profile produced obliquely upward and anteriorly in an
acute angle, almost pointed at tip ; antennae subequal to length of head,
first joint about as broad as long, second joint flattened, with sides
straight, expanding distally, apical margin asj'mmetrically excavate, a
single short vermiform appendage attached at base. Pronotum very
short, anterior margin angularly convex, posterior margin still more
acutely concave, lateral fields broad, quadrate, median carina present,
and a carina on each side between eye and tegula ; me«onotum convex,
depressed in posterior third, median carina feeble, lateral carinae obso-
lete except for two flangelike vertical eminences in middle; tegulae rel-
atively large. Tegmina with sides expanding distally, apical margin
oblique, costal margin sinuat<^, with a prominent recurved rounded
eminence near base, vein M leaving Sc + R one-seventh from base,
Sc + R forking two-sevenths from base, M with five branches reach-
ing apical line of transverse veins, clavus narrowly open, but common
claval vein not passing beyond its apex. Wings four-fifths as long as
tegmina.

Genotype, Iquttoaa shannoiu^ new species.

IQL'ITGSA SHANNONI. new Hprrin

PI.ATC 2. FI0URE8 23-26; Platk :{. Fkjukes 31M1

MaJe. — Length, 3.2 nnn. ; togmen, 5.0 mm.

Head, pronotum except margins, mesonotum except carina, a patch
on each lateral field, and posterior margin brown; median carina of
mesonotum and margins of pron(*tum and mesonotum, sternite^s, and

102 PROCEEDINGS OF THE NATIONAL MUSEUM vol. w

legs testaceous to stramineous, except for a narrow fuscous band at
apex of profemora and a similar band in middle of protibiae. Teg-
mina mostly piceous, a dull testaceous spot on each side of piceous
costal eminence, apical cells from node to Ms and intervening veins dull
yellow with two fuscous interruptions, middle portion of subapical
veins paler than corium, veins of corium chiefly red, basally infuscate
and interrupted with dull yellow spots. Wings smoky, veins con-
colorous, margin red.

Anal segment narrow, lateral apical angles slightly deflexed.
Aedeagus tubular in basal half, with a minute bicuspidate process
near base of flagellum, two stout spines at base of flagellum, one on
right side, the other distally, both directed dorsally, a transparent
spine on flagellum at middle, apex of flagellum lobate with a large
spine below it on left ; two small spines on right. Genital styles nar-
row, membranous dorsally distad of middle, with a scroll-like process
in middle of dorsal margin.

Type.—U.S.^M. No. 57099.

Described from one male collected at Iquitos, Peru, by R. C. Shan-
non (March 1931). The basal reflection of the costal margin recalls
the similar modification found in Sayiana Ball, but Iquitosa differs
from Ball's type, with which it has been compared, in the more pointed
head, the shape of the antennae, and the shape and venation of the
tegmina.

Family ACHILIDAE

Subfamily APATESONINAE

Genus ATESON Metcalf

Ateson Metcalf, Bull. Mus. Comp. Zool., vol. 82, No. 5, pp. 367, 369, 1938. (Geno-
type, A. marmoratum Metcalf, loc. cit., p. 369.)

ATESON SEMILUTEUM, newr species

Plate 3, Figures 42-47

Male. — ^Length, 5.5 mm. ; tegmen, 8.0 mm.

Frons with lateral margins ampliate and raised between level of
antennae and suture, median carina terminating distally against a
transverse ridge, median carina of clypeus distinct except at base.

Testaceous ; abdominal sclerites infuscate. Tegmina yellowish, fus-
cous at base and chiefly so distad of nodal line, with a pale area in the
subapical cells of M and Cuia, transA^erse veins mostly pale. Wings
hyaline at base, smoky near apical margin, veins fuscous.

Anal segment of male short. Pygofer with lateral angles scarcely
produced, pointed, slightly falcate, medioventral process twice as
broad as long, distally shallowly trilobed, the middle lobe largest.

NEW LAXTERXFLIES FROM SOUTH AMERICA — FEXXAH 103

Aedeagus tubular, sli^litly taporiun; distally, pcriandiium rocurved
dorsally at apex into a sinuate spine, penis when everted with two
curved spines directed ventrally and two straight spines porrect
caudad. Genital styles distally expanded, in profile with apical mar-
gin sinuately oblique, dorsal margin sinuately horizontal, folding in-
ward with a short pointed lobe.

Tijpe.—JJ.SSM No. 57100.

Described from one male taken at Para, Brazil, by P. R. Uhler
(coll. No. 132). This species is distinguished by the carination of
the frons and by the pattern on the tegmina, as well as by the shape of
the genitalia.

ATESON LUTEOSPERSUM, new species

Plate 3, Fiouurs 48, 49

Fe77iaJe. — Length, 7.0 mm. ; tegmen, 8.5 mm.

Lateral margins of frons not much raised above level of disk, disk
of vertex scarcely depressed, margins scarcely raised.

Vertex, frons, clypeus, pronotum, and mesonotum fuscous, heavily
speckled with yellow spots, genae testaceous, sternum and legs fuscous.
Tegmina fuscous, heavily marbled and spotted with yellow on mem-
brane, a small patch basad of stigma and over junction of claval veins
yellow, corium wholly fuscous, veins spotted with yellow. Wings
smoky, veins fuscous.

Ovipositor with first valvulae bearing two teeth dorsally near apex
and a curved apical spine; apex of second valvulae rounded in profile.
Genital chamber (bursa copulatrix) with three pairs of sclerotized
plat<'.s, each of the posterior pair, in dorsal view, elongate-quadrate,
each of the middle pair larger, curved and triangular, each of the
third (dorsal) pair small, narrow, sinuately tapering as shown in
figure.

7y/>c.— U.S.N.M. No. 57101.

Descril)ed from one female taken at Cabima, Panama, by A. Busck
(May 28, 1911). Thi.s species is distinguished by the weak carination
of the frons and vertex, by the shape of the second valvulae of the
ovipositor and of the sclerites of the genital chamber, and by tho
distribution of the yellow spots on the tegmina.

ATESON MARMORATUM Metralf
Pr.ATi: 3, FifiUKES r»0-54

AtCHon mannoralum MfrrcAu. Bull. Miih. Coinp. Z<)oI., vol. 82, No. 5, p. 300, 1938.

For comparison with the preceding species figures are given of the
male genitalia of a paratype specimen and of tho distinctive portions
of the female genitalia.

EXPLANATION OF PLATES

Plate 2

1,7-10, Pintalta quadrimaculata, new species: 1, Tegmen; 7, vertex; 8, anal segment,
pygofer, and right genital style; 9, aedeagus, right side; 10, aedeagus, left side.
2, 11-15, Pintalia curvivitta, new species: 2, Tegmen; 11, vertex; 12, anal segment and
pygofer; 13, right genital style; 14, aedeagus, left side; IS, aedeagus, right side.
3, 16-19, Pintalia obliquivitta, new species: 3, Tegmen; 16, vertex; 17, anal segment, pygofer,
and right genital style; 18, aedeagus, right side; 19, aedeagus, left side.
4,22, Pintalia falcata, new species: 4, Tegmen; 22, anal segment, pygofer, and right
genital style.

5, 20, Pintalia marmorata, new species: 5, Tegmen; 20, vertex.

6, 21, Pintalia daedala, new species: 6, Tegmen; 21, vertex.

23-26, Iguitosa shannoni, new genus and species: 23, Anal segment, side view; 24,
aedeagus, left side; 25, aedeagus, right side; 26, right genital style, side view.

Plate 3

27-29, Pintalia falcata, new species: 27, Anal segment, left side; 28, aedeagus, left side;
29, aedeagus, right side.

30-32, Pintalia marmorata, new species: 30, Anal segment, pygofer, and right genital
style; 31, aedeagus, right side; 32, aedeagus, left side.

33-35, Pintalia daedala, new species: 33, Anal segment, pygofer, and right genital style;
34, aedeagus^ right side; 35, aedeagus, left side.

36-38, Pintalia vomerifera, new species: 36, Anal segment, posterior view; 37, anal seg-
ment, pygofer, and left genital style; 38, aedeagus, right side.

39-41, Iquitosa shannoni, new genus and species: 39, Head and pronotum (right antenna
incomplete); 40, head in profile; 41, tegmen.

42-47, Ateson semiluteum, new species: 42, Anal segment and pygofer; 43, medioventral
process of pygofer; 44, aedeagus (everted), right side; 45, right genital style;
46, head, frontal view; 47, tegmen.

48,49, Ateson luteospersum, new species: 48, Right second valvula of ovipositor; 49,
sclerotization of genital chamber.

50-54, Ateson marmoratum Metcalf : 50, Right second valvula of ovipositor; 51, sclerotiza-
tion of genital chamber; 52, right genital style, ventrolateral view; 53, medio-
ventral process of pygofer; 54, aedeagus (retracted), left side.

104

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 2

NEW South American Fulgoroidea,
For explanation 8cc page 104,

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 3
33

South American Fulgoroidea.
For explanation see page 104.

U. S. GOVERNMENT PRINTINS OFFICE: I94B

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

h the

SMITHSONIAN INSTITUTION
IS NATIONAL MUSEUM

Vol. 96 Washington: 1945 No. 3190

TTIF. OKXT'S FT'XDELLA ZELLKK: A COXTRTBrTIOX
low AIM) A IJFAISIOX OF THE AMEKTCAX PVKALI-
DoID Moril-^ OF rilF F.V.MILY PHYCl'lTD.VF

B\- (".\i;r. IIi;im;ii ir

This paper i.s oftVred now because two of the species treated liave
iMcn discovered in the United States, and one of these {pellurcnf<) is
rated as a l)ean pest of some importance in the Tropics and may easily
become one in our Gulf States. Also, there has been considerable mis-
identificaHon of at least three of the species due to the wide variability
of the markings and the mi>ai)i)lication of the name peUucens in eco-
nomic and taxonomic literal iin-. Tlic >])('(ies are markedly dilfeicnt in
theii" genitalia, but herelofoic tlic'-e organs have not l)een figured, nor
have the structuial iiecnliaril ies of the several species been indicated.

The genus is conliii'il \n ilic Niw A\'(irl(|. h'ivc species nrc recog-
nized of which one is iier<' described for tiie jii-.^t time. Two older
names an; reft ri'ed to synonymy. T'lie >l udy is l);i<e(l upon material in
the United Slates National Museum, a considcr;il>le collect imi id' speci-
mens from Puerto Kieo in the Cornell Uidxcrsity collection, and a few
s|iecimens from the .Janse and ]*riti~h .Museum collections.

(Jcmis II NDKI.I.A /»ll.r

rmiili llii Zki.i.kk, I.sIs voii (»kfii, \(»1. 11, p. stu;, l.SJs,— U.vi.onoi', .Mciiioircs sur le.s
I/-pi(l(iI(terf'S, vol. 7, p. UlU. lW.i. — .Iansk., Jeiirii. Kiit. Soc. Soiilli Africa, vol.
•I, p. 1W. 1041. ((Jennfyi)o: Fundrlla jkUuciiik Zellcr.)

lUillnvin DvAU. I'rr»c. U. S. Niif. Mils., vol. 41, p. '.VS.\, VM'.\: Iii.sociilor Tiisciii.ic
Mfiistnuis, vol, 7, p. in, UM'.*. («;i'iintyp<' : Itnllnrin cisHin iiiiis Dyar.)

Anleima of male slightly piibe.-cent, base somewhat enlarged, shaft
laterally (lattened ami very slightly excavate at ba.se (pi. 4, fig. li)

•'.2024.". 4.'i in.'.

106 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

(except in k/nohUis and aliemora) and -with a very small blackish scale
tuft in the excavation (except in ignohilis) ; of female slender, simple.
Front of male head deeply grooved to hold labial palpi ; of female
rounded. Lal;ial palpus upcurved, reaching to vertex, clothed with
broad appressed scales; in male closely appressed to face, with second
segment over three times as long as first and with third segment very
short (about one-sixth the len.gth of second) ; in female with second
segment shorter and third about one-third the length of second. Max-
illary palpus minute. Forewirig smooth ; 11 veins, 10 from cell, parallel
for some distance but not approximate to stalk of 8-9, 9 short, 6 from
below upper angle of cell, straight, 4-5 connate or ajjproximate at base,
3 approximately equidistant from 4 and 2, 2 from before lower angle
of cell. Hind wing with 8 veins, 7 and 8 closely approxinuite beyond
cell for less than half their lengths, 4 and 5 long stalked, 3 from stalk
of 4-5 or closely approximate for some distance, 2 from near lower
angle of cell, cell short, discocellular slanting and slightly curved; in
male anal area, involving veins la and lb, thickened and folded under
to form a pocket enclosing enlarged scales and hair tufts. Eighth
abdominal segment of male bearing a thin, short pair of ventiolaterai
hair tufts.

Male genitalia with uncus long, curved, strongly sclerotized, con-
stricted at middle and broadly divided at apex (hammer-clawed) ;
gnathos terminating in a short, stout hook or a short, broad plate
{ahemora) ; harpe rather short, with clasper ; vinculum narrow, short;
aedeagus stout w4th long, stout, projecting, curved spine or spines
at apex (except in argent ina) ; cornutus a single, strong spine.

Female genitalia without signum {jyellucens) or with signum well
developed and consisting of a large oval or pear-shaped cluster of
thornlike spines {^argentina^ agapella), or curved sclerotized bands
armed with stout, thornlike spines {ahemora^ ignohilis) ; bursa large;
ductus bursae short, broad (narrowest in agdfeUa) ; area surround-
ing genital opening strongly sclerotized, the dorsal sclerotization in
the form of a band connected with the supporting rods of eighth
segment collar, and armed with two or four spinelike projections
(except in ignohilis and some examples of argentlna) ; ductus semi-
nalis from caudal area of bursa.

This genus is easily distinguished by its striking male characters:
the strongly sclerotized, long-stemmed, bifurcate (hammer-clawed)
uncus; the large pocket on anal area of hind wing; the long, embedded
labial palpus with very short third segment; and minute maxillary
palpus. A similar bifurcate uncus is not found in any other Ameri-
can genus except Defundella Dyar. In the type species of the latter
{corynophora Dyar) the uncus is somewhat produced and exhibits a
slight bifurcation at apex; but other species, which must also be

THE (.KM - IINDKI.I.A HKIXHK II 107

1 t'ft'iTcil to I >< fiiiuli'llii, l.itk I Ills cluu'ncii'r. I >( fumhlJa soparatos
rt'Mflily nil otlu-r male st i"iutiirc> : Its irn-allv in I need irnatlios, j^tron^rly
linokcil. partially free sacciiliiv of liarpc it> rniiiidcd linns, and the
1. arrow. vti"on!_dy >-cIt'ini izt'd. deeply iii\ aeinatfd pmkfi of i he .-icriiitc
id" tlu' I'iirhtli alulouiiiial >('i:iih'1ii .

In Fuiiililht tilt' wiii<2' patirrii \;iric- ^-o imiili within aii\- !.M\('n
-pt'cies tliat it atl'ofds no irlialilc chniacit'r I'or >pcfitic iih'iii ilicai ion.
and the sm'iM'al -pccics can he -cpai-alrd with ct'ilaintx onl\- h\ thi ii-
•ronilalia.

KEY TU Till-: Sl'KCIKS UF l-UNDKLLA

MALES

]. ('l.isiM-r ;i str;ii};lii >|iiiir; m<m|.',i;:us simple argentina Dyar

• 'laspor a curveil (li;rinis: ,h'iji":il:us aiiiiPtI with curved, strongly sclerotized

spiii«> nr spines at or near ai»ex 2

_' W itii larjje, stronfily sclerotized snbanal plate; a cluster of several spines

from apex of aedeagus; <-laspei- short pellucens Zeller

Witlmut srlerotized sut»aiial plate; no more than two spines from aedeagus

at or near ajiex; clasper long 3

'■\. (inatlios terminating in a l)road plate; aedeagus with a pair of spines from

apex ahemora Dyar

(Inathos terminating in a short, stout hook; aedeagus with a single spine
from hel(»w ajK'X ignobilis, new siK'cies

FKMALKS

1. r.nrsa copnlatrix wiilii>iit slgnum : (liniiis Imrsae sclerotizi'd throughout
leiigtli, scjerntizaliiin involving part of bursa adjacent to ductus Iiursae

aiirl (hielns si'ininalis pellucens Zeller

r.nrsa eopnlairix with strong signa ; ductus bursae at most only partially

sr-lernt i/.Ml 2

-' Simia iti fmiii nf lar^'e <>\al nr iiear-s|i,i|pei| cjuslers of spines argentina Jtyar

agapella Schaus

SiL'iia ill fiiriii nf bands beariiiir sloiil si)iiies 3

'■'■. Signa eniisisiiiig of If rallier slinri bands, eaeli arnietl witli a rnw of bmg
spines: duel lis Itiirsae willi a slrmigly seleroti/.ed median ndlar; a widely
sjiaee*! pair of long, curved spines from sclerotized area immediately lu'liind

geniial opening ahemora Dyar

Sigiiji <*<>nsisling (if '1 loni:, partially fused bands eaeli ariiieil with a inw nf
slmrl, stniit, llHtriilike sjiines ; diielns bursae with median area uiisclero-
lize«l; no spiiM-s adjacent to genital ojM'iiing ignobilis, new si)ecles

Kl Mil I I \ I'l II I I I NS /. ll.i
IM.MI I. Fk.I Kl > 1 I. t; t;.(. 7 7' ; I'l Ml Ci. Kk.IKI s 1". 1."../

I iiii'lillii iiflhniiiM Zn I Kit. Isis vun nken, Mil. II, p. sCiC, |s)S: llmae See. lint.
Kossieae, Vi>l. pi. p. !.•:{(;, ISSl. |{A(iOMiT. .Memnires slir les l/''lii(l"|ileres.

vol. 7. p. 2ln, ]su:\.
Hitll'iii)! rixtipriiuiM Dv mj. I'hk-. I'. .»<. Nat. Mils., vel. H, p. .•{::;',. P,ii:{.

108 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Fundella cistipennis (Dyar) Dyar, lusecutor Inscitiae Meustruus, vol. 7, p. 40,
1919.~WoLCOTT, Jonrn. Dept. Agr. Puerto Rico, vol. 17, pp. 241-255, 1933;
Journ. Agr. Univ. Puerto Rico, vol. 18, p. 432, 1934; vol. 20, p. 477, 1936.—
Scott, Journ. Agr. Univ. Puerto Rico, vol. 24, pp. 35-47, 1940. (New
synonymy. )

Male. — Antennal shaft with very small black basal tuft (pi. 4,
fig. 3). Palpi, thorax, and forewing grayish fuscous more or less
dusted with whitish and with interspersed reddish-brown scales (in
many specimens the ground color is reddish brown), giving the moth
a distinctly gray or gray-brown appearance to the naked eye. Fore-
wing with a conspicuous, round, darker brown or fuscous sj)ot in
the center of the area usually occupied by the antemedian line, this
dark spot more or less obscured in some specimens but in typical
examples outlined by whitish areas inwardly and outwardly and not
reaching to inner margin or costa of the wing; discal mark at end of
cell obscure, often absent; subterminal line (when distinguishable)
faint, white, indented at vein 6 and at submedian fold ; a row of dark
spots along termen (present only in specimens having an appreciable
dusting of white scales). Hind wing white, translucent, a faint
fuscous border along costa and (in some specimens) a fuscous line on
termen for a short distance from apex; cilia white ; anal pocket yellow-
ish white. Midtibia with a fringe of pale hairlike scales along dorsum.
Hind tibia with a rather long and slender tuft of pale (whitish ochre-
ous) , hairlike scales from the knee joint (pi. 4, fig. 4) .

Alar expanse 19-23 mm.

Genitalia (pi. 4, figs. 7-7c) with a large, strongly sclerotized sub-
anal plate, constricted before and beyond its middle. Harpe with
apex notched below costa; clasper short, curved, situated near middle
of harpe and armed with several setae at its knobbed apex. Aedeagus
Avith a cluster of several long, curved spines from apex ; cornutus long,
straight, stout.

Female. — Essentially like the male in color and markings except
that the dark spot near the base of the forewing is more diffused,
sometimes reaching to the costa. Hind wing usually with a dark shade
along termen.

Alar expanse 19-24 mm. :

Genitalia (pi. G, figs. 15-15a) with bursa copulatrix finely scobinate
but without signum ; ductus bursae flattened, broad, twisted and con-
stricted near genital opening, sclerotized throughout, the sclerotiza-
tion involving bursa adjacent to ductus bursae and ductus seminalis;
sclerotized band behind genital opening armed with four long, stout,
projecting spines; collar of eighth segment invaginated at dorsal
margin to form a sclerotized pocket (pi. 6, fig. 15«).

Types. — In British Museum {peJluce?is) ; United States National
Museum ( cistipennis ) .

THE GENUS FUXDELLA — HKIXIUCH 109

Tiipe localitks. — St. Thomas-. l?iiii>li AVest Indies {pclluecn-s) ;
B:i rl )atlos {cist i pen n As ) .

Food plants. — Vigna un</iii<-uhita ((•(•wpi-as. l)l;i(k-fyt'(l pt'iin, and
<:ardei\ i^cas), Cunavalia cnsiformis (sword beans), Canavalla mari-
time (black bean), Cnjan cajan (pigeon poa), Pha'^cohis hnmtiis (cul-
tivated ami wild lima beans), PJiascohis sp. (Brazilian specimens),
Cassia occidcntalis (one reared siiecimcn I'mni McCubbins Mills,
Puerto I\ico. before me; most records from this last plant are doubtful
aiul proiiably the result of a misidentification of FundclJa argcntina as
( istipcnnis).

I distribution. — I'nited States: Florida. Hobe Soiuid. ^Sliami. Jupi-
ter. Coconut Grove, Marco Island, AValton, Jensen (U.S.D.A. rearings
Ik .in lima beans, February 1944), Vero Beach (J. R. Malloch, Decem-
ber r.>4r). B.MMVMHis. IlArrr : Damien (Deceml)cr, Fel)ruary). Bort-
au-Piince. Montsi:i:i;at (January). Cuba: Santiago, Matanzas.
\'ii;(.iN Islands: St. Croix (March). Puerto Rico: San Juan, Rio
Picclias (^larch-May), Isabella, Catano (July), Vieques Island
(April), Bkazil: Bahia (May), Ceara.

Xinetj'-six specimens examined.

Zeller had two species before him when he described pcJhiccns., and
the one he figured (fig. 411) in Horae Soc. Ent. Rossicae is his ''var.
!»,*• which is Dvar's argcntina. Through the courtesy of Messrs. Riley
and Tams. of the British Museum, I have been able to examine the
genitalia of the male paraty]»o de<i^nated "var. b" by Zeller and a
typical male pclluccns of the Zellei' material from ISIai'aiinita. The
latter proved to be what Dyar described as cistipcnnis and the species
that has ajipeai-ed in economic literature under that name. What
Mtisehler and (»thers have identified as pcllucens could not be deter-
mined without a genitalic examination of their specimens. Probably
in many instances they ha<l mistaken argcntina for pclluccns. since
bfdh species occur in the AVest Indies and Brazil.

Aeeonling to Scott the favored host of pclluccns { — cistipcnnis) in
I'uei-to Rico is the cowpea {Vigna unguirulata), and the species while
iieqiient in lima Ix'ans seldom does serious damage. Potentially it is

an insect of er-oiiMlnic Inipni | ;i|He. Tlie l;il\;ie are pliliiil iil\- tuxl

borers but also bore into the stems and feed on the (lowers i^i their
hosts. They attack. a>^ far as known, oidy Legumino>ae.

I I Mil.l.l.A AIM.IM I\ \ K.Mir
I'l.Mi ."., Kri.cuKs S Sf.- IM.AiK <i, Kl(iri!i:> IJ, 1.'!

i'lDuh Ihi iiillin'(HM ZKr i> R (in p;irl, "v;ir. li"), Isis von < >ki'ii, vnl. |1, p. sCT, 1S4S;

Iliinif Sof. Kilt. Hossicii(>, vol. ]<», p. 2^1, Wk. HI*, I'^^sl (new syimnyniy).
FuuiliUn nifimtina Dyau, Inseeiifor Inscitinc .Menslnnis, vol. 7. i). I<>, VMW.
J'lniihlhi lUfiiHiH I>YAi:, ln.'«eniinr Ifiseiline Menslnnis, vol. 7, ji. I", ini'.t ( new

svimnyniv ).

110 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Male. — ^Anteniial shaft with even smaller black basal scale tuft than
that of pellucens. Forewing gray without the reddish brown, inter-
spersed scaling characteristic of typical examples of peUucens,' entire
basal area to antemedian line dark fuscous gray (with but very slight
dusting of whitish scales toward base in some specimens) ; this dark
basal patch contrasted against the paler gray color of the remainder of
the wing, extending from costa to inner margin and bordered out-
wardly by a narrow whitish line. Otherwise not distinguishable,
superficially, from pellucens.

Alar expanse 15-20 mm.

Genitalia (pi. 5, figs. 8-8c) without sclerotized subanal plate. Ter-
minal projection of gnathos varying from round to pointed (pi. 5,
fig. 8«) at apex. Harpe tapering to bluntly pointed apex; clasper a
single, straight, slightly roughened, appressed spine, situated beyond
middle of harpe. Aedeagus simple; cornutus a single, straight spine.

Female. — Essentially like the male in color and markings except
that the basal area of forewing is concolorous with or contrastingly
paler than the remainder of the wing. A narrow dark line or a
diffused dark shading outwardly bordering the obscure antemedian
line.

Alar expanse 15-23 nnn.

Genitalia (pi. 6, figs. 12, 13) with signum well developed and con-
sisting of a large pear-shaped cluster of thondike spines; sclerotized
band behind genital opening, divided in the middle, simple (pi. 6,
fig. 12) in Argentinian and Brazilian specimens, or armed with a pair
of median, spinelike projections (pi. 6, fig. 13), rather long in West
Indian specimens or short and disappearing in Mexican and Vene-
zuelan specimens.

Types. — In United States National Museum {argentina and
eucasis).

Type loeaUtics. — Tucuman, Argentina {argentina) ; Caracas,
Venezuela {eucasis).

Food plant. — Cassia spp. (reared examples in National Collection
from Cassia hicapsularis and C. corymbosa).

Distribution. — United States: Florida, Biscayne Bay (May),
Coconut Grove (April); Texas, Brownsville (November). Mexico:
Several examples reared from pods and blossoms of Cassia hicapsularis
at Brownsville, Tex., quarantine station. Cuba: Baragua (March),
Habana, Matanzas, Santiago Province. Puekto Rico: Bayamon
(March, September), Vieques Island (April, July), Coamo Springs
(April) , Aguirre Central (August) , San Gernnin (August), San Juan
(November). Haiti: Petionville (June). Jamaica. Venezuela:
El Valle (June). Brazil: Bahia (May). Argentina: Tucuman
(March).

Seventy-three specimens examined.

1 1 f: ( . I- \ 1

\i)ii.i.\ iii:i\i;i( II

111

In rolh'ctioiis [\\\> >|n'fU'> \\;\> ii[)[>VMVi\ liin>t i'l('(|iltiit 1 y lIIidiT \\\0
luiiiH' fi) H III! lis. liotli (iri/) iitlna :nnl jtiJhiciits \\\\\v .iliiiiii tin- >iiiii<'
tlisi i-iliiit'niii and ai't- ahnmlaiit in tlif \\'f-l Imlirs. (Ii niiili. t'loni ni;i-
tfiial ai hand. /" Ihirt i\s si'cins (<» hi' rarer on tlic niiiinhimh 1 hn»u<_di-
• Mit ii> riiiiiH' (irqeiit'iuii shows considcrMlili' variation in fcnialo
ijt'nitalia. A\'c^i Indian -]ifiiin<Mi> haxc rather con-i lii iious spinidikc
e.\ten>ions of the >eh'rot i/.i'd hand hi'hind tlie L^eiutal o|»enin<i. These
are entirely laekiiii: in I>ra/,ilian specimens, and if one liad oidy those
e\treine> he wonhl he jii-tilieil in a---nnii iii:" ihat they wcir at h'a>t
I'aeially distinet. Ilowexi-r. \'i'ne/.nehiii and .Mexican e.\aiii|)lf- -how
an intermedial*' form with veiv short |)roir(i ions, and Central Anieii-
can >iiecimen>. when i'eco\ eicd in >nllicient nnnilicr-. will |Moltalily
-liow all inter:iradation>. The male (.'enitalia ai-e remaiUahly nni form
thronirhont the ranne of the specirs. exhihit injj; only minor individual
\ariatioii- in the shape (d" the terminal projection of the ^nathos.
The type (d' Dyar's t iira.sis i> oidy a -mall, -omewhat faded male (d"
<U(jtiitin(i.

FLNOKLI-A A<;AI'KI.I.A Schnus
Pl.ATK <!. FKilliK 11

/'ini'hilil <l<l>ll>< ll<l .•<( MAT >. /,cinln-ii;t. M <] . ."., Nu. L', \K J7. VJ'SA.

Fi null, . — I*al|)i. liea(l. thorax, and foreuinLT whitish jiray: dark
marking's drali ;:iay: (ran-\er>e antemeilian line (d' foicwinn white,
detiiie(| chiefly liy it> narrow, dark outer hoi-der, sharjdy simiate,
indent<'d a trifle just i»e|ow co-ta. more dccjilv at to]) of cidl and still
more clecpjy at fol(| helow cell: »li-cal dot at end (d' c(dl oh-cur-e;
white -uhterndnal line indented at Ncin (i and at suhmedian fold, hor-
deied inwardly hy a di-tinct dark -hade a- hroad as the white line
il-elf and outwardly hy a -iniilai'. fainter, dark shadinjf. the latter
con-picnoii- only at apc\. Hind w in^^ ;,.. ji, th,. other species of
h II 1x1 1 1 1 II.

-Mar e\|.an-e j-j nun.

(ienitalia (pi. Ci. lii:. 11) like tlio-e of intermediate example- of
iiri/i nfiiiii ex<'ept that the sijjnmn i- ci .n-ider-ahl \ -lualler in propor-
tion Id the -ize of the hursa.

^ III"- — ^" I'niled Slates National Mir-euin.

Tijin Jdriil'ilii. — '{'a^nis Cove, AlU'liiarle, (iala|»a;ros I.-land.s.

I'linil j>liiiif . — I nknow II.

Known only from the female type. SupeilicialK a ili-iinct -pecies.
The female <i:enitalia. however, would indi<ate that <i<i,i i>, llii \~~ onlv a
race of nrifintiiui. .\ male will he needi-d for exaci placement, and
initil it i- axailahle we -hall have to treat nqii jn lln a- a -|iei-ies.

112 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

FUNDELLA IGNOBILIS, new species

Plate 5, Figures 9-9(/ ; Plate 6, Figure 14

Male. — Antennal sliaft without any trace of black basal scale tuft.
Otherwise partaking of the pattern markings of bath fcUucens and
argentina; in some specimens dark basal patch of forewing round
and reaching neither costa nor inner margin (as in typical pellucens) ^
in majority of specimens, however, basal patch occupying whole basal
area (as in typical argentina) ; median and outer areas of wing averag-
ing a trifle paler than in argentina and without the reddish-brown
scaling of pelJucens.

Alar expanse 13-20 mm.

Genitalia (pi. 5, figs. 9-9<:Z) with gnathos terminating in a short,
stout hook. Harpe with apex truncate; clasper moderately long,
curved, and weakly haired at apex. Aedeagus with a single, long,
strong, curved spine from below apex; cornutus a short, stout, curved
thorn.

Female. — Superficially similar to argentina except a trifle paler on
the average.

Alar expanse 15-22 mm.

Genitalia (pi. 6, fig. 14) without spines adjacent to genital opening.
Bursa copulatrix with signa, consisting of a pair of partially fused
bands, each armed with a row of short, stout, thornlike spines; ductus
bursae short and broad, with median area unsclerotized ; eighth seg-
ment collar completely sclerotized except for a small, round, trans-
parent spot on midventer, sclerotization extending to and over area
behind genital opening.

Type and paratypes. — U. S. N. M. No. 57185. Paratypes also in
British Museum and in Cornell University and Janse collections.

Type locality. — Oaxaca, Mexico.

Food 2^1 ant. — Unknown.

Described from the male type and 2 male and 4 female paratypes
from the type locality; 4 male and 8 female paratypes from Tehua-
can, Mexico (May, June, July) ; 3 male and 7 female paratypes from
Orizaba, Mexico; 1 male and 6 female paratypes from Cordoba,
Mexico (May); 1 female paratype from Guadalajara, Mexico; 1
female paratype from Jalapa, Mexico ; 1 male paratype from Cayuga,
Guatemala; 1 female from Costa Rica; 1 male and 1 female paratype
from Santiago, Cuba (June) ; 1 male and 1 female paratype from
Sierra Miestra, Cuba (May) ; 1 male paratype from Cuba without
other locality label; 1 male paratype from Aguirre Central, Puerto
Rico; and 1 female paratype from Petionville, Haiti. Most of the
foregoing were in the National Collection under either pellucens or
argentina. The species is quite distinct and easily recognized in
either sex by its genitalia.

THK (.K\IS II \|)|.l.l.\ IlKIMiK II 113

FlM)i:i,I.A AlllMoltV D.Mit
PiAiK I. Fi(.ri:i: r. : I'l ah: .'i, I"i<,i i;i:s in Id.-; I'l \ri:<;, FiuruE 16

Fitmltlla iilu mora Dyau. I'mc. U. S. .\:it. Mus.. vnj. J7. p la'^. I'.il }.

Mill*. — Aiit<'nna with siii:ill Klark -r.ilc lull al hasc of shaft.
Forewiiiji with no or a \iry faini daik ha-al ])atch (when present
coverinir i)asal area to anteiiieclian liiu') : aiitemcdian line whitish,
veiT faint; subtenninal line white, faint but less oliscure than antc-
niedian. without dark borders exeej^t for an inner and an oiiii r dark
spot at inner margin of wing; veins from cell rather strongly outlined
by dark scaling (the most conspicuous superficial character of the
>pecies). A thick, dark (brownish) hair tuft covering outer surface
of fore tibia ( ]•!. I. tig. '> » . a ihalc rliai-actcr not fnund in oilier species
of the genus.

Alar expanse 18-23 nun.

Genitalia (pi. 5, figs. 10-lOr) with giuithos terminating in a broad
tonguelike plate. Harpe somewhat tapering but with apex truncate;
a stronii tuft of long- scales from costa: clasper long, curved, slender,
with a few hairs at ajiex. Aedeagus with a paii- of long, curved,
llattenecl spines from ai)ex; cornudis a long, straight, sh'uder >pinc.

Fr/no/r. — Essentially like the male in color and markings.

Alar expanse 18-23 nun.

Genitalia (])\. Ct. fig. IC) with a pair of long, widely sjiaced, l)asally
curved s]iin('s from sclerotiz('(l area iniiiiedialely beliind genital open-
ini.'. r>iir-a id| Milat ri.\ with >igiia <<>ii^i^tiiig ol iwn ratliei' >hoi't
baiid>. each arm«Ml with a row of long >pines. Ductus bursae bulged
in the mi<Idle and with a strongly s( leiotized nie<lian collar. Collar
of eighth segment partially sclerotizeil and fuscii veni rally.

Ti//tr. — In United Stale- Xatioiial Mu~euni.

T>/pc lornTtty. — Ori/.aba. Mexico.

Food qyJaut. — UnkiHtw II.

Ifistrihutioti. — Mi:.\ico: ()ri/,ai)a. .lalajia. 'I'eapa (December), Cor-
doba (April. December). Cuernavaca (July), Gi atkm.m.a : Qiiiiigua
iXfarch). Cayuga (January. May). Paiuiha (July). Costa Ivica:
Juan Vinas (Xovember),

•Nineteen sj)eciniens examined.

Sup«'rficially the most easily di-i ingui>lied specie- in (he genus.
The large forelibial tuft at once idetililies the male, and both sexes
can be >e|iaraled bv the r.iiiiei- cori-|>icu<Hi- dark oiillining of the
veins. The vein- .iie -iiiiilarly dark -c.iled in the other sjiecies, but
(he contrast of the daik xcin- again-t the pale iiit er\ enular aieas is
iiiofc marked in uln mnrd.

EXPLANATION OF PLATES

The drawings of figures 7-7c, 8-Sc, d-9d, 10-lOc, 11, 12, and 13
for the plates accompanying this paper were made by Mrs. Eleanor
A. Carlin, formerly with the Bureau of Entomology and Plant Quar-
antine. Figures 1, 2, 3, 4, 5, 6-6c/, 14, 15-15f/, and 16 were drawn by
Mrs. Sara H. DeBord, of the Bureau of Entomology and Plant
Quarantine.

Plate 4

1-4, 6^7c, Fundella pellucens Zeller: 1, Side view of male head; 2, basal segments of male
antenna, denuded to show depression in shaft; 3, same, showing scale tuft on
shaft; 4, hind tibia of male; 6, wings of male showing venation; 6a, ventral
view of anal pocket of male hind wing, opened to show scale tufts; 7, ventral
view of male genitalia with aedeagus omitted; 7a, lateral view of tegumen,
gnathos, subanal plate, and uncus; 7b, aedeagus; 7c, sternite and tergite of
eighth abdominal segment of male.
5, Fundella ahemora Dyar: Tufted foretibia of male.

Plate S

8-8c, Fundella argentina Dyar: 8, Ventral view of male genitalia with aedeagus omitted;
8a, terminal projection of gnathos showing extreme of variation; 8^*, sternite
and tergite of eighth abdominal segment of male; 8c, aedeagus.
9-9d, Fundella ignobilis, new species: 9, Ventral view of male genitalia with aedeagus
omitted; 9a, lateral view of tegumen, gnathos, and uncus; 9b, aedeagus; 9c,
sternite and tergite of eighth abdominal segment of male; 9d, anellus.
10-lOc, Fundella ahemora Dyar: 10, Ventral view of male genitalia with aedeagus omitted;
10a, lateral view of tegumen, gnathos, and uncus; lOb, aedeagus; 10c, sternite
and tergite of eighth abdominal segment of male.

Plate 6

11, Fundella agapella Schaus: Ventral view of female genitalia.
12, 13, Fundella argentina Dyar: 12, Ventral view of female genitalia of type with bursa
copulatrix omitted; 13, ventral view of female genitalia of West Indian speci-
men.
14, Fundella ignobilis, new species: Ventral view of female genitalia.
15-15a, Fundella pellucens Zeller: 15, Ventral view of female genitalia; 15fl, collar of
eighth abdominal segment of female, dorsal view.
16, Fundella ahemora Dyar: Ventral view of female genitalia. .
114

U.S. GOVERNMENT PRINTING OFFICE: 1945

U. S. NATIONAL MUSEUM

.77NV

PRCX:EEDINGS. vol. 96 PLATE 4

-. 1

ahe mora

7 pellucens

The Genus Fundella.
For explanation sec pavrc 114.

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 5

ahemora.

The Genus Fundella.
For explanation see page 114.

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE6

14 ifnobiliJ

15 pellucens 16 ahemora.

THE GENUS FUNDELLA

For explanation sec pa^^:c 1 14.

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued ft'-t^iVvA- sii»JB h the

SMITHSONIAN INSTITimON

U. S. NATIONAL MUSEUM

Vol. % Wwhington: 1945 No. 3191

A M:\V (iENUS AND TWO NEW SPECIES OF PEKCOID
FISHES FROM XEW GUINEA, FAMILY CENTROPOMIDAE

Hv Lion Aiv'i) P. Schultz

Lt. Jmiik's R. Simon, U.S.N.R., irccntly donated to tho Unitod States
National Museum a small collection of lishes jtriven to him hy his friend
Capt. Rail)]) F. Honess, U.S.A.. who collected them in New Guinea
in 1044. In this lot. lotalin*; '27 s])ecimens, ^Yere eijiht examples from
fresh water that I consider to helonj; to an undcscribed pei-coid <;enus
invoh in<_' two new species. These are described herein. The other 19
.sp»'cinicn< arc of known foiins oc('nn'in;Lr in the re<ri<>ii.

XENAMBASSIS, new genus

Geiwtypi . — Xr)u//)iha.y.s/',<< lifrytpss'/. new species.

Body ronij)ressed. covered with cveioid scales of moderate si/e every-
where except on top of head: basal half of caudal fin scaled and a few
-cales on base of peel oi'al : lateral line arched ^\\^'\• j)e<'loral fin so that
It is concurrent with doisal pi'ofile. then extendin;:; alon^ midaxis of
body posteriorly; each lateral line tube straiefht. not extendine; entire
len^jth of scale ; anterior profile concave ovci- orbits : intei-orbital space
a little convex; a low predorsal ridejc exteiidinir alon^r middorsal line
endin«r at occiput: mouth teiminal. a little obruiue; premaxillary
diLditly [irotractile : maxillaiy withoiil supplemental bone, mostly t'X-
posed, o(dy the doj-sal edp- slipi)in;j: sli^dit ly mulei- ed<fe of preorbital
bone; maxillaiy with its {posterior ed^'e conca\e: jaws e(|ual or nearly
-CI. the lower sli^htlv in front id' upper: iccth in both jaws in a \illi-
t'orm band, with the out«*r row consisting; of somewhat e;darjie(l conical
teeth, the band of teeth becoming' narrower on sides of jaw. endinjj
in onlv one or t wo rows on lower jaw : vomer with a patch of villi form
teeth and palatines with a nairow row of \ illifoini teeth : no teeth on

620241— 4.^ 11''

116 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

tongue; cheek and operculum scaled; posterior and lower margins
of orbit denticulate and two denticulate ridges on preorbital bone
above maxillary, the upper one forming part of the serrae around
orbit ; preoperculum with double edge, the outer one serrated on both
posterior and ventral borders, the inner one serrated at lower angle
and a little on each side of the angle above and forward; inter-
operculum denticulate ; operculum not spinate ; gill membranes united
far forward, free from isthmus; branchiostegals 6; pseudobranchiae
present ; gill rakers of moderate length, of rather heavy build ; a single
dorsal fin with the spiny portion scarcely longer than soft portion of
VIIT or IX spines, the second or third longest, the next to the last
shortest; spinous dorsal fin preceded by a short recumbent spine,
directed forward, hidden in the skin beneath the scales ; anal with 3
spines and usually 10 soft rays ; both fins with a sheath of scales along
their bases, one scale in width along dorsal and one or two along anal,
these fins partly depressible between the sheath ; pectorals ii, 13, asym-
metrical, upper 2 or 3 branched rays longest ; pelvic fins, I, 5, inserted
under base of pectorals and a little in front of dorsal origin; pelvic
spine slender, not reaching to anus and about two-thirds length of
first branched pelvic ray, which ends in a short filament; axillary
scale of pelvic small, about length of diameter of pupil; caudal fin
forked, lobes more or less pointed.

Other characters are those of the genotype, X. honessi, described
below.

This new genus is related to the percoid fishes usually referred to
the family Centropomidae but sometimes separated from them and
grouped in the family Ambassidae or Chandidae. Xenainbassis is
especially close to Tetra^entrmn Macleay, Synecliopterus Norman,
and Amhassis Cuvier and Valenciennes, differing from them as in-
dicated in the accompanying key. Ambassis differs from the new
genus and from Tetracentrum and Synechopterus by having the first
dorsal fin made up of VII spines and joined to the base of the first
spine of the second dorsal, whereas in the other three genera the first
dorsal is joined at least halfway out the last spine so that the two
fins are continuous. Chanda Buchanan-Hamilton, 1822, with G. lala
as the type as restricted by Fowler (Proc. Acad. Nat. Sci. Philadel-
phia, 1905, p. 500) differs from the above-mentioned genera chiefly in
having 14 to 17 soft rays in the anal fin. Pseudoarribassis Castelnau,
1878 {— Austrochanda Whitley, 1935), a substitute name supposed by
Whitley to be preoccupied by Pseudarribassis Bleeker, 1876; and Vel-
anihassis, proj)osed by Whitley (Rec. South Australian Mus., vol. 5,
No. 3, pp. 356-365, 1935) , are here considered as subgenera of Amhassis
Cuvier and Valenciennes. Indeed, they may even be synonyms of
Ambassis, since the generic differences appear so slight. I also refer

NEW PERCOID FISHES I'HOM NEW GUINEA SCHULTZ 117

Acanthoperca Castoliuiu, 1S78, ami lilandoirskieUa Irediile and Whit-
ley, 1932, as subgenera of Aiiibassh. Priopidichthys Whitley, 1935,
with teeth on the tongut.'. may well he a valid grnus, as none of the
other irenera seem to have lingual teeth.

Named Xena//>h(issi.s, meaning a strange or different Ambassis.

KEY TO THE GENERA AND SPECIES OF NEW GUINEA
CENTROPOMIDAE RELATED TO TETRACENTRUM

la. Anal rays IV, f>; dorsal rays IX, 10, dorsal tiu loiitiiuious ; preorbital witli 2
serrated ridges; suborbital and postorbitnl ring of bones witb denticula-
tions ; inner double edge of preoi>ercular bone strongly toothed at angle
and along its lower edge; outer edge of preoperculum strongly denticulate
aluug its entire border; least depth of caudal peduncle Ir.i in its lenytii :
profde concave over orbits ; pelvics inserted under i^)ectoral base; postorbital
length of head 1.1 in length of caudal peduncle.

Tetracentruni ' apogonoides (Macleay)
lb. Anal rays 111, ."S lu 11 (see table 1).
2a. Dorsal fin continuous, no deep notch in front of last spine, membrane be-
twt^n la.st two spines connected over halfway out last dorsal spine; sec-
<»nd dorsal spine not reaching anywhere near base of last dorsal spine
when fin is dtpressed.
3n. Orbital rim without serrae; profile over orbits convex; inner ridge ot
preoperculum with 1 or 2 serrae at angle; pelvics inserted just behind
pectoral base; anal origin under next to last dorsal spine; least deptli
of caudal peduncle 1% in its length; postorbital length of head 1.3 in
length of caudal peduncle; dorsal rays IX, 10; 14 gill rakers on lower
half of first gill arch ; pectt)ral with 14 rays.

Synechopterus caudovittatus Norman ^

:ib. Orbital rim denticulate; profile over orbits a little concave; inner ridge

of preoperciilinn with lower and posterior sides near angle denticulate;

Itelvic iiLsertion under pectoral fin base; anal origin under base of last

dorsal spine or base of first soft dorsal ray.

.}a. Dor.sal rays VIII, 10 or 11; a blackish band along nudaxis of body

commencing behind head and becoming more intense on caudal

[jeduncle, its width about equal to that of pupil, ending in a dark

blotch at base of caudal fin Xenanibassis honessi, new speci<>s

■'ih. Dor.sal rays IX, 10; midaxis of body with a narrow dark streali on
caudal region not ending in a large dark lilol( h at base of caudal tin.

Xenanibassis simoni, new specie's
2h. Dorsal lln dcply notclied, first portion connected at ba.se of first spine of
second dorsal fin; dorsal rays VII-I, 8 to 14; second dorsal spine long,
slender, when depressed Its tip reacliing past base of last dorsal spine.

Aiubassis ' Cuvier and Valenciennes

' Whitlfy, 1935, propoHed VcgamhaHsit to replace Tctracentrum, which ho Hnid was pre-
oi('ii|il.-.| l)y Trtrnrrntron BrainT, 18t!.1. but the apt-llinR U not In conflict, acconllnK to
oplnioiiH 147 anci \\H of the Intcriuitioiial CotnmlHRlon on Zoological Nomenclature, and bo
Tctracentrum muHt Htand aH a valid name.

' Copela. lOa.',. No. 2. pi>. 01-0:j. flj;. 1.

•For a key to the Hpecles of AmhatitiH Hee W'elxT and (!<• Iti-aufort, FIhIh-h of the ludn
.\ufltrallnn Arrhlpelajco, vol. 5, p. 398, 1029.

118

PROCEEDINGS OF THE NATIONAL MUSEUM
XENAMBASSIS HONESSI, new species

Figure 3

HoJoti/pe.—V.S.^M. No. 122830, a specimen 87.6 mm. in standiird
length, collected in 1944 by Capt. Ralph F. Honess, U.S.A., in either
the Samboga or the Giriia River at Buna, New Guinea (long. 148°
30' E.; lat. 8°45' S.).

Paratypes.—U.S.^M. No. 122831, 3 specimens, T7.5, 80.5, and 82
mm., collected along with the holotype and bearing same data;
U.S.N.M. No. 121832, 2 specimens, 58 and 87.5 mm., bearing same
data as holotype.

Descriptio7i. — Detailed measurements were made, and these data,
expressed in hundredths of the standard length, are recorded first for
the holotype and then for two paratypes in parentheses, respectively.
Standard lengths in millimeters 87.6 (82; 87.5).

Table 1. — Counts recorded for certain species of Centropomidae from New Guinea

Number of fin rays

Number of scales

Sppcios

Dorsal

Anal

Pectoral

Lateral line

Above

lateral

line

Below

lateral

line

VIII,

IX, 10

III, 10

IV, 9

ii, 12

ii, 13

1
1

1
6
2

1
1
6
2

1
1
5
1

Syn echopter us cau dorittaius

11
4

XeTianibassis siiuoni

Greatest depth of body 46.2 (45.1; 44.6) ; length of head 37.7 (37.8;
37.4) ; length of snout 9.25 (9.39; 10.2) ; diameter of eye 11.0 (11.3;
11.8) ; least width of interorbital space 9.82 (9.76; 9.02) ; length from
tip of snout to rear tip of maxillary 13.7 (13.4 ; 14.7) ; postorbital length
of head 18.3 (18.8; 17.7); least width of preorbital opposite tip of
maxillary 2.28 (2.20; 2.28) ; least depth of caudal peduncle 14.4 (14.4;
13.7) ; length of caudal peduncle from base of last anal ray to mid-
caudal fin base 18-8 (18.7; 17.4) ; length of procumbent embedded spine
at origin of dorsal fin 4.56 (4.27; 4.34); lengih of longest ray of
pectoral fin 25.5 (27.1 ; 25.7) ; longest soft ray of pelvic fin to end of fila-
ment 25.7 (26.8; 25.1) ; length of pelvic spine 15.6 (16.5; 16.6) ; length
of first dorsal spine 8.10 (6.95; 7.77) ; of second dorsal spine 19.6 (19.8;
18.3) ; of last dorsal spine 13.8 (14.5; 15.2) and next to last spine 13.5
(13.0; 13.2) ; longest soft ray of dorsal fin 20.5 (20.1 ; 21.7) and of anal
fin 21.6 (22.0 ; — ) ; length of first anal spine 8.56 (6.22 ; 8.34) , of second
12.0 (12.8; 15.6), and of third anal spine 14.8 (14.0; 14.8); longest
caudal fin ray 34.2 (33.5; 30.0) ; shortest midcaudal fin ray 16.8 (17.4;
16.1) ; length of longest gill raker on first gill arch 3.77 (4.27; 3.65) ;

NEW PEHCOll) FISHES IKOM NEW CTINEA — StlllLrZ

IIU

(listaiK-e from ti|) of snout to dorsal orijriii 4(').S (47.0; 47.4) : snout to
niuil oii<:iu G")!) (G4..'5; 65.7): snout to pectoral insertion 34.2 (30.8:
3(5.0); snout to jjelvie insertion 3i>.0 (39.0: 11.5): snout to center of
:inus .'m.C) (5r..7 : ."rj.l ) : ceiitei- of anus to anal ori<i;in 7.<t.") (7.32: ,s.()() i.

The follo\vin«r counts were made, respectively : Dorsal rays VI 11. 11
(VIII. 10: VIII. 11: VIII. II: \TI1. 11: VIII. 11) : anal rays III, 10
in all six specimens; pectoral rays ii. 13 in each side of all six s[)eci-
mens: pelvics always I. 5: iriH rakers 6+1 + 13 in all the types; scales
in lateral line 31 (30; 31; 31: 30; 30) ; scales ahove lateral line 4 (4:4;
4; 4; 4) and helow lateral line to anal ori«;in 7 (7: 8; 7: 7; 7) ; zigzag
.scales around caudal i)eduncle 15 (l6: IC); 16; 15; 16) ; branched rays
of caudal fin always 8 + 7. totaling i:».

\^fl^_v

Figure 3. — Xenambassis honessi, new species: Holotype (U.S.N.M. No. 122830), standard
length X7.6 mm. Drawn by Mrs. Aimc \I. .\wl.

Depth of body about 2'/). head 2%, both in standard length; snout
shorter than eye 3»'- to 4. eye 3yi„ to IP/-,, both in length of head; in-
terorbital about e<|ual to snout; maxillary with its rear margin a little
concave, not quite reaching to below middle of pupil; three series of
scales on the che<.»k ; dorsal origin over rear of base of pectoral Hn ;
first dorsal spine short, second aixl third of nt-arly e(|ual length, those
following gradually shorter, the next to last sjjine shortest, almul
three-fourths length of last : anterioi- soft rays of both dorsal and anal
tins longest, longer than any of the spines in the same fin>; first anal
>pine about two-thiids length <»f second and third, the ladcr s|)ine
usually a trifle longer than the second ; -pines of dorsal fin about ecjual
to f)ostorbital length of head: pelvic spine a little shorter than |)o>t-
orbital length of head and the first soft ray a little pr<i(|uccd. reaching
about opjiosite the anal origin: pectoral fin iieaily or (|nitc reaching
lo opposite anal origin; vertebrae 10 i 17.

120 l'lU)C;iOEDlNCiS OF THE NATIONAL MUSEUM voi-. »b

OoTonrJion. — Goncrnlly brownish, with the center of each scale
pnU'i-; a pule, hroiul, wedge-shaped bh)(ch extends from in front of
anus to pelvic base, upward to behind pectoral fin base; a blackish
laleral band extends alono- niidaxis of body, more intensely blackish
l)os(eriorly, eniliiig in a black blotch at base of caudal lin ; a dark blotch
about size of eye or smaller occurs just dorsally to anterior base of
anal fin ; all (ins dusky, the pip;ment more intense basally on interradial
membranes ol" soft I'ays of each median lin; clieek pale; operculum
with dusky blotch above and below, w hich are palei- areas; peritoneum
black.

lieinarka. — This species may be separated from all rehited forms by
the foregoing key.

Named honessl in honor of its collector, Capt. Ilalj)h F. Honess,
U.S.A.

XENAMBASSIS SIMONI. ni-w KpetiiN

Figure 4

llolotype. — U. S. N. M. No. l'J282.S, a male specimen 80 nnn. in
standard length, collected in either tlie Samboga or the Girua Kiver
at Huna (long. 148^'U)' E., lat. 8°45' S.), New Guinea, by Capt.
Uali)h V. lioness, U.S.A., in 1944.

Pamti/pe. — U. S. N. M. No. 122829, a fenuile si)ecimen, 08.5 nmi.
in standai'd length, taken along with the holotype and bearing
same data.

Description. — Detaiknl measnrements were made, and tliese data,
expressed in hundredths of the standard length, are recorded first
for the holotype and then for the para(yi)e. Standard lengths in
millimeters 80 and (58.5.

(JreatesI depth of body 45.0 and 48.8; length of head ;H).9 and 40.7;
length of snout 9.25 and 9.78; diameter of . eye 10.9 and 13.0; least
width of inlerorbital space 9.38 and 10.4; length from tip of snout
to rear tij) of maxillary 13.5 and 14.2; post orbital length of head
17.4 and 19.9; least width of i)reorbital opposite tip of maxillary
2.00 and 2.34; least depth of caudal i)e(luncle 15.1 and 15.0; k'ugth
of caudal peduncle from base of last anal ray to midcaudal tin base

17.3 and 18.7; length of embedded procumbent dorsal spine at dorsal
origin 4.25 and 3.94; length of longest ray of i)ectoral fin 25.3 and
24.8; longest soft ray of pelvic lin to end of lilament 27.1 and 2G.7;
length of pelvic spine H').2 and 17.4; length of first dorsal spine 8.50
and 7.44; of second doi-sal spine 19.2 and 21.3; of last dorsal spine

14.4 and 11. 5; of next lo last dorsal spine 14.1 and 11.5; longest soft
ray of dorsal fin 19.5 and 19.0; of anal lin 20.0 and 20.3; length of
first anal spine R.50 and 7.88; of second 13.3 and 14.9; of third anal
spine M.4 and 14. 0; longest caudal lin ray 33.1 and 34.0; shortest
midcaudal lin ray 10.2 and 10.3; length of longest gill raker on first

NEW PERCOin FISHES FROM NEW (UTIXEA SCHULTZ

121

g\\\ arch ;i.l3 iind i5.65; (.listanoe from snout tip to dorsal ori«xin 44.4
and 49.G; snout to anal origin 64.0 and 04.8; snout to pectoral inser-
tion 33.8 and 37.7; snout to pelvic insertion 39.3 and 43.0; snout to
center of anus 56.8 and 58.8; center of anus to niial orif^iii 7.02
and 5.55.

The following counts were made, respectively: Dorsal rays IX, 10
and IX, 10; anal rays III, 10 and III, 10; pectoral rays ii, 13-ii, 13
and ii, 13-ii, 13; pelvics always I, 5; gill rakers on first arch 6+1 + 13
and 6 + 1 + 13; scales 30 and 31; scales above lateral line 4 and 4, and
below lateral lino T and 8; 7,igzag scales around caudal peduncle
15 and 10; branched caudal fin rays 8 + 7 and 8 + 7.

Fu;lre 4. — Xenambassis simoni, new species: Holotype (U.S.N. M. No. 122828), standard
length 80 mm. Drawn by Mrs. Airae M. Awl.

The .shape of the body, length of fins, and otiior characteristics
except certain counts and coloration are so similai- to X. hoiusK}
that it is not deemed necessary to repeat thom here, since dct.iili'fj
measurements are given ai)ove.

Coloration. — (leneral coloration light brownish with centers of
.scales paler; lateral line i)lackish on caudal region but no black
blotch at base of caudal fin; area behind pectoral ami above |)elvic
fins pale to midaxis of l)0(ly; median fins dusky, more intensely
pigmented on interradial membranes basally; paireil fins light dusky,
pelvics darker distally; peritoneum black.

Rrwarh-H. — This new species may be distinguished IVoiu <losely
ulatefl forms by the foregoing key.

Named nlwoni in honor of Lt. James R. Simon, U.S.N.H., formerly
of the Wyoming fiame and Fish Commi.ssion, who donated the
specimens collected by Captain Honess to the National Museum.

n » «OVIItllMCNT PXINTINa OfrirC l>4>

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued |^r»(vA- QlJfil h ^'

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM

Vol. 96 Washington: 1945 No. 319 2

THREE NEW SCIAEXID FISHES OF THE GENUS OPHIO-
SCTON FROM THE ATLANTIC COASTS OF CENTRAL
AND SOUTH AMERICA

By Leonard P. Schultz

During my studies of the sciaenid fishes of Venezuela I found a
specimen of phioscion (U.S.N.M. No. 86710) from Uruguay, col-
lected by the late Dr. Hugh M. Smith in 1922, with 28 soft rays in the
dorsal fin. This count did not agree with the statements of recent
authors who have reported on fishes from Panama and from the
West Indies under the name of adustus. Although Jenyns and
Agassiz each counted 28 soft dorsal rays for adustus from Maldonado
and Montevideo, Jordan and Eigenmann in their review of the
Soiaenidae (Rop. U. S. Comm. Fish and Fisheries for 1886, pt. 14,
p. 403, 1889) decided that Agassiz's count was incorrect and should
have been 22 or 23, or perhaps 18 or 19. Thus the species of the western
Atlantic were confused in the first review and the name adustu,'i has
been applied to two or three Atlantic species.

This contribution discusses the relationships of the western Atlantic
species of the genus Ophioscion and describes three new species from
Panama, Venezuela, and Brazil.

Drawings of figures 6 to 8 were made by Mrs. A. M. Awl.

GenuH OPHIOSCION Gill

OlihioMfion (JiLr.. I'roo. Afnd. Nat.'Sci. I'liiliidcliihia, vol. ITi, p. 1(V5, 1863. (Type:
OphioJtritm typU-HH GUI, Itasj'd on U.S.N.M. No. iJ2SGl, west con.st of Panama.)

No at(eni})t is made to include heroin the numerous refenMiccs to
jocjilitv r*"'>!-.U of ~t,ifi<-t; nf (li(> gi'iuH f) pit i/>sc'n>ti from the western

123
62o4-12 -45

124 PROCEEDINGS OF THE NATIONAL MUSEUM vol. b6

Atlantic Ocean, because this would require a study of the specimens
on which such reports were based, and these specimens are not now
available.

The genus Ophioscion is closely related to Bairdiella Gill and
Stellifer Oken. Meek and Hildebrand (Marine fishes of Panama,
vol. 2, p. 611, 1925) separated Stellifer from Bairdiella and O'phioscion
in their key on the basis of the skull being "excessively cavernous,
spongy to the touch." I had some difficulty separating the various
species referred to these genera by this character, and so 1 made a
dissection of the upper surfaces of the skull, removing the scales and
skin, thus exposing the nature of the cavernous skull. Bairdiella
chrysura (Lacepede), Ophioscion typicus Gill, and Stellifer rastrifer
(Jordan) all have cavernous skulls dorsally, and also around the orbits
occur narrow bony bridges or stays supporting the overlying skin
and scales. Stellifer has a broader interorbital space, and thus the
caverns are a little broader and by touch can be felt a trifle more easily
than the slightly narrower caverns in the other two genera. My dis-
sections indicate that the caverns are weJl developed in all three gen-
era ^ and are of little value as a diagnostic character in the separation
of these three genera, especially if the specimens are weU hardened
in preservation.

Ophioscion is said to differ from Bairdiella by having the lower
spine of the preopercle pointing straight backward and a little down-
ward, whereas in Bairdiella it is said to be hooked downward. The
lower preopercular spine in these genera is so variable among the
various species of Bairdiella and Ophioscion that I cast serious doubt
on its usefulness as a character. None of the species of Ophioscion has
any of the preopercular spines hooked downward. However, some
individuals, especially young examples of Bairdiella chrysura^ like-
wise do not have the lower preopercular spine hooked downward,
although in adults of Bairdiella that spine is hooked downward.

I have searched for characters to separate these three genera but
have found indications of overlapping, and so the following char-
acters are not wholly satisfactory, although the genera can be sepa-
rated by them when taken together :

Bairdiell'G, with an obliquely terminal mouth; both jaws of nearly
same length; lower jaw with minute teeth in a narrow band of two or
three rows forward and in a single row of slightly enlarged teeth
posteriorly ; the pair of small pores at tip of chin close together and
lying more or less in a shallow depression (fig. 5, &) and the margin of
the snout lacking the small lobes at each side of the median pore

* I removed the skin and scales from specimens representing various species usually
referred to the following genera and found the dorsal part of the skull to be cavernous :
TJmhrina Cuvier ; ilicropogon Cuvier and Valenciennes ; Plagioscton Gill ; Macrodon Schinz ;
Gynoscion Gill ; Corvula Jordan and Bigenmann ; Larimus Cuvier and Valenciennes. In
Mentlcirrhus Gill the caverns were much smaller than in the other genera examined.

THREE NEW SCIAEXID FISHES — SCHULTZ

125

(fig. 5, a) ; the «2:ill rakers moderately long and slender, contained less
than twice in the diameter of the eye; the first soft ray of the pelvics
ending in a filament.

Stellifer^ with an oblique mouth, somewhat intermediate between
BairdieUa and Ophios^cion^ a little more inferior in position than in
BairdieUa but not ventral in position as in Ophioscion; the snout
projecting a very little in front of the tip of the lower jaw; low^er jaw
with teeth in a narrow villiform band, the inner row of which is a
little enlarged; the pair of small pores near the tip of chin separated
by a small bony knol) (fig. o, d) ; the margin of the snout lacking a
lobe at each side of the anterior median pore near margin of snout

b d

Figure 5. — Diagrammatic sketches of the tip of the snout and of the anterior part of the
underside of the lower jaw of three species of sciaenid fishes: a. Snout tip of BairdieUa
chrysura (Laccpede); h, lower jaw of B. chrysura; c, snout tip of Stdlifer rastrifer (Jor-
dan); d, lower jaw of S. rastrifer; e, snout tip of Ophioscion typicus Gill (type, U.S.N.M.
No. 22861); /, lower jaw of 0. typicus.

(fig, 5, c) ; the gill rakers slender and long, equal to eye or contained
in it fewer than two times; the first soft ray of the pelvic fins ending
in a filament.

Op/i/osc!on, wilh the mouth in a somewhat ventral position, the
lower jaw included and jaws nearly horizonlal, with the snout pro-
jecting a little beyond the tip of the lower jaw ; lower jaw with a wide
Ijand of villiform teeth, none of whicii is enlarged; the pair of small
l)ores at midtij) of hnver jaw close together and usually lying in a
sliallow depression (fig. 5, /) ; the front margin of the snout bearing
a short, blunt lobe each side of the anterior median |)()re (fig. 5, e) ;
file gill rakers usually short, not slender, and coiitaiiied more than
•J. 4 to times in the eye; (he first soft ray of the |»el\ir liiis ending in
a filament, usually white.

Thorugh the courtesy of Dr. Tiiomas Barbour, of the Museum of
Comj^arative Zoology, I ha\e been able to examine some of (lu^ speci-
mens dewribed by Jordan and Kigenmann in l.ss!) in llieii- review of

126 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

the Sciaenidae. ^''Stellifertts naso Jordan," based on specimens from
Cachiura, Brazil (M.C.Z. Nos. 4583 and 10808), definitely belongs
to the genus Ophioscion. Specimens of Stellifer microps Steindachner
(M.C.Z. Nos. 4581 and 1031), from Para and Tonteboa, are Ophioscion
microps (Steindachner).

Jordan and Eigenmann identified M.C.Z. No. 22417 as Sciaena
adusta Agassiz, but my reexamination of this specimen indicates that
it is Ophioscion punctatissimus Meek and Hildebrand.

The following key is intended to separate the species of Ophioscion
occurring in the Atlantic along the coasts of Central and South America
and in the West Indies :

la. Dorsal rays usually X-I, 28 or 29; anal rays II, 8; gill rakers on first gill
arch 9+1+16; scales 50 to 57; dark streaks commencing at upper part of
back, passing forward and obliquely downward, then a little above lateral
line bending abruptly downward, almost vertically, disappearing near mid-
axis of body ; tip of spiny dorsal fin dark, base of dorsal fin with a narrow
pale or whitish band, above which the fin is abruptly darker ; opercle dusky

(mouth of Rio de La Plata) Ophioscion adustus (Agassiz)

16. Dorsal soft rays fewer than 25.
2a. Anal rays II, 9.

3a. Dorsal rays X-I, 21 ; gill rakers on first gill arch 8 or 9+1+13, totaling
22 or 23 ; scales about 45 or 46 ; eye diameter 1.2 to 1.8 in interorbital

space Ophioscion brasiliensis, new species

8&. Dorsal rays XI-I, 21 or 22; gill rakers on first gill arch 9 to 11+1+17 to
19, totaling 28 to 31 ; scales about 47 to 51, eye diameter from 1% to 2%

times in interorbital space Ophioscion microps (Steindachner)

26. Anal rays II, 7 or 8.

4a. Scale rows above lateral line usually 51 to 57.

5a. Gill rakers on lower part of first gill arch usually 16 to 18 including
rudiments ; dorsal rays XI-I (rarely XII-I) 21 or 22 ; anal rays II, 8 ;
scale rows 52 to 54; color more or less plain grayish above, paler

below Ophioscion venezuelae, new species

56. Gill rakers on lower part of first gill arch 11 to 13 (see table for counts) .
6a. Dorsal rays X-I (occasionally XI-I) , 22 to 24 ; anal rays II, 7 (rarely
II, 6) ; gill rakers 7 or 8+1+11 to 13; scale rows above lateral
line about 54 to 57.

Ophioscion punctatissimus Meek and Hildebrand
66. Dorsal rays X-I, 20 or 21 ; anal rays II, 7 (rarely II, 8) ; gill rakers
7 to 9+1+13 ; scale rows above lateral line about 51 or 52.

Ophioscion panamensis, new species

46. Scale rows above lateral line 45 to 49 ; anal rays II, 8 ; dorsal rays XI-I,

21 ; gill rakers 8 or 9+1+14 or 15; scales 46 to 49; eye diameter 1.0 to

1.1 in interorbital space Ophioscion naso (Jordan)

OPHIOSCION ADUSTUS (Agassiz)

Corvina adusta Agassiz, in Spix and Agassiz, Selecta genera et species . . .

Brasiliam . . ., p. 126, 1831 (Atlantic Ocean off Brazil).— Jenyns, The

zoology of the voyage of H. M. S. Beagle, pt. 4, Fishes, p. 42, 1842 (Maldonado

and Montevideo).
Ophioscion adustus Tortonese, Boll. Mus. Zool. Anat. Comp. Univ. Torino, ser. 3,

vol. 47, No. 100, p. 130, 1939 (Rio de Janeiro and Montevideo).

THREE NEW SCIAENID FISHES — SCHULTZ

127

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128 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

? Ophioscion icoodwarcU Fowled?, Proc. Acad. Nat. Sci. Philadelphia, vol. 89,
p. 311, fig., 1937 (Port-au-Prince, Haiti).

Corvina adusta Spix, pi. 70 in Spix and Agassiz (1831), has but
X-I, 19 dorsal fin rays and 52 vertical scale rows above the lateral
line; the anal rays as shown are II, 7. Obviously Agassiz did not
describe the fish figured by Spix, since he gave dorsal rays as X-I,
28 and anal rays II, 9. The most significant character in certain re-
spects as clearly shown in plate 70 is the direction of the scale rows
below the lateral line. None are shown parallel with the axis of the
body, but the scale rows run obliquely upward and backward to the
lateral line and the scale row arising from the rear base of anal fin
meets the lateral line just behind a vertical from the rear base of the
dorsal fin. The direction of these scale rows strongly suggests that
plate 70 in Spix and Agassiz (1831) may be some species of
Plagioscion^ perhaps near P. pauciradiatus Steindachner (1917).

The following references list O. adustus^ but their counts disagree
with those for the true adustus: Ribeiro, "Fauna Brasiliense Peixes,"
Arch. Mus. Nac. Rio de Janeiro, vol. 17, family Sciaenidae, p. 23, 1915,
and.Devincenzi, Ann. Mus. Nac. Montevideo, ser. 2, pt. 5, p. 239, 1924.
It must be concluded, therefore, that their descriptions must apply
to some other species or that they were erroneously drawn up.

The description by Jenyns fits very well the specimen before me
(U.S.N.M. No. 86710), which is 120 mm, in standard length and was
collected by Dr. H. M. Smith in Uruguay in 1922.

Berg (Ann. Mus. Nac. Buenos Aires, vol. 4, p. 52, 1895) listed
Sciaena adusta (Agassiz), but his counts did not agree with those of
Agassiz or of my specimen, and they need reexamination to determine
the identity of his material.

Ophioscion tuoodtcardi Fowler {loc. cit.), described from Haiti in
1937, probably is a synonym of adustus. I have not seen Fowler's
types, but the number of fin rays places it with adustus, and the white
area along base of dorsal fins abruptly set ofi' by the blackish area
distally along basal part of dorsal fin rays, as in our specimen
(U.S.N.M. No, 86710) from Uruguay, indicates that woodwardi and
adustus are the same.

OPHIOSCION BKASILIENSIS, new species
FiGUEE 6

Uolotype. — U.SN.M. No. 87742, one specimen, 77 mm. in standard
length, taken over a sand bar at Santos, Brazil, September 12, 1925,
by Dr. Waldo L. Schmitt.

Paratype. — U.S.N.M. No. 122611, one specimen, 89 mm., taken with
the type and bearing same data.

THREE NEW SCI.VENID FISHES — SCHULTZ 129

Descriptioji. — Certain meapurcmeiits were made, and these data,
recorded below, are expressed in hundredths of the s(an(hutl k'ngth,
fii-st for the holotyjie, then for the paratype in parentheses, respec-
tively. Standard len<rths in millimeters, 77 ^89).

Length of head 33.8 (33.2); greatest depth of body 32.5 (30.8);
diameter of eye 7.53 (7.42) ; length of snout 9.22 (8.70) ; distance
from front of upper lip to rear tip of maxillary 12.2 (11.8); least
preorbital width 4.1G (3.93) ; postorbital length of head 18.8 (18.4) ;
width of bony interorbital space 9.74 (9.10) ; length of caudal peduncle
or distance from base of last anal ray to midcaudal fin base 22.1 (23.G) ;
least depth of caudal peduncle 10.0 (9.r>5) ; length of base of second

Figure 6. — Ophioscion brasiliensis, new species: Ilolotype (U.S.N.M. No. 87742).

doi-sal fin 32.5 (31.2) and of base of anal fin 12.5 (12.9) ; longest dorsal
spine 175 ( — ) ; length of second dorsal spine 12.3 (11.8); length of
second anal spine 16.2 (13.8) ; longest ray of pectoral fin 24.4 (22.5) ;
longest soft ray of pelvic fin 25.1 (25.8) ; length of pelvic spine 12.1
(11.7); longest midcaudal fin ray 2G.0 (23.0); length of longest gill
raker 3.64 (2.81) ; distance from snout tip to dorsal origin 37.1 (36.5)
and to anal origin 68.3 (70.8) ; snout (o pelvic in.sortioii 3r).4 (3"). 5) and
to pectoral insertion 31.4 (33.7).

The following counts were made, respectively : Dorsal rays X-I, 22
(X-I. 21) ; anal rays II, 9 (II, 9) ; pectoral rays ii, 16-ii, 17 (ii, 17-ii,
17) ; pelvics always I, 5; vertical .scale rows counted above lateral lino
•16 (46) and pores in lateral line 45 (46) ; scales from dorsal origin to
lateral line 5 (5) and fi'oni base of first soft dor.^al ray to lateral lino
5 (5) ; scales from lateral jiiie to anal (iriiiin 7 (7) : zig/ag .'-cale rows
around caudal peduncle 17 (17).

Snout blimtly roiuided, projecting a liltic in front of the mouth, tho
latter inferior in [josition; lower jaw included: intciorbital space
broad, a little convex, its width about equnl to Iciigdi of snout; an-

130 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

terior profile from dorsal origin to between eyes nearly straight, or a
very little convex; ventral profile curves to pelvic insertions, then
nearly straight backward to anal origin; body compressed pos-
teriorly; greatest depth of body at dorsal origin; eye 4:% in head,
1% in interorbital space, and 2% in postorbital length of head;
posterior nasal opening close to eye, larger than the anterior one;
tip of lower jaw without barbels but with a median pit contain-
ing two minute pores lying in this porelike depression, and laterally
two pairs of pores as in panamensis; pores and lobes on front of snout
as described for fanaineTisis ; anus a little over two-thirds closer to
anal origin than to pelvic bases; tip of filament of pelvic soft ray
reaching to anus ; pectoral fins reaching to opposite anus ; gill rakers
moderately short, the longest equal to diameter of pupil ; preopercular
spines numbering 9 or 10, none hooked downward, those dorsally
smaller than those near lower angle of preopercle ; skull with the usual
cavernous spaces as found in other genera ; least depth of caudal pe-
duncle a little more than twice in its length ; teeth in villif orm bands in
both jaws, the outer row of upper jaw a little enlarged; pseudo-
branchiae well developed; scales ctenoid; lateral line broadly curved
over pectorals, then running a straight course along midaxis of body
posteriorly, and extending on the caudal fin; the fourth scale row
below lateral line anteriorly is the first one continuing to base of caudal
fin; second dorsal spine only slightly heavier (enlarged) than follow-
ing spines ; second dorsal spine \y^ in second anal spine and reaching
more than halfway to tips of third or fourth dorsal spines and 1% in
postorbital length of head; second anal spine moderately enlarged,
and not reaching to tips of soft rays ; pelvic spine equal to length of
second dorsal spine ; distal margins of all fins a little rounded, that of
caudal fin double truncate, with the middle rays longest.

Color. — In alcohol, pale brownish above, lighter below; anal and
spiny dorsal dusky; pelvic soft rays dusky distally, the filamentous
ray white ; other fins very pale brownish ; peritoneum with numerous
black pigment cells; the types are not well preserved and the colors
have faded.

ReTYiarhs. — This new species of OpMoscion may be separated from
other Atlantic species of this genus by the foregoing key.

Named hrasiliensis in reference to the country along whose shores
the types were collected.

OPHIOSCION MICROPS (Steindachner)

Corvina microps Steindachner. Sitzb. Akad. Wiss. Wien [Ichth. Notizen No. 1],
vol. 49, p. 6, pi. 2, fig. 2, 1864 (Guiaua).

I have made measurements on two specimens from Para (M.C.Z.
No. 4581), and the results are recorded below in hundredths of the
standard length. Standard lengths in millimeters, 66.3 and 60.3.

THREE NEW SCIAENID FISHES — SCHULTZ 131

LenfiTth of head 32.1 and 33.'2; greatest depth of body 30.2 and 31.5;
diameter of eye C.64 and G.47; length of snout 9.95 and 8.62; distance
from front of upper lip to rear of maxillary 11.3 and 11,9; least pre-
orbital width 4.98 and 4.48; postorbital length of head 18.5 and 19.1;
bony interorbital space 10.4 and 11.3; length of caudal peduncle or
distance from ba.se of last anal ray to raidcaudal fm base 26.8 and 24.9;
least depth of caudal peduncle 9.95 and 10.1; length of base of second
dorsal fin 33.2 and 35.6; length of anal fin base 12.5 and 12.6; length
of longest or third dorsal spine 19.6 and 19.8; length of second dorsal
spine 14.0 and 13.4 ; longest soft anal ray 10.6 and 21.5 ; length of second
anal spine 18.1 and 18.6; longest pectoral ray 21.1 and 25.7; longest
soft ray of pelvic fin 23.1 and 23.7 ; length of pelvic spine 12.4 and 13.3 ;
tip of snout to dorsal origin 36.9 and 38.1 ; snout to anal origin 65.4 and
66.8; snout to pectoral insertion 32.1 and 33.3; length of longest gill
raker 2.71 and 2.65.

The following counts were made : Dorsal rays XI-I, 19, and XI-I,
21 ; anal rays II, 9 and II, 9 ; pectoral rays ii, 16-ii, 16 and ii, 17-ii, 16 ;
pelvics always I, 5; gill rakers on first gill arch 11 + 1 + 19 and
11 + 1 + 18; vertical scale rows above lateral line 51 and 50; scales
from dorsal origin to lateral line 5 and 5, and from base of first soft
dorsal ray to lateral line 4 and 4; scales from anal origin to lateral
line 7 and 8; zigzag scale rows around caudal peduncle 18 and 18.
Additional counts are recorded in table 1.

This species has a very small eye, smaller than in any other species.
Diameter of eye is contained in young li/> to 1% and in adults 2 to
2% times in interorbital space.

OPHIOSCION VENEZUELAE. new species

FlOUKE 7

Holotype. — U.S.N. M. No. 121749. one specimen, 139.5 nmi. in stand-
ard length, collected by Leonard P. Schultz near mouth of Cano de
Sagua, 25 km. north of Sinamaica, Venezuela, May 12, 1942.

ParatypcH. — U.S.N.M. No. 121750, six specimens, 57 to 150 mm.,
collected with the holotype and bearing same data.

Description. — Certain measurements were made, and these data, re-
corded below, are expressed i)i liundrcdtlis of the standard length, first
for the holotype and then for the thiee paratypes in i)aren(h('ses, re-
spectively. Standard lengths in millimeters, 139.5 (68.8; 150; 139).

I^^ngth of head 28.6 (30.5; 32.4; 30.2) ; greatest depth of body 30.1
(27.0; 31.2; 30.9) ; diameter of eye 6.24 (7.12; 5.93; 6.11) ; length of
snout 8.74 (8.14 ; 9.34 ; 8.63) ; distance from tip of snout to rear edge of
maxillaries 13.1 (12.6; 13.0; 13.2) ; least width of preorbital 3.05 (3.63;
4.13; 39.5) ; postorbital length of head IH.O (15.8; 17.9; 18.5) ; width of
bony interorbital space 9.68 (9.16; 9.66; 10.3) ; length of caudal pe-

132 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

duncle 25.1 (25.1 ; 24.3 ; 25.4) ; least depth of caudal peduncle 10.7
(9.88; 10.9; 10.9) ; length of base of second dorsal fin 32.6 (32.5; 32.1;
32.4) ; length of base of anal fin 11.0 (11.6; 11.7; 11.2) ; length of long-
est dorsal spine 18.6 (21.1; 18.5; 19.3) ; length of longest soft dorsal
ray — (13.1; —; — ; 13.2) ; longest soft anal ray 16.1 (17.0; — ; 14.7) ;
length of second anal spine 16.3 (17.4; — ; 15.8) ; longest pectoral fin
ray 25.2 (23.1; 22.7; 25.2); longest soft pelvic ray 13.0 (18.6; 12.3;
13.9); length of pelvic spine 9.32 (11.5; 8.34; 8.85); longest or
middle caudal fin rays 25.9 (26.9; 22.7; 25,5) ; distance from tip of
snout to dorsal origin 37.1 (35.9; 38.5; 37.7); snout to anal origin
67.2 (65.4; 65.1; 66.2); snout to pectoral insertion 32.6 (31.1; 32.2;
31.6) ; snout to pelvic insertion 32.6 (30.5; 30.6; 80.9) ; length of long-
est gill rakers on first gill arch 1.58 (2.76; 1.66; 3.22).

Figure 7. — Opkioscion venezuelae, new species: Holotype (U.S.N.M. No. 121749).

The following counts were made, respectively : Dorsal rays XI-I, 21
(XI-I, 22; XII-I, 21; XI-I, 21; XI-I, 21; XI-I, 21; XI-I, 22);
anal rays on all types II, 8; pectoral rays ii, 17-ii, 17 (ii, 16; ii,
l7-ii, 17; ii, 17-ii, 17; ii, 16) ; pelvics always I, 5; number of vertical
scale rows above lateral line 52 (53; 52; 54) ; scales from dorsal origin
to lateral line 6 ( — ; 6; 6) and from base of first soft dorsal ray to
lateral line 6 ( — ; 6; 6) ; scales from lateral line to anal origin 8 ( — ;
8; 8) ; scales in a zigzag row around the caudal peduncle 19 ( — ;
19; 19) ; number of gill rakers on first gill arch 9+1+16 ( — ; 10+1
+ 18; 10 + 1 + 18; 9 + 1 + 16; 10 + 1 + 16).

Head depressed forward but rounded dorsallj^, the interorbital space
convex, broad, about equal to the snout; body compressed; anterior
profile nearly straight but the dorsal contour curved, the ventral con-
tour but slightly curved backward to anus; back highest at base of
spiny dorsal fin ; eye about 2% in postorbital length of head, 14^ in
interorbital space; posterior nasal opening rounded, slightlj^ larger
than the anterior one; tip of lower jaw without barbels; anal origin
equidistant between pelvic insertion and midcaudal fin base; pelvic

THREE NEW SCIAENID FISHES — SCHULTZ 133

fins reaching halfway to anus, tlie first soft ray ending; in a short fihi-
ment; preuperole uith eight or nine short spines, the lowest one
strongest but not hooked downward; caudal peduncle least depth 2V^
in its length; tips of pectoral fins reaching a trifle i)ast anus; teeth in
jaws in bands, the outer row of upper jaw a little enlarged; pseudo-
branchiae well developed; gill rakers short, not quite so long as
pupil diameter; scales strongly ctenoid; lateral line curved over pec-
toral fin, then running a straight course on caudal peduncle along
its midaxis; fourth scale row below lateral line, anteriorly, the first
one extending to base of caudal fin ; first dorsal spine rudimentary,
second 2\-2 in third, the latter nearly as long as the fourth; second
and eighth to eleventh and the next spine heavier than the third to
seventh spines of dorsal fin ; fourth or longest dorsal spine about equal
to postorbital length of head; distal margin of spiny dorsal fin trun-
cate or a very little concave, that of soft dorsal probably a trifle rounded
(the tips of the soft rays are lacking and this cannot be determined
accurately) ; middle rays of caudal fin longest, edges of lobes more or
less truncate to rounded (double truncate) ; distal margins of anal
and pelvic fins a little rounded; pectoral fins somewhat pointed, the
fourth branched ray from above longest.

Color. — In alcohol the upper sides and back are grayish brown,
white below; dorsal, anal, and pelvic fins dusky, more intensely pig-
mented distally ; soft dorsal and cj^udal fins dusky; pectoral fin darker
than other fins except tip of spiny dorsal; lower jaw and upper lip
white; peritoneum white. In the smaller paratypes the dusky upper
sides are broken up with several pale blotches, which appear to have
a small cyst at their centers.

Remar/cs. — This new species differs from all otlicr known species
of OpJiioscion^ excei)t 0. adustus and 0. niicrops, both from the west-
ern Atlantic, in having more numerous gill rakers, 1(> to 18 on lower
part of first gill arch, but adustus has 28 soft dorsal rays and vene-
zuelae only 22 to 24. O. microps has II, 9 anal rays and 0. venczuelae
II, 8. The key will serve for distinguishing the seven species now
recognized in the western Atlantic.

Named venezuclae in reference to the country where the specimens
were collected.

OPHIOSCION I'UNCTATISSIMUS Mick and Hildcbrnnd

OphioHcion punclntUiHimuH Mi;i;k and IIildkhuam), Mariiii' llshi'S of I'niiaina, vol.

2. p. 044, pi. f«. WlTi (Cristobal, Torn Point, and Colon, I'anania).
OphioMcion adunta Evkkmann and Maksii, Fislit's of Porto Kico, U. S. Fish Conim.

Bull., vol. 110 (IDOO), pt. 1. p. 1:H>, lUOli ( Viftpics Island).

I have examined the following specimens in the national collections:
U.S.N.M. Nos. 817GG, the, holofype, and r>()l(;i and 120188, four
specimens from Vieques Island off Puerto Kico.

134 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

U.S.N.M. No. 104297, one specimen, from Recife, Pernambuco,
Brazil.

U.S.N.M. Nos. 80765 and 80766, two paratypes, from Panama.

OPHIOSCION PANAMENSIS, new species
FiGXJEE 8

Ophioscion adustus (in part) Meek and Hiujebrand, Marine fishes of Panama,
vol. 2, p. 639, 1925 (Fox Bay, Colon, Panama).

Holotype. — U.S.N.M. No. 122612, one specimen, 52 mm. in standard
length, collected in Fox Bay, Colon, Panama, January 27, 1912, by
Meek and Hildebrand.

Paratypes. — U.S.N.M. No. 81204, three specimens, 31.5 to 43 mm. in
standard length, from Fox Bay, Colon, Panama, January 5, 1911,
Meek and Hildebrand ; U.S.N.M. No. 81205, four specimens, 23.3 to
30.5 mm.. Fox Bay, Colon, Panama, March 31, 1911, Meek and Hilde-
brand; U.S.N.M. No. 81207, one specimen, 33 mm., from Porto Bello,
Panama, March 17, 1912, Meek and Hildebrand ; U.S.N.M. No. 81206,
one specimen, 42 mm., collected along with the holotype and bearing
same data; U.S.N.M. No. 128260, one specimen, 35.5 mm., from Fort
Sherman, Canal Zone, Panama, collected March 3, 1937, by Dr. S. F.
Hildebrand.

Description. — Certain measurements were made, and these data,
recorded below, are expressed in hundredths of the standard length,
first for the holotype, then for a paratype in parentheses. Stand-
ard lengths in millimeters 52 (42).

Length of head, 33.3 (34.3) ; greatest depth of body 32.7 (32.2) ;
diameter of eye 7.30 (7.86) ; length of snout 9.80 (9.76) ; distance from
front of upper lip to rear tip of maxillary 10.4 (11.4) ; least preor-
bital width 4.80 (4.76) ; postorbital length of head 17.9 (19.0) ; width
of interorbital space 9.62 (10.2) ; length of caudal peduncle or dis-
tance from base of last anal ray to midcaudal fin base 22.3 (23.1) ;
least depth of caudal peduncle 11.2 (10.7) ; length of base of second
dorsal fin 35.4 (35.7) and of base of anal fin 10.6 (11.9) ; longest dorsal
spine 18.3 (16.4) ; longest soft ray of dorsal fin — (7.2) ; longest soft
ray of anal fin 20.8 (18.6) ; length of second anal spine 18.8 (19.3) ;
longest ray of pectoral fin 23.1 (22.4) ; longest soft ray of pelvic 21.7
(25.0) and of pelvic spine 12.1 (12.6) ; longest midcaudal fin ray 30.0
(31.0) ; length of longest gill raker 2.11 (2.38) ; distance from snout
tip to dorsal origin 38.3 (39.3) and to anal origin 71.9 (67.4) ; snout
to pelvic insertion 36.0 (35.6) and to pectoral insertion 34.6 (32.9).

The following counts were made, respectively : Dorsal rays X-I, 20
(X-1, 21) ; anal rays II, 7 (II, 7) ; pectoral fin rays ii, 17-ii, 17 (ii, 17-ii,
17) ; pelvics always I, 5 ; scale rows above lateral line 51 (52) and pores
in lateral line to midcaudal fin base 50 (50) ; scales from dorsal origin

THREE NEW SCIAENID FISHES — SCHULTZ 135

to lateral line 5 (5) and from base of first soft ray of dorsal to lateral
line 5(5); t^cales from lateral line to anal origin 9 (8) ; zi<;za<j^ scale
ro>Ys around caudal peduncle 19 (17). Additional counts are recorded
in table 1.

Snout bluntly rounded, projecting a little in front of mouth, the
latter inferior in position, lower jaw included; interorbital space
broaiK a little convex, its width about equal to length of snout ; anterior
profile nearly straight from dorsal origin to between eyes or a trifle
convex ; ventral profile a little convex anteriorly, then nearly straight
to anal origin; body compressed posteriorly; greatest depth at dorsal
origin; eye about 4V'> in the head, 21/5 m postorbital length of head,
and 1V4 ill interorbital space; posterior nasal opening close to eye, a

,„-^*!»"?^*'S!><>

MiM.

Figure 8. — Ophioscion panamensis, new species; Holotype (U.S.N.M. 122612).

little larger than anterior one; tip of lower jaw without barbel, but
with three pairs of pores, the median pair minute and in a porelike
depression; tip of snout with two pairs of lobes, a pore lying between
the middle pair of lobes and the outer pair being separated from the
median lobes by a pore on each side, a third groovelike pore lying lat-
erally to the outer lobe; then dorsally to the margin of the snout occur
a median pore and another pore at each side; anus two-thirds closer to
anal origin than to pelvic insertion; j)elvic fins not quite reaching to
anus; the first soft ray of pelvics ending in a short filament; pectoral
fins reafhing opposite tip of pelvics; gill rakers short, a little less than
one-half the piij)il; preopercle with eight or nine small spines, those
dorsally snmllei- th.m those near the lower angle of preopercle; none
hooked downward; skull with the tisual ()|)en spaces or sinuses between
the narrow bony biidges; least depth dl" caurjal peduncle l^^jo i" its
iength; teeih in villiform bands in both jaws, the outer row of upper
jaw slightly enlarged; i)seudol)ran<'hiae well developed; scales
strongly ctenoid; lateral line broadly curved over pectorals, then run-
ning a straight course along midaxis of body posteriorly, extending on

136 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

caudal fin ; the fifth scale row below lateral line anteriorly is the first
one continuous to base of caudal fin ; second dorsal spine enlarged, its
length ly^ in second anal spine, and reaching more than halfway to
tip of third or fourth dorsal spine; second dorsal spine 1% in post-
orbital length of head; second anal spine enlarged, not reaching to
tips of soft anal rays ; pelvic spine a trifle shorter than second dorsal
spine ; distal margins of dorsal, anal, and of paired fins a little rounded,
that of the caudal fin double truncate, the midcaudal rays longest.

Color. — In alcohol the body is brownish everywhere, paler brown
ventrally; paired fins and anal and dorsal fins blackish, with the
first spine in these fins, except pectorals, whitish ; tip of first soft ray
in pelvic fins white, especially the filament ; soft dorsal and caudal fins
brownish with numerous black pigment cells; undersides of head
and breast pale ; lips pale ; peritoneum white.

In the smaller specimens of this species the coloration of the median
fins differs from the larger ones. At a standard length of 24 mm.
the caudal fin is white, except for a few scattered brown pigment cells
located near the center of the fin, the caudal fin base is abruptly dark
brown with the pigment extending backward a little on middle rays ;
the anal and dorsal fins have a brownish band extending across rays,
with the margin of the fins white, and below this bar is another white
area separating the brownish base of these fins from the brown band ;
these fins gradually fill in with brown pigment so that at 42 mm. the
fins are plain brownish.

Bemarks. — This new species may be separated from other Atlantic
species of Ophioscion by the key and traces down to O. adustus in
Meek and Hildebrand's key to the species of Ophioscion in their
"Marine Fishes of Panama" (vol. 2, p. 636, 1925) .

Named panamcnsis in reference to the region where it has been
collected.

OPHIOSCION NASO (Jordan)

Stelliferus naso Jordan, in Jordan and Eigenmann, Rep. U. S. Comm. Fish and
Fisheries for 1S86, vol. 14, p. 395, 1889 (Cachiura, Brazil).

I have made measurements on two of the types (M.C.Z. No. 4583)
from Cachiura, and the results are recorded below in hundredths
of the standard length. Standard lengths in millimeters, 75.5 and
70.5.

Length of head 31.1 and 30.5; greatest depth of body 31.8 and 28.4;
diameter of eye 8.60 and 9.22 ; length of snout 9.27 and 9.22 ; tip of pre-
maxillaries to rear of maxillary 10.1 and 9.78 ; least width of preorbital
3.97 and 3.97; postorbital length of head 15.9 and 15.6; least width
of bony interorbital 9.14 and 9.22 ; length of caudal peduncle (base of
last anal ray to midbase of caudal fin) 25.0 and 24.4; least depth of
caudal fin 10.6 and 10.8; length of base of second dorsal fin 31.8 and

THREE NEW SCIAENID FISHES — SCHULTZ 137

36.2; leno^th of anal fin base 12.4 and 12.1; lonf^th of longest dorsal
or tiiird spine 22.3 and 21.0; longest soft dorsal ray 17.2 and — ;
length of second dorsal spine 10.6 and 11.3; longest soft ray of anal
fin 19.9 and 17.9; length of second anal spine 17.0 and 16.5; longest
ray of pectoral fin 2r),2 and — ; longest soft ray of pelvic fins 21.2
and 21.3; length of pelvic spine 12.6 and 12.3; longest or middle rays
of caudal fin 28.2 and 28.4; tip of snout to dorsal origin 38.3 and
37.2; snout to anal origin GG.2 and 68.8; snout to pectoral insertion
31.8 and 31.9; longest gill raker 1.99 and 1.99.

The following counts were made, respectively : Dorsal rays XI-I, 21
and XI-I, 21; anal rays II, 8 and II, 8; pectoral rays ii, 16-ii, 16 and
ii, 16-ii, 16; pelvics always I, 5; gill rakers on first gill arch 8 + 1 + 14
and 8 + 1 + 14; vertical scale rows above lateral line 46 and 46; scales
above lateral line at origin of spiny dorsal fin 4 and 4, and at base of
first soft dorsal ray 5 and 5; scales below lateral line from anal
origin to lateral line 8 and 8; zigzag scales around caudal peduncle
18 and 18.

u. t. «ov(iiNiiiNi rniNTiNs orricci i*4f

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM

Vol. 96 Wwhington: 1945 No. 3193

THE ICHNEUMON-FLIES OF THE GENUS CRYPTANURA
liRULLE, MAINLY TROPICAL AMERICAN

P>V R. A. CUSIIMAN

Because of BruUe's juistukcn idcu tluit the specimens on which he
based the genus C ryptanxLva were fenuiles "with concealed ovipositors,
this genus of ichneumon-flies was misunderstood until Roman (1910)
identified it as the male of Polydeinis Cresson and synonymized the
latter genus with Crijptanura.

I'he genus dates from 1845, when the figure of Cryptanura nigripes
Brulle was published in the atlas of Lepeletiers 'Tlistoire Natnrelle
des Insectes," although tlie description did not appear until the fol-
hjwing year. It is therefore a monobasic genus with nigripes Brulle
as g<'notype. On this basis, and assuming that nigr/pcfi and .sfriafa
Brulle are not congeneric, Viereck took exception to Roman's synon}'-
mizing of Polyaenttfi Cresson with Cryptanura. Viereck's statement
that these two species are not congeneric must have been based on the
fact that Brulle does not state definitely that nigripes has the two
small frontal horns characteristic of striata; but by inference he
certainly does ascribe this character to nigripes^ for in the description
of all the species that follow sfn'tifa he mentions only the cliuiinters
by which they differ from it and from one another.

Cresson and Cameron were in error in their inter])retati()ns of the
genus, the species assigned to it by those authoi's Ijeing properly re-
ferred to such genera as Trapr.zonaHs Szcpligeti, GUxlianus Cameron,
I'liotocryptuft Viereck, and prol):ibly other genera. Roman and Brues
iippear to be the only ones who have interpreted the genus correctly.

624514—45 1 130

140 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9«

Cryptanura is a large genus apparently confined in its distribution
to the Western Hemisphere and there very largely to the tropical
region, only two species being known to occur north of the Mexican
border.

Three species from the Old World Tropics have been referred to
Polyaenus: cingvlatus Tosquinet from New Guinea, spiniferus Cam-
eron from Borneo, and striatum Szepligeti from Formosa, but none
appears to be properly referable to the genus. This is certainly true
of cingulatu^ and spiniferus^ for I have been able to identify the
latter definitely and the former without much question in the Baker
collection. These si)ecies represent an apparently hitherto unde-
scribed genus. It is described in an addendum to this paper.

I have been unable to identify striatus Szepligeti, but its smooth
and polished mesoscutum, longitudinally impressed propodeum with
the apophyses apparently represented only by carinae, and elongate
postpetiole would seem to exclude it from Cryptanura. Aside from
the new genus mentioned above, to which striatus obviously does not
belong, only one other Oriental genus is known to me that has two
frontal horns. This is Ceratocryptus Cameron, in which striatus
appears to be equally out of place.

SPECIES WRONGLY REFERRED TO CRYPTANURA

The following species described in Cryptanura by Cresson and
Cameron do not belong to the genus. Unfortunately, most of them
are unknown to me. The apparently proper status of the few that I
have been able to place is indicated.

{Cryptanura acolhua Cresson) =Glodianns acolhua (Cresson), new combi-
nation.

C albispina Cameron.

C. cinctipcs Cameron.

C. curtispina Cameron.

C. delecta Cresson.

C. fascia tipennis Cameron.

C. incauta Cameron.

C. interrupta Cameron.

C. laticarinata Cameron.

C. ornatipennis Cameron.

(C pachymene Cresson) =Pliotocryptus pachymene (Cresson).

C. pedicata Cameron.

(0. sumichrasti Cvesson)—TrapezonaUs suniichrasti (Cresson), new combi-
nation.

Genus CRYPTANURA Brulle

Cryptanura Brulle, Histoire naturelle des insectes : Hymenopteua, Atlas, pi. 41,
fig. 6, 1845; vol. 4, p. 242, 1846.— Roman, Ent. Tidskr., 1910, p. 154.— Brues,
Ann. Ent. Soc. Amer., vol. 5, p. 200, 1912. — Townes, Mem. Amer. Ent. Soc.
No. 11, pt. 1, p. 288, 1944.

ICHNEUMON-FLIES OF GENUS CRYPTANUIU — CUSHMAN 141

Pulyacnus Crksson, Proc. Acad. Nat. Scl. riiihulolphia, 1873, p. 149.— Cameron,
Kiologia Cvatrali-Aiufricana, Ilyuit'iioptfra, vol. 1, p. 244, 188G; Journ. Hoy.
Agr. Coium. Soc. British Guiana, sor. 3, vol. 1, p. 16G. 1911.— St u.\nEm:KNKcnT,
Genera insoctoruui, fasc. 7r>, p. G7, 19<,)8. — Sz^i-luieti, Ann. Mns. Nat. Hun-
garici, vol. 14, p. 264, 1916.— Cushman, Proc. U. S. Nat. Mus., vol. 74, art. IG,
p. 38, 1929.

Polyacnidia Viekeck, Proc. U. S. Nat. Mus., vol. 46, p. 381, 1913.

Hcsostcnus authors, part.

Head from above transverse, the temples receding; frons with a
carina medially and with two small horns, frequently arising from a
common base; malar space distinct; clypeus strongly convex; eyes
large and strongly convex; antenna in female frequently more or
less distinctly thickened between middle and apex, the thickened por-
tion flattened below. Thorax stout; epomia distinct and usually ex-
tending to upper margin of pronotum, where they form carinate or
conical projections; nolaulices deep and complete; scutellum small^
usually convex, rarely subconical or flattened ; propodeum with basal
carina complete, apical carina usually represented only by two prom-
inent apophyses, sometimes, especially in mule, distinct and witli
apophyses less developed ; propodeum usually strongly rugose or
transver.Ncly striate, rarely without sculpture, spiracles elongate;
wings large; areolet small, complete, quadrangular, broadening to-
ward apex, recurrent interstitial or somewhat antefurcal ; nervulus
antefurcal; second discoidal cell broad at base; nervellus broken near
bottom and perpendicular or weakly reclivous; logs long and usually
rather slender; front tibia in female rarely slightly inflated. Abdo-
men in female fusiform, in male small and narrow; spiracle of first
segment far beyond middle; ovipositor sheath from a half to fully as
long as abdomen; ovipositor subsagittate or swordlike at apex.

Head and thorax ornamented with white or 3'ellow on a black or
red ground; abdcjmen black and yellow or largely red; wings immacu-
late hyaline or dilutely infumate.

The color pattern of the head and thorax nearly throughout the
genus is so similar as to constitute almost a generic character. In
order to avoid much repetition a description of what may be called
the normal or basic color pattern is gi\en here. In the specific descrip-
tions only variations from this pattern are indicated.

Ground color of head and thorax black, rarely red or partly red,
with pale-yellow or whitish markings as follows: Oibilal ring, broad
on cheek, much narrower or interrupted on upper temple and in malar
space; face, clypeus (apical margin always dark), labrum, mandible
basally, palpi, aninilus on antenna; pr()i)leura more or less; anterior
and humeral margins of pronotum; meso.scutum either immaculate,
or with a single spot on disk, or with paired lines along inner margins
of lateral lobes, or with a nairow line on each lateral margin oppo-

142 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9e

site tegulae, or with cuneiform markings on anterior margins of lat-
eral lobes, or with combination of two or more of these; scutellum,
its subtending carinae, and posterior margin of its f renum ; postscutel-
lum and margin of its f renum ; tegulae ; subalar tubercles ; an oblique
band on mesopleuron from near anterior margin to middle coxa; a
larger or smaller spot on each side of mesosternum; upper division
of metapleuron and its lower division largely, the latter with the
ground color showing only below; and two broad marks posteriorly
on propodeum embracing the apophyses, very rarely confluent ante-
riorly. In the descriptions this color pattern is referred to as black
(or red) with yellow or white markings and the exceptions noted.

The abdomen is either largely red or the ground color is black, typi-
cally with broad apical and lateral margins of tergites 1-7, lateral
margins of tergite 8, and the petiole more or less yellow or whitish ;
venter white with sternites darker. In the following descriptions this
color pattern is referred to merely as black, the tergites margined
with yellow and the exceptions noted.

SPECIFIC CHARACTERS

For the most part Cryptanura is a very homogeneous group with
few striking structural characters that will serve to divide it into spe-
cific groups, and it has been found necessary to depend to a consider-
able extent on color in the construction of the following key to species.

The form of the scutellum, the structure of the propodeum and
pronotum, and the form of the ovipositor have been found useful as
group characters, though the last, of necessity unisexual, does not
appear to be accompanied by a companion character in the opposite
sex.

A considerable number of the described species have not been avail-
able for study and have been omitted from the following key, but to
assist in the identification of these species I have constructed a key to
all the species, based largely on color. This will be found following
the descriptions of the species examined.

KEY TO SPECIES EXAMINED

1. Scutellum strongly subconically elevated; metapleuron also with a tuberculate

elevation 1. tuberculata, new species

Scutellum and metapleuron not strongly elevated 2

2. Apical carina of propodeum distinct medially at level of apophyses ; abdomen

polished and unsculptured 3

Apical carina usually absent, sometimes distinct in male, but then arching
high above apophyses ; abdomen usually distinctly, finely sculptured— 5

3. Abdomen and propodeum entirely red 2. dicostata, new species

Abdomen and propodeum black and yellow 4

4. Mesoscutum with yellow marginal markings, but without discal markings- 5
Mesoscutum with paired discal markings, but without marginal markings.

5. bicarinata, new species

ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 143

5. Mesoscutal markings in furm of narrow marginal linos opposito togulae;

torgite 2 yellow at apex, the j-ellow band broader medially; tegnlae yellow

basally 3. quadrimaculata, new species

Mesoscutal markings in form of cuneiform spots laterad of origins of nolau-
lices; tergite 2 with a uniformly broad subapieal yellow band; tegulae
entirely black 4. mediostrigosa, new species

6. Ilunjeral margin of pronotum conicnlly prominent anteriorly; scutellum

broad and transversely flat, with coarse, deep punctures; abdomen pol-
ished, without trace of sculpture; front tibia in female subinflated; hind
femur stout, not or barely two-thirds as long as tibia; pale markings of

propodeum extending broadly forward to basal carina 7

Disagreeing with all or nearly all above characters 10

7. Abdomen black and yellow (or white) 8

Abdnmen largely or entirely re<l 9

8. Legs red, coxae marked with yellow; thorax reddish piceous and yellow.

6. piceothorax, new species
Legs and thorax black and yellow 7. planiscutellata, new species

9. All coxae and front and middle femora black and yellow.

8. politigaster, new species
Hind coxae and aJl femora red 9. conica, new species

10. EiKimia not extending upward to humeral margin of pronotum, the humeral

njargin not at all carinate or tuberculate anteriorly ; ventrolateral carina
of petiole distinct to base; second tergite with a median white spot at

base 11

EiK>mia forming a carinate elevation on humeral margin of pronotum ; ventro-
lateral carina not distinct to base; second tergite without a median white
spot 12

11. Ht-ad and thorax red and yellow; antenna without white annulus; abdomen

polished 10. ruficeps, new species

Head and thorax black and yellow; antenna with white annulus; abdomen
beyoml first tergite opaque shagreened, second and third tergites also

punctate 11. septentrionalis, new species

IZ M'soscutum with two yellow marks discally 13

Mesoscutum with a single median spot, rarely flanked by small traces of

yellow on inner margins of lateral lobes 16

L3. Petiole and hind femur entirely black 12. apophysis, new species

Pftiole and hind femur partly yellow 14

H. I'ropodeal yellow spots not abruptly narrowed before apophyses, the latter

long 15

Proixtdeal yellow spots abiniptly narrowed before apophyses, the latter

short 15. bilineata, new si>ecie3

IT). Petiole yellow above, black bilow ; Iiiiul femur entirely black above.

13. mexicana (Cresson)
Petiole Itlack above, yellow below; hind femur with a narrow median yellow

line above 14. orizabensis (Canjeron)

10. Thorax red and yellow 16. rufa, new species

Tlt'irax black and yellow 17

17. Proiwideul spots each with a narrow forward extension, sometimes (male)

confluent medially 18

ProiKxleal .sjwt.s either not extending forward or not abruptly narrowed
before ap^iphyses 22

18. Abdomen black and yellow 19

Abdomen largely or entirely red 20

144 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m

19. Front and middle femora red ; hind femur red ( $ ) or entirely black (S) ;

hind coxa red ( 9 ) or black ( $ ) with a yellow spot above.

17. spinaria (Brull6)

All femora black and white, hind femur white below, black above, with a

narrow median white line; hind coxa white below and above, black

on each side 18. lineatifemur, new species

20. Hind coxa black and white 19. coxata, new species

Hind coxa largely or entirely red 21

21. Humeral margins of pronotum prominent, subtuberculate anteriorly; post-

petiole red . 20. boliviensis, new species

Humeral margins of pronotum merely carinate anteriorly, not prominent;
postpetiole piceous, margined with yellow 21. isthmus, new species

22. Ovipositor nearly as long as abdomen, stout, much deeper toward apex than

at base, not sagittate at apex 23

Ovipositor slender, its upper and lower margins parallel except at the sagit-
tate apex, usually much shorter than abdomen 25

23. Abdomen red 22. excalibur, new species

Abdomen black and white 24

24. Hind coxa largely and femur red 23. acinaces, new species

Hind coxa yellow and black ; femur black or piceous above, yellowish below.

24. propinqua (Cresson)

25. Front and middle femora entirely red 25. pretiosa (Viereck)

Front and middle femora piceous posteriorly 26

26. Hind leg very slender, femur apparently nearly 8 times as long as deep;

temples in dorsal view very strongly receding and slightly concave.

26. gracilipes, new species

Hind leg stouter, femur apparently not more than 6 times as long as deep;

temples flat or weakly convex or concave 27

27. Temple at middle of eye distinctly more than half as broad as short diame-

ter of eye and very weakly convex 28

Temple not or barely half as broad as short diameter of eye and fiat or weakly
concave 29

28. Head in side view with occipital carina very nearly parallel to posterior

margin of eye ; apical margin of clypeus weakly curved.

27. gracilis, new species

Occipital carina and posterior margin of eye distinctly divergent below;

apical margin of clypeus perfectly straight 28. genalis, new species

29. Hind femur rather slender, at least 6 times as long as deep and more than

three-fourths as long as tibia ; flagellum in female much thickened beyond

middle and flattened below, the joints there strongly transverse 30

Hind femur rather stout, distinctly less than 6 times as long as deep and
less than three-fourths as long as tibia ; flagellum in female only slightly
thickened and flattened beyond middle, the joints there weakly trans-
verse 32

30. Occipital carina nearly parallel to posterior margin of eye ; ovipositor sheath

hardly as long as abdomen beyond first tergite 29. variegata (Brull6)

Occipital carina and posterior margin of eye distinctly divergent below;
ovipositor sheath nearly as long as abdomen 31

31. Temple at middle of eye distinctly less than half as broad as short diameter

of eye ; flagellum very slender at base, first joint more than 6 times as long

as thick at middle 30. paranensis, new species

Temple about half as broad as short diameter of eye ; flagellum stouter, flrst
joint not 6 times as long as thick at middle.

31. tenuiterebrata, new species

ICHNEUMON-ILIES OF GENUS CHYPTANURA — CUSHMAN 145

32. Face entirely yt'Uow ; frttnt and middle coxae piceous behind, yell*)w in

front, metathorax not at all red 32. incerta (Cresson)

Face with longitudinal iniprt'ssions broadly black ; front coxa largely
piceous, middle coxa entirely red; nielai)lenron below and nietasternum
red 33. maculifrons, new siiecies

1. CRYPTANURA TUBERCULATA, new speciei

Evidently clctsely allied to .scutellaris (Szopligeti) and possibly syn-
onymous with tliat species. However, it seems unlikely that 8zepli-
geti would have failed to mention tlie very evident tuberculiform
metapleura charaeteristic of the present species if his species were of
similar structure.

Female. — Length 19 nun., antenna 17 mm., ovipositor sheath 8 mm.

Head in dorsal view with temples very slightly concave, occipital
carina flangelike, sinuate at lower extremity; frontal horns very
small; scrobes very deep; a distinct, radiately rugose impression sur-
rounding the ocelli except behind; postocellar line hardly as long as
ocellocular line, stemmaticum longitudinally rugose; eyes parallel
within, faintly sinuate opposite frons; face with a median subhemi-
spherical elevation flanked on each side by a transversely rugose im-
pression; clypeus smooth, with scattered punctures, roundly convex
with a broad reflexed margin, broadly truncate; malar space very
nearly as long as basal width of mandible ; antenna only slightly thick-
ened beyond middle, 38-jointed. Thorax polished, with notaulices,
margins of mesoscutum, sternaulices, and groove along posterior mar-
gin of mesopleuron foveolate; scrobe of pronotum, upper anterior por-
tion of mesopleuron, metapleuron, and sides of propodeum obliquely
striate; mesopleuron below and sternum sparsely punctate; humeral
margin of pronotum smooth, anteriorly tuberculate; scutellum very
strongly elevated, subconical, in profile with its posterior slope deeply
concave ; metapleuron with a high rounded tubercle in the middle ; legs
very long and slender, hind femur reaching distinctly beyond apex
of abdomen and fully eight times as long as deep; coxae polished and
sparsely punctate. Abdomen almost exactly as long as head and
thoi-ax, rather narrow, very finely shagi'eened, subopacjue beyond first
tergite; i)etiole distinctly depressed; ovipositor ratiier stout, becoming
gradually slightly deeper toward apex, where it is swordlike rather
than subsagittate.

Head and thorax black with yellow maculation (see description of
color patern, p. 141), abdomen ferruginous, legs ferruginotis and
])arlly yellow and piceous; wings stibhyaline, with blackish venation;
orbital ring interrupted on upper temples and in mahir sjiace; facial
impressions and outline of clypeus black; anmilus on antenna embrac-
ing flagellar joints (5)r)-12( b'J) ; two yelU)w lines on disk of mesos-
cutum; apex and anterior face of the elevated part of scutellum black;
sternum yellow only along sternaulices; lower division of metapleuron

146 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

with only the tubercle and its apex yellow ; stripes on propodeum ex-
tending only a short distance basad of apophyses. Front coxa behind
and all femora behind piceous, this color reduced on hind femur to a
narrow stripe on the inner side and the apex; front legs otherwise yel-
lowish, as are also the middle coxa, tibia, and tarsus, the hind tibia
except at apex, and a stripe on inner side, which are blackish, and the
hind tarsus. Ovipositor sheath black,

Tyfe locality. — St. Laurent du Maroni, French Guiana.

Types. — A holotype and a paratype, both females, taken at the above
locality by Audoit in 1862. The type is in the Paris Museum ^ and the
paratype in the U. S. National Museum (No. 57080) .

The paratype is a little larger than the type but is otherwise like it.

2. CRYPTANURA DICOSTATA, new species

Apparently related to shnilis (Szepligeti) and perhaps the same
but having the apical carina of the propodeum more or less distinctly
developed, a character not mentioned by Szepligeti.

Female. — Length 12 mm., antennae 11 mm., ovipositor sheath 4 mm.

Temples very narrow and strongly receding, flat; occipital carina
neither especially prominent nor sinuate at lower extremity; vertex
and frons deeply concave, ocelli distinctly below level of superior
tangent of eyes, frontal horns small; face nearly flat, transversely
rugulose and very minutely shagreened, subopaque; clypeus strongly
convex, with very narrow reflexed margin, polished, with sparse
punctures; malar space about two-thirds as long as basal width of
mandible, finely opaque; head elsewhere polished; postocellar line
and diameter of an ocellus about equal and much shorter than ocel-
locular line; antenna 33-jointed, slender, only slightly thickened and
flattened beyond middle. Thorax anteriorly subopaquely sculptured,
posteriorly polished and almost without sculpture ; humeral margin of
pronotum smooth, anteriorly angulate, scrobe striate; mesoscutum
punctate, prescutum transversely striate on each side next to the
notaulices, lobes flattened; scutellum convex, polished, with a few
punctures ; mesopleuron striatopunctate above, punctate below, smooth
posteriorly; sternum punctate; metapleuron polished and sparsely
punctate, with a few rugae posteriorly; propodeum smooth and
polished, more or less roughened medially between carinae; apical
carina distinct and straight between apophyses; basal carina only
slightly curved medially ; basal area not at all defined ; legs long, hind
femur about six times as long as deep and reaching slightly beyond
apex of abdomen; coxae polished. Abdomen distinctly longer than
head and thorax, polished, unsculptured, rather slender ; first segment
without trace of dorsal carinae, petiole slightly broader than deep,

1 All specimens indicated as being in, or received from, the Paris Museum are retained
In the U. S. National Museum for the duration of the war.

ICHXEUMOX-FLIES OF GENUS CRYPTANURA — CUSHMAX 147

I>ostjietiolo barely as ^vi(le as lonp:; second teririto as ]oujz ^^ fii'^t and
about twice as lon<j: as broad at base; ivopositor sheath shorter than
abdomen beyond first segment; ovipositor slender, of even depth
throughout, apex weakly subsagittate.

Head and anterior portion of thorax black with yellow markings
(see description of color pattern, p. 141) ; thorax posteriorly, mesoster-
num, abdomen, and legs ferruginous; antennal annulus embracing
flagellar joints (4) 5-11 (12), two narrow yellow streaks on disk of
mesoscutum, scutellum yellow only in basal angles and at apex, meso-
pleuron yellow except the prepectus, the impression below tubercle,
and an oblique streak running down from this, which are black;
upper division of metapleuron yellow tinged with ferruginous; propo-
deum, lower section of metapleuron, and sternum, except prepectus,
ferruginous; front coxae anteriorly and all tibiae and tarsi yellow;
front and middle femora piceous below and behind; wings yellowish
hyaline, venation dark brown; abdomen entirely ferruginous, sheath
biack.

7^l/pe locality. — Kamakusa, British Guiana.

Type!i. — Four females, the holotype and three paratj'pes in the
U. S. Natitmal Museum (No. 57058).

The type and two paratypes were taken at the type locality by H.
Lang. The third paratype is from Port-of-Spain, Trinidad.

One specimen of what is probably the male of this species was taken
by W. M. Mann at Cavinas, Beni, Bolivia, during the Mulford Bio-
lr)gical Exploration of 1921-22. It has the mesopleuron and sternum
almost entirely yellowish stramineous, the antennal annulus embrac-
ing fl.igellar joints (5) 6-19 (20), and the apical carina of propodeum
only faintly indicated medially. It is excluded from the type series.

8. CRYPTANURA QUADRIMACULATA. new species

Apparently closely' allied to cctypa (Cresson), from the description
of which it differs in its possession of yellow lateral lines on the meso-
scutum. Also Ciesson makes no mention of the presence, in his
species, of the apical carina of the propodeum.

Frmalr. — Length 13 mm., antennae (broken), ovipositor sheath 4.5
nun.

Temples flat, short and very sharply receding, occipital carina
moderately prominent, bent abruptly inward at its lower extremity
to meet the hypostomal carina far back from base of mandible; vertex
and frons concave, lateral ocelli slightly below superior tangent of eyes,
frontal horns very acute and arising from common base; postocellar
line and diameter of an ocellus equal and distinctly shorter than ocel-
locuiar line; eyes large and bidging, weakly fonvergent lielr»\v ; 'i\n-\.' :md
rlyjieus minutely shagreened, sub()j)aque, face laterally and dyiu'Us

C24.'>14— 45 2

148 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

sparsely punctate, face dorsally irregularly striate; clypeus strongly
convex, in profile subnasute; malar space hardly two-thirds as long
as basal width of mandible. Thorax rather slender, generally polished ;
pronotum striate in lower part of scrobe, its upper anterior margin
only weakly tumid, smooth, carinately angled; mesoscutum sparsely
punctate, rugose in posterior middle, prescutum more densely punctate
and transversely striate anteriorly along notaulices, lobes flattened;
mesopleuron largely and sternum sparsely punctate, space below tu-
bercle more or less striate ; scutellum convex, with scattered punctures ;
metapleuron very sparsely punctate and with a few vertical striae
posteriorly; propodeum with sparse, coarse punctures basally and
more or less distinctly longitudinally striate medially behind basal
carina, basal area obsoletely defined, basal carina bent sharply for-
ward medially, apical carina distinct between apophyses ; legs slender
but hind femur less than two-thirds as long as tibia and not reaching
apex of abdomen ; coxae elongate, polished, and sparsely punctate, are-
olet narrow. Abdomen longer than head and thorax, polished, with
scattered punctures on basal tergites ; first tergite narrow, petiole not
depressed, dorsal carinae absent, postpetiole barely as broad as long,
with a median impression between spiracles; second tergite as long
as first and nearly twice as long as broad at base ; sheath nearly as long
as abdomen beyond first tergite ; ovipositor slender, of nearly uniform
depth, and subsagittate at apex.

Black and yellow (see description of color pattern, p. 141) ; cheeks,
frontal orbits, an annulus beginning on flagellar joint 6 (missing
beyond joint 8) yellow; anterior margin of pronotum black except
for a small spot ; humeral margin entirely black ; narrow lines on lateral
margins of mesoscutum above tegulae, only basal angles and apex of
scutellum, tegulae basally, and mesopleuron except prepectus largely
yellow, the last confluent with mark on sternum, metapleuron except
suture between upper and lower divisions and a dash near apex en-
tirely yellow, large yellow spots in basal lateral areas of propodeum,
apical markings of propodeum with triangular projection anteriorly,
apical bands on basal tergites triangularly broadened medially;
petiole yellow above, black below ; venter yellow. Legs yellow, with
front and middle coxae behind, hind coxa within and medially above,
middle trochanter and femur behind, hind trochanter largely, and
femur except a yellow streak on each side black or piceous; wings
hyaline, venation blackish.

Male. — Essentially like female, the frontal horns much longer.

Type locality. — San Bernardino, Paraguay.

Types. — A female holotype, an allotype, and a female paratype in
the U. S. National Museum (No. 57059). All were collected at the
above locality by K. Fiebrig.

The paratype female is essentially like the type.

ICHNEUMON-FLIES OF GENUS CRYPTANURA CUSHMAN 149

4. CRYPTANURA MEDIOSTRIGOSA, new species

Female. — Length 14 mm., antenna 12 mm., ovipositor sheath 4.5 mm.

Differs from the above description of qiuulrimaculata principally
as follows: Mesopleuron polished, with a small area of rather coarse
j)unctiires anteriorly; metapleuron rather coarsely* punctate-rugose
over most of its surf race; propodeum polished basally, longitudi-
nally striate medially between the transverse carinae and in i)etiolar
area; hind femur fully two-thirds as long as tibia and fully reaching
apex of abdomen; abdomen polished and entirely without sculpture;
petiole depressed; postpetiole fully as broad as long, flat between
spiracles.

Color pattern as in quadrim-amdata except frontal orbit broader
and extending over top of eye; antennal annulus beginning on joint
5; anterior margin of pronotum entirely black, humeral margin with
a white mark at the angle; mesoscutum with cuneiform markings
extending bacJvward from origins of notaulices; tegulae entirely
black; mesopleuron and sternum and metapleuron and sternum stra-
mineous rather than yellow, this color including also most of the pre-
pectup; iJropodeum dorsally yellow except for a cruciform black
mark with its crossbar on the basal carina ; yellow bands of abdomen
subapical and of nearly uniform width; front and middle legs yel-
low, with femora posteriorly and apical half or more of tarsi black-
ish; hind leg stramineous, trochanter and femur black above.

Male. — Essentially like female.

Type locality. — Barro Colorado Island, Canal Zone.

Types. — Two females collected at the type locality by S. W. Frost,
the holotype on March 10, the paratype on March 22, 1937; and one
male allotype taken by A. Busck on March 12, 1912, at Alhajuela,
Canal Zone. All are in the U. S. National Museum (Xo. 57UC0).

S. CRYPTANURA BICARINATA, new fpecies

Though related to qu/idrhnaculata in its possession of the distinct
apical carina and in its highly polished abdomen, this species is very
tlistiiict, especially in the minute frontal horns, stouter body, and
ditferi'ut arrangement of color. It seems uidikely that this can be
championi (Cameron), also from Panama, for it is apparently much
stouter and has the black on (he inner side of the hind fcnuir rather
than on the ui)per side. Moreover, Cameron does not mention the
I)resenco in his species of the apical carina of the propodeum.

Fnnah. — Li'ngth 13 mm., an(enna«' 12 nun., ovipositor sheath 5.5
mm.

Temples flat, narrow, sharply receding, occipital ciuiiia rather
prominent, ils lower extremity curving slightly to join the hyposlo-
mal carina back from the base of the mandible; vertex and frons con-

150 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

cave, ocelli slightly below superior tangent of tlie eyes ; frontal horns
obsolete, their common base distinct ; eyes large, bulging, very faintly
convergent below ; face rather flat, with a transversely striate impres-
sion on each side of middle, the upper striae extending across middle,
laterally opaque»shagreened, medially polished ; clypeus polished and
sparsely punctate, in profile strongly convex ; malar space hardly two-
thirds basal width of mandible; antennae 32-jointed, only very
slightly thickened, and flattened beyond middle. Thorax rather stout,
polished and sparsely punctate except as noted below ; humeral mar-
gin of pronotum smooth, carinately angled anteriorly, scrobe striate
below ; lobes of mesoscutum rather flat, prescutum transversely striate
along notaulices; scutellum weakly convex; mesopleuron anteriorly
and metapleuron in lower posterior angle obliquely striate; propo-
deum with basal area defined, basal carina arching forward medially,
apical carina distinct between the very stout, compressed apophyses,
middle of propodeum both before and behind apical carina obsoletely
transversely rugose, and with a few coarse rugae radiating from the
apical margin ; legs rather stout, hind femur hardly five times as long
as thick, less than two-thirds as long as tibia and barely reaching -
apex of abdomen; areolet rather broad, its lower side distinctly
angled by the strongly antefurcal recurrent. Abdomen polished,
stout, petiole distinctly depressed, postpetiole broader than long, dor-
sal carinae absent ; second tergite shorter than first and barely a half
longer than broad at base ; ovipositor sheath about as long as abdomen
beyond first tergite, ovipositor rather slender, of uniform depth, and
weakly subsagittate at apex.

Black and yellow (see description of color pattern, p. 141) ; orbital
rings of unusually uniform width behind eyes; annulus embracing
flagellar joints (5) 6-10 (12) ; yellow of anterior margin of pronotum
reduced to two small spots near middle ; two small streaks on disk of
mesoscutum ; scutellum with an elongate black mark in basal middle ;
mesopleuron entirely yellow; yellow spots in basal lateral areas of
propodeum and the apical bands extending forward nearly to basal
carina; petiole yellow both above and below; broad subapical bands
and lateral margins of tergites 2 and 3 yellow, the color extending
inward at base of second ; remaining tergites entirely and venter yel-
low. Front and middle legs yellow, with femora and middle tro-
chanters piceous posteriorly ; middle coxa piceous on each side above ;
hind coxa similarly marked but more reddish beneath; trochanter
stained with piceous; femur reddish on outer side, piceous within;
tibia and tarsus yellow. Wings hyaline, venation black.

Type locality. — Trinidad Kio, Panama.

Type. — One specimen taken by August Busck, March 16, 1912. It
is in the U. S. National Museum (No. 57061).

ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 151
6. CRYPTANURA PICEOTHORAX, new species

Female. — Longtli 1-2 iniu.. rtntcnniie 8.5 inin., ovipositor shealli
3.5 mm.

Temples narrow, flat, and sharply receding; occipital carina prom-
inent, especially below, where it joins the hypostomal carina not far
from the base of the mandible; vertex nearly straight across except
for a groove at each side of ocelli; ocelli at level of superior tangent
of eyes; frons moderately concave, slightly tumid laterally, horns
prominent on a common base; eyes moderately large and somewhat
bulging, weakly convergent below; face with a rather deep striate
impression on each side of middle, the middle roundly convex and
with sparse, coarse punctures; clypeus sculptured like middle of face,
strongly convex, in profile subnasute; malar space flat, nearly as long
as basal width of mandible; antennae stout, slightly thickened and
flattened near apex, 29-jointed. Thorax stout, generally coarsely
punctate to striatorugose; pronotum with humeral margin smooth^
conically produced anteriorly, scrobes coarsely striate; mesoscutum
coar.-^ely punctate, prescutum somewhat rugose, lobes rather flat;
scutellum flat and coarsely punctate above, apex rather abrupt and
impuMctate; frena of both scutellum and postscutellum coarsely
foveolate; pleura and propodeum coarsely striatorugose; sternum
coarsely punctate; basal areas partly smooth; basal median area
l)artly defined, basal carina nearly straight; legs rather stout, hind
femur little more than two-thirds as long as tibia and not extending
to apex of abdomen; hind coxa coarsely punctate above; areolet short.
Abdomen broad, polished; petiole strongly depressed, post petiole
much broader than long, with a weak elevation on each side of middle
between spiracles, but without dorsal carinae; second tergite a little
shoiter than first and only about a third longer than broad at base;
sheath sliorter than abdomen beyond first tergite; ovipositor rather
thick, in profile rather slender, a little deeper near apex, the apex not
at all sagittate.

Head and thorax black to rcdtlisli piceous, both with yellow mark-
ings (see description of color pattern, p. MI); amiuhis emliiacing
flagellar joints (5) 6-9 (10) ; single spot on disk of mesoscul urn ; only
apex (tf scutellum yellow; most of nies(»pleuron and sternum and
nietaj)leiiron except suture sepai'al ing its two divisions yellow ; pro[)o-
deMJ marking extending broadly forward to basiil caiina, but con-
stricted before apophyses. Legs ferruginous, coxae and trochanters
partly yellow. Wings hyaline, veins blacU, stigma brown. Abdomen
lilack with white markings; petiole, except vent rally, white ; three spots
t lan.-vei-sely airange(] at base of second teigite and a bioad subapical
band also white; the extreme a[)ex i)lack.

152 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se

Type locality. — Baragua, Camagiiey, Cuba.

Types. — Two females, the holotype taken at light June 4, 1932
(Christenson), the paratype at Sole, Cuba, March 2, 1925 (George
Salt). Both are in the U. S. National Museum (No. 57063).

The paratype is smaller, with the sculpture less heavy and the dark
color of the thorax more reddish.

7. CRYPTANURA PLANrSCUTELLATA, new species

Very similar in form, structure, and color pattern to piceothorax
but at once distinguishable by the color of its thorax, which is not at
all red, and of the legs, which are black and yellow. Differs further
from piceothorax as follows :

Female. — ^Length 12 mm,, antennae 8.5 mm., ovipositor sheath
5.5 mm.

Sculpture of thorax, especially metapleuron and propodeum, in-
cluding the basal areas, more reticulate-rugose than striatorugose ;
scutellum less abruptly sloping at apex ; postpetiole and tergites 2 and
Z coarsely, sparsely punctate, especially laterally.

Antennal annulus longer, extending to flagellar joint 13; pale mark-
ings of abdomen yellow rather than white; legs to apices of femora
black and yellow, tibiae and tarsi pale stramineous ; front and middle
coxae yellow, black behind, hind coxa yellow with base below and
streaks on upper and inner sides black; trochanter yellow, basal joint
above and apical joint at apex more or less black, hind trochanter with
basal joint largely black; front and middle femora yellow, black be-
hind; hind femur yellow on outer and inner sides, black above and
below.

Type locality. — Puerto Castilla, Honduras.

Type.— One female dated March 26, 1924. It is in the U. S. National
Museum (No. 57062).

This may be the same as compacta (Cresson).

8. CRYPTANURA POLITIGASTER, new species

This species is similar in structure and in color pattern of head
and thorax to the preceding two species but differs from both in its
red abdomen and in other details.

Female. — Length 11 mm., antennae 9 mm., ovipositor sheath 3 mm.

Temples rather long, slightly but distinctly concave, sharply reced-
ing, occipital carina high, especially at lower extremity, where it
passes over a prominence in the posterior margin of the cheek, and
joins the hypostomal carina rather close to the base of the mandible;
cheek transversely striate along carina, vertex flat with a shallow
groove next to ocelli; frons deeply concave, tumid next to eyes, ob-
liquely striate above, horns small but elevated on a high common base ;

ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 153

eyes large, somewhat buliriiig, weakly convergent below; face nearly
smooth laterally and medially with a rugose impression on each side
of middle, the rugae extending across middle at top; clypeus smooth
with large scattered punctures, nasute in profile; malar space flat and
fully as long as basal witlth of mandible; antenna rather stout, weakly
thickened and flattened near apex, 30-jointed. Thorax stout; humeral
margin of pronotum smooth, conically prominent anteriorly, scrobe
striate; mesoscutum coarsely punctate, lobes rather flat, present um
very low anteriorly; scutellum sloping from base to apex, polished,
with a few large punctures; frena coarsely foveolate; mesopleuron
longitudinally striate below tubercle, obliquely and indistinctly so
below, elsewhere polished and sparsely punctate, prepectus rugu-
lose-punctate, sternum polislied and sparsely punctate; metapleunm
coarsely rugose above, polislied and punctate below, as is also its upper
division; propodeum largely rugose, more or less transversely so
behind apophyses, basal median area defined, basal carina curved
medially; legs rather slender but not long, hind femur barely two-
thirds as long as tibia and not reaching apex of abdomen; areolet
short. Abdomen broad, polished, with a few large punctures at sides
of postpetiole and second tergite; first segment stout, petiole flattened
above but not distinctly depressed, postpetiole much broader than
long, without trace of dorsal carinae; second tergite nearly as long
as first, but barely a third longer than broad at base ; ovipositor sheath
distinctly shorter than abdomen beyond first segment; ovipositor of
uniform depth; apex weakly subsagittate.

Head and thorax black with yellow markings (see description of
color pattern, p. 141) ; annulus embracing flagellar joints (5) 6-11
(13); anterior margin of pi-onotum medially and humeral margins
before tegulae black; spot in center of mesoscutum; scutellum yellow
only at apex; mesopleural band divided into a large spot anteriorly and
a smaller one posteriorly; a large spot on each side of sternum; both
npj)er and lower divisions of metapleuron largely yellow; propodeal
markings extending broadly forward to carina but constricted before
apophyses. Front and middle legs yellow, with their coxae, tro-
chanters, and femora black behind; hind coxa black and yellow, tro-
chanter and femur ferruginous, tibia and tarsus yellow. Wings
hyaline, venation black. Abdomen fei'iiiginous; petiole and apex of
fii-st tergite yellow, postjM'fioh! ferruginous and black; stcrnites pale
ferruginous margined wilh whitish.

Type locality. — Kurrenabaque, Itif) Beni, I'olivia.

Type. — One female, taken in October by W. M. Mami on the
Mul ford P.iological E.\i)loration of 191^-22. It is in the U. S. National
Museum (No. 570C5).

154 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9»

9. CRYPTANURA CONICA, new species

Similar to politigaster but distinguishable at once by its much more
strongly flexed occipital carina and by its entirely red femora and
hind coxae.

Female. — Length 12 mm., antennae (tips broken off), ovipositor
sheath 3.5 mm.

Temples rather long, slightly concave, and sharply receding; occi-
pital carina high, especially below, where it passes over a strong angu-
lation in the posterior margin of the cheek; vertex concave, ocelli dis-
tinctly below superior tangent of eyes; frons moderately concave,
with rugae radiating from ocelli, horns small but elevated on a broad
common base ; eyes bulging, very weakly convergent below ; face with
a shallow impression on each side of middle, medially coarsely rugose,
laterally smooth; clypeus with scattered coarse punctures, in profile
subnasute; malar space fully as long as basal width of mandible, flat;
antennae distinctly thickened and flattened beyond middle. Thorax
stout, largely coarsely sculptured; pronotum with humeral margin
transversely striate, conically produced anteriorly, scrobe coarsely
striate; mesoscutum coarsely but not densely punctate, lobes, espe-
cially prescutum anteriorly, flattened ; mesopleuron striate, the sculp-
ture changing to punctation toward lower posterior angle, sternum
polished and sparsely punctate; scutellum roundly sloping from base
to apex, polished, with sparse punctures, frena foveolate; metapleu-
ron obliquely punctatostriate, its upper division smooth and sparsely
punctate; propodeum polished before carina, coarsely rugose behind,
the rugosity becoming transverse on apical slope, basal median area
defined, basal carina curved forward medially; legs rather slender
but not very long, hind femur barely two-thirds as long as tibia and
not reaching apex of abdomen, hind coxa polished, with sparse punc-
tures; areolet narrow. Abdomen rather narrow, polished; petiole
flat above but not depressed, postpetiole much broader than long, with
no trace of dorsal carinae; second tergite about as long as first and
about a half longer than wide at base ; sheath much shorter than abdo-
men beyond first tergite; ovipositor slender, of uniform depth, and
subsagittate at apex.

Head and thorax black with yellow markings (see description of
color pattern, p. 141) ; orbital ring broadly interrupted on temples;
annulus on flagellar joints (5)6-10(13) ; anterior margins of prono-
tum entirely and humeral margin posteriorly black; mesoscutum im-
maculate ; apex of tegulae piceous ; mesosternum immaculate ; yellow
marking of lower division of metapleuron reduced to a streak in the
dorsoposterior half; propodeal markings extending from apex to
basal carina and narrowing and divergent from apophyses to carina.
Legs ferruginous, tibiae and tarsi and front and middle coxae ante-

ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 155

riorly yellow. AVings hyaline, venation black. Abdomen ferrugi-
nous, petiole and apex of postpetiole stramineous.

Type localitj/. — Kamakusa, British Guiana.

Type. — One female taken by H. Lang. It is in the U. S. National
Museum (No. 57066).

10. CKYPTANUKA RUFICEPS, new spcciea

This and the next following species differ from all others known to
me in lacking any trace of angulation of the tumid humeral margin
of the pronotum; and the present species is unique in the genus for
its red head and is very unusual in its lack of a pale annulus on the
antenna.

Female. — Length 13 mm., antennae 10.5' mm., ovipositor sheath
4.5 mm.

Temples short, distinctly convex, sharply sloping, occipital carina
rather high, especially at lower end, hypostoraal carina very high;
vertex straight across, ocelli elevated above general level; frons not
deeply excavated, horns very small, on a broad, low, common base;
eyes not bulging; parallel within; face subpolished, with coarse punc-
tures, especially in sliallow impressions on each side of middle, median
portion in profile nearly as strongly convex below as is clypeus,
which is not so strongly convex as usual and is sculptured like the
face: malar space a little shorter than basal width of mandible; cheeks
in front view slightly convex, in side view fully twice as broad as
malar space; antenna slightly thickened and flattened beyond mid-
dle. Thorax robust, sculpture mostly coarse rugosity; pronotum with
scrobe coarsely striate, epomia strong, but terminating below the
smooth humeral margin, which is less strongly tumid anteriorl}' than
posteriorly; mesoscutum with lobes very weakly convex and notaulices
very shallow, polished and sparsely, coarsely punctate; scutellum
strongly convex but not prominently so, polished, with a few punc-
tures, frena foveolate; mesopleuron largely coarsely rugosostriate;
sternum coarsely punctate; metapleuron and propodeum, except its
punctate basal areas, transversely striate, basal median area not de-
fined, basal carina curved forward me^lially; apophyses unusually
short and obtu.se; legs rather slender, hind femur three-fourths as
long as tibia and reaching apex of abdomen, hind coxa polished and
punctate; areolet broad. Abdomen stout, i)olislied; petiole distinctly
flepressed, ventrolateral carina strong, with a foveolate groove above
it, postpetiole much broader than long, with low rounded elevations
repre.sentmg the dorsal carinae; second tergite about a fourth longer
than broad at base; ovipositor sheath not quite so long as abdomen
beyond first tergite, ovipositor rather stout, subsagittate at apex.

824514— 4!5 3

156 PROCEEDINGS OF THE NATIONAL MUSEUM vol, 66

Head and thorax rufous with yellow markings (see description of
color pattern, p. 141) ; orbital ring not abruptly narrowed on temples;
facial impressions, clypeal suture, and margin of clypeus ferruginous ;
scape rufous, flagellum entirely black; mesoscutum with a single
median spot; scutellum with a rufous spot in basal middle; meso-
stemum and both upper and lower divisions of metapleuron largely
rufous; propodeal markings each with a narrow tongue-shaped ex-
tension before the apophyses and reaching nearly to basal carina.
Antenna black, scape rufous, without white annulus; small areas in
rufous coloring of head and thorax more or less piceous; legs entirely
ferruginous ; wings yellowish hyaline, venation black. Abdomen black
and white ; three white spots at base of second tergite, the middle one
large and broadly transverse, the lateral ones small and longitudinal ;
venter mostly white ; sheath black.

Type locality. — Pico Turquino, Cuba.

Types. — A holotype and four paratypes, all females, collected by
S. C. Bruner and C. H. Ballou, July 10-20, 1922, at elevations ranging
from 2,900 to 6,000 feet. They are in the U. S. National Museum (No.
57067).

11. CRYPTANURA SEPTENTRIONALIS, new species

Related to unifjceps in the lack of the angulation of the humeral
margins of the pronotum and in the color pattern of the abdomen, but
distinct from that species in the black and white head and thorax
and in many details of structure and color.

Fem-ale. — ^Length 15 mm., antennae (broken), ovipositor sheath
4,5 mm.

Differs from ruflceps principally as follows : Frons obliquely striate
above, horns not on a common base ; postvertex densely, temples, cheeks,
and sides of frons more sparsely, punctate; face rugose except later-
ally, less strongly convex medially; malar space much shorter than
basal width of mandible; occipital and hypostomal carinae weaker;
sculpture of thorax largely coarse punctation, densest and coarsest on
mesoscutum and metapleuron, humeral margin of pronotum coarsely
punctate ; propodeum posteriorly irregularly transversely rugose, scu-
tellum polished ; notaulices deep ; apophyses longer ; abdomen beyond
first tergite opaque shagreened, second and third tergites and post-
petiole at sides punctate ; ventrolateral carinae of petiole subobsolete.

Differs from ruflceps in color as follows: Head and thorax black
and white ; face entirely white ; antennal annulus embracing flagellar
joints (5) 6-11 (12) and underside of scape white: propodeal white
marks narrowly confluent across median line anteriorly ; coxae white,
the front and middle ones more or less black behind, the hind coxa
black on outer and inner sides toward base and with a median black
stripe above, trochanters largely white, legs otherwise ferruginous,
with tarsi paler.

ICHNEUMON-FLIES OF GENUS CRYPTANURA CUSHMAN 157

Type locality. — Cleveland, Ohio.

Types. — One female (the holotype) captured September 19, 1932, by
Frank D. DeGant; one female paratype, St. Louis, Mo., June 29, 1938;
and one female paratype received from H. A. Scullen, Corvallis, Oreg.,
but probably eastern. A\\ are m the U. S. National Museum (No.
57068).

This is the second species of this tropical genus to be found in the
United States.

12. CRYPTANURA APOPHYSIS, new specieii

Distinct from all tlie other species known to me in its extraordinarily
large propodeal apophyses. It is also almost unique in its entirely
yellow front and middle legs and black hind femur.

Female. — Length 15 mm., antennae (missing), ovipositor sheath
5.5 mm.

Temples flat to slightly concave, long, sharply receding; occipital
carina very high, but becoming abruptly lower just before its junction
with the very high hypostomal carina; vertex concave, top of ocelli
below superior tangent of eyes; frons with short rugae radiating
from ocelli, deeply concave below, horns short, stout, not on a com-
mon base; eyes very large, bulging, very weakly convergent below;
face medially elevated, with an impression on each side of middle,
opaque .shagreened, with sparse punctures, obliquely rugose below
antennae; clypeus subnasute, subpolished, with a few coarse punc-
tures; malar space nearly as long as basal width of mandible, straight
in front view. Thorax distinctly compressed ; pronotum with humeral
margin transversely striate, carinately angulated anteriorly, scrobe
striate; mesoscutum subpolished and rather densely and coarsely
punctate, lobes low, notaulices deep and narrow; scutellum strongly
I on vex, polished, very sparsely punctate; mesopleuron subpolished,
-par.sely punctate below, scrobe polished below and striate above;
mesosternum more densely punctate; upper division of metapleuron
coarsely punctate, lower division coarsely obliquely rugosostriate ;
propfMleum rugose laterally and between carina and apophyses,
otherwise largely polished, apopl»y.ses very large, long, conical, nearly
twice as long as their distance from basal carina; legs very slender,
hind femur fully eight times as long as deep and reaching beyond apex
of abdomen; areolet elongate. Abdomen s](!nder, linely shagreened
and suliopaque beyond first tergite; petiole ahnost exactly .square in
cioss-section. {)ost[)etiole nearly as long as broad; second tergite fully
two-thirds longer than l)road at base; sheath as long as abdomen
beyond first tergite; ovipositor rather stout, distinctly subsagittate at
apex.

Black and yellow (see description of color pattern, p. 141) ; orbital
ring nearly interrupted in upper temple and in malar space; two

158 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

narrow lilies on disk of mesoscutum ; a large spot anteriorly on meso-
pleuron and a smaller one before middle coxa ; yellow of lower division
of metapleuron reduced to a large spot in the middle of its upper
margin; propodeal spots extending only a short distance anterior to
apophyses ; front and middle legs except their coxae at base behind,
a large spot on upper side of hind coxa, hind tibia except narrowly at
base and broadly at apex, and probably hind tarsi (though these are
missing) yellow; petiole entirely black; wings yellowish hyaline,
venation blackish.

Male. — Essentially like female, but more highly polished ; the pro-
podeum with striae radiating from bases of apophyses, which are
somewhat more slender than in female; postpetiole fully as long as
broad and second tergite fully twice as long as broad at their junction ;
legs even more slender.

The antennae, though not entire, exhibit a yellow annulus beginning
on the eleventh flagellar joint, and the hind tarsus is entirely yellow.

Type locality. — Colombia.

Allotype locality. — Huascaran, Peru.

Types. — One of each sex, the type female, having no further data ;
the allotype taken September 21, 1911, by C. H. T. Townsend. Both
are in the U. S. National Museum (No. 57069) .

13. CRYPTANURA MEXICANA (Cresson), new combination

Mesostenus mexicanus Cresson, Proc. Acad. Nat. Sci. Philadelphia, 1873, p. 157,
female, male (part).

Cresson's type series was not all of the same species, for a male
eotype in the National Collection is referable to orizdbensis (Cam-
eron). A female eotype agrees with Cresson's description, and ap-
parently is the true meooicana.

Three additional specimens are before me as follows : Argas,
Panama, April 28, 1911 (A. Busck) ; Costa Rica and Santa Marta,
Colombia, 1852 (Fontanier), the last belonging to the Paris Museum.

Female. — ^Length 14 mm., antennae 14 mm., sheath 5.5 mm.

Temples flat or slightly concave, occipital carina high, slightly
sinuate below; vertex weakly concave, ocelli a little below superior
tangent of eyes ; postocellar line much shorter than ocellocular line ;
frons moderately concave, with a few rugae radiating fi'om the
ocelli, horns short and broad, not on a common base ; eyes large, bulg-
ing, weakly convergent below; face convexly elevated medially, pol-
ished and sparsely punctate, with a transversely striate impres-
sion on each side of middle; clypeus strongly convex but not subnasute,
polished and sparsely punctate; malar space nearly as long as basal
width of mandible, in front view straight; antenna about 38- jointed,
strongly thickened and flattened below beyond middle. Thorax
stout; humeral margin of pronotum transversely striate, carinately

ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 159

aiighnl anteriorly, scrobe striate; niesoscutuiii polished, sparsely
t^oarseiy puiu-late. notaulices deep and narrow; scutellum narrow,
convex, polished, with a few punctnres; mesopleuron obliquely striate
alx)ve, the striation chan<rin<T £rradually to punctation below, sternum
punctate; upper division of metapleuron punctate, lower division
coarsely, obliquely rugosostriate ; propodeuni with basal median area
obsoletely defined, basal carina curving forward medially, apophyses
long and slender, basal areas punctate and posteriorly more or less
rugose, space between carina and apophyses with oblique striae con-
verging toward middle, posterior face more coarsely, transversely
striate; legs lt)ng and slender, hind femur reaching apex of abdomen,
hind coxa sparsdly, coarsely punctate; areolet elongate. Abdomen
beyond fii-st tert^ite finely shagreened, subopaque; petiole not de-
pressed, post petiole broader than long, with median and lateral shal-
low impressions; sheath a little longer than abdomen beyond first
tergite ; second tergite about a half longer than broad at base; oviposi-
tor subsagittate at apex.

Black with yellow markings (see description of color pattern,
p. 141) ; orbital ring interrupted in upper temples; annulus embracing
flagellar joints (5) 0-10 (12) ; two elongate marks on disk of meso-
scutum ; scutellum yellow only at apex, mesosternum largely yellow :
lower division of metapleuron very largely yellow; jiropodeal mark-
ings extending forward nearly to carina; petiole yellow above, black
below. Legs yellow ; front and middle coxae and femora behind, base
of hind coxa and a broad stripe above, and apical joint of hind tro-
chanter and the fennir above black. Wings hyaline, narrowly, weakly
infumate apically, venation blackish,

I have not seen a male of this species, but according to Cre^ison il
differs from the female in being more slender, with antennae and
legs longer, the knees and the apex of the hind tibia black.

U. CRYPTANLRA ORIZABENSIS (Cameron), new combination

SfiHoxtenua mcTinimiH Ckkhson, Proc. Acad. Nat. Sfi. Pliiladelpbia, 1873, p. 1157,

mule (part).
I'olyafnun orizabrnxiH Camkkun, Hlologla Centrall-Aiuericaiia. Hyuienoptera, vol.

1, 188G, p. 240, female.

As stated above, the National Museum male colype of iiwxicana
(Cre.s.son) belongs to this species. A female, also from Mexico, from
the C. F. Baker collection is also before me.

The female differs from that of tncx-i/iana in structure principally
as follows: Occipital carina not sinuate below; postocellar line nearly
as long as ocellocular line; frontal horns longer and more acute;
sculpture at middle of proi)odeum irregularly transverse rugosity;
ovipositor and sheath distinctly .shorter; areolet shorter, with recur-
rent interstitial.

160 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Differs from mexicana in color as follows : Mesosternum yellow only
along sternaulices ; propodeal markings extending only a little before
apophyses; hind coxa black above with a large oval yellow spot;
basal joint of hind trochanter largely black, apical joint yellow, hind
femur black above but with a narrow median yellow line, hind tibia
somewhat blackish or brownish at base and apex ; petiole black above,
yellow below.

The male is very much like the female but has the frontal horns
longer and more acute and the hind tibia more broadly black at apex.

15. CRYPTANURA BILINEATA, new species

Very similar to the two preceding species, but distinct from both
in the form of the propodeal markings, which extend forward to the
carina but are abruptly narrowed before the apophyses; in the less
distinctly thickened and flattened antennae; in having the frontal
horns set rather high on a common base; in the broader and more
weakly convex scutellum; and in the short, obtuse apophyses. It is
like Tnexicana in general in the color of the legs but has the hind coxa
colored as in orisabensis; the petiole is, as in mexicana, yellow above
and black below; and the mesosternum, as in oriBobensis, is yellow
only along the sternaulices. In the long postocellar line and in the
sculpture of the propodeum it is more like ori^abensis and in the
length of the ovipositor and in venation of the wings it agrees with
mexicana.

Type locality. — Cuernavaca, Morelos, Mexico.

Type.—On& female in the U. S. National Museum (No. 57064),
taken by E. G. Smyth.

16. CRYPTANURA RUFA, new species

Distinct from all other species known to me and apparently from
all other described species in the arrangement of color, black and
yellow head, red and yellow thorax (not black dorsally), and entirely
red abdomen.

Female. — ^Length 14 mm., antennae 12 nmi., ovipositor sheath
5 mm.

Temples weakly convex, occipital carina not high, with a shallow
notch just before its junction with the hypostomal carina ; vertex flat,
ocelli as high as superior tangent of eyes ; f rons rather weakly concave,
with rugae radiating from ocelli, horns short, stout, rather widely
separated ; eyes weakly convergent below ; face polished and sparsely
punctate, weakly convex medially, with a shallow, transversely striate
impression on each side of middle; clypeus polished and sparsely
punctate, subnasube in profile; malar space three-fourths as long as
basal width of mandible ; cheeks in front view weakly convex. Thorax
robust ; humeral margin of pronotum transversely striate, carinately
angled anteriorly, scrobe sparsely striate ; mesoscutum rather densely

ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 161

punctate, notaulices deep and narrow, lobes rather flat; scutellum
convex, polished, and obsoletcly punctate; mesopleuron obliquely stri-
ate, this sculpture changing below and posteriorly to rather sparse
punctation. sternum more densely pun'-tate; upper division of meta-
pleuron polished and sparsely punctate, lower division coarsely,
obliquely rugosostriate ; propodeum polished and punctate before and
transversely rugosostritite behind carina, posterior fare transversely
striate, apophyses moderately long and slender; legs rather stout, hind
femur hardly reaching apex of abdomen, hind coxae polished and
sparsely punctate. Abdomen ■^ubpolished, inconspicuously shagreened
beyond first tergite; petiole slightly depressed, postpetiolc much
broader than long, with faint rounded elevations between the spiracles ;
second tergite about a third longer than broad at base; sheath shorter
than abdomen beyond first tergite, ovipositor rather stout, subsagit-
tate at apex.

Head black (see description of color pattern, p. 141) with orbital
lings interrupted in upper temple and in malar space; facial impres-
sions and outline of clypeus piceous. Thorax ferruginous, slightly
stained with piceous in postscutellar frena and in speculum and with
yellow markings; single spot on disk of mesoscutum; scutellum yellow
only at apex: upper divisioii of metapleuron and dorsal half of its
h'wer division yellow; propodeal markings including the apophyses
and two small spots just in front of and mesad of these. Front and
middle legs yellow, with front coxa, trochanter, and femur piceous
behind, middle coxa ferruginous with yellow spot in front; hind leg
ferruginous witli tibia and tarsus yellow. Wings hyaline, venation
blacki.sh. Abdomen entirely red.

Type locality. — Estero de Sao Paulo, Brazil.

Typea. — A female holotype and two female paratypes tnken in 1910
by E. R. Wagner. Tlie holotype and one paratjqoe are in the Paris
Museum, and the second paratype is in the U. S. National Museum
(Xo. 57081). There is also a female with l)roke,n antennae in the Paris
Mu.seum, taken in 1010 at Cliaeo de Santiago del Estero, Argentina.

17. f RYPTANURA SPINARIA (Brullc)

Mr.nnHtenuft Mpinariuif BRt:T,l.f;, Hislnjrf njitun-llr' dos ins<'<-fos, Hyiin'iioplern, vol.
4. p. 227, 184f5, feraalo.

^fC80fltfmuH alhnpirtus rBERBON hiul Smith), I'ror. Ent. Sor. rhiladelphin. vnl. 3.
p. 312. 1804. male.

}friiO!iU-nu.<< drinii nrrnsia Dali.a Toiire. CntnloRus liyiiiciiojifornninT, i>. HIO. lfK)1~
02.

I'Dlfinriix spiti'irifix SriiMlKOKK M' iii. (IfiK-rn iiist'rtnruiii. f)iH<'. 7"t. p. C,S. IJtilS.

\frsoKtrnuH spituiriiiH ViESKCK, in Sinitli, Inserts nf Now Jorsoy. p rC^O, 1f>1(t.

^fc808tenid^a (Vnlynrnun) spinaria Vikkdi;k, The HyiiHMioptcra <>r wnsp-liko In-
sects of Connef.tlcut, p. 320, 1917.

Pnlyncnun Kpinnriim fxTSiiMAN, Joiirn. ^^■nsllinKtn^ Acnrl. Sri., vol. 15. p. 301. 10"Jr» :
Pror. U. S. Nat. Mus., vol. 74. nrt. 10. p. .3S. fl«s. U. 3fc, W, 1020.

Crupl">iiirn ^pinnrin Tdwnks, M<>m Aiikt. Knf. Soc , No. 11, jit. I. j). L'MS. 1011.

162 PROCEEDINGS OF THE NATIONAL MUSEUM vol,. M

I have described this North American species too recently and too
fully to make further discussion necessary here. It may well be men-
tioned, however, that the mesoscutum frequently has, in addition to
the single median yellow spot, traces of tlie yellow lines along the
inner margins of the lateral lobes and sometimes those along lateral
margins next to the tegulae.

18. CRYPTANURA LINEATIFEMUR. new species

Fennale. — Length 14 mm., antennae 11 mm., sheath 5 mn\.

Except that it has a single median spot on the mesoscutum instead
of two yellow lines and that the propodeal markings extend narrowly
forward to the basal carina, this species is very similar in color and in
large part in structure also to the three species mexicana^ orizabensis,
and hilineata. The hind legs are colored exactly like those of oriza-
iensis, as is the mesosternum, while the abdomen is colored like that
of mexicana. The temples are not at all concave, the cheeks are more
convex, tlie vertex is flat, the occipital carina is lower, the antennae
are shorter, the petiole is thicker, with the lateral carinae, both dorsal
and ventral, more distinct, and the ovipositor longer ; otherwise very
similar to oiizabensis.

Type locality. — La Caja, near San Jose, Costa E-ica.

Types. — Two females, the holotype and .i paratype, taken at the
above locality by M. Valeria on July 1, 1931, at an altitude of 900
meters, and one female paratype captured by Schaus and Barnes at
Cayuga, Guatemala. All are in the U. S. National Museum (No.
55071).

The Costa Rican paratype is smaller and the Guatemalan paratype
larger than the type, but otherwise they are similar.

Mesostenus veraepacis Cameron, which seems to be a Ci^ptarmra,
appears to be very similar to this species but differs in that the hind
femur is entirely black above without the yellow median line.

19. CRYPTANURA COXATA, new species

Female. — Length 16 mm., antennae 12 mm., ovipositor sheath
5.5 nam.

Temples flat, short, and very strongly receding, occipital carina
moderately high, somewhat higher and slightly sinuate below ; vertex
nearly straight across ; f rons rather deeply concave above, with striae
radiating from ocelli, horns short and broad, close together but not on
a common base ; eyes slightly bulging, weakly convergent below ; face
polished and sparsely punctate, somewhat rugose below antennae,
impressed on each side of middle; clypeus strongly convex, polished,
sparsely punctate ; malar space about two-thirds basal width of mandi-
ble; cheeks in front view weakly convex; antenna 32-jointed, dis-
tinctly broadened and flattened beyond middle. Thorax stout; hu-
meral margin of pronotum transversely striate, carinate anteriorly.

ICHNEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 163

scrobe striate; mesoscutuni polished aiul sparsely punctate, lobes some-
what flattened; scutellum convex, polished, very sparsely punctate;
upper division of nietapleuron polished and sparsely punctate, lower
division coarsely, obliquely ru^osostriate; pro[)odeuni transversely
striate behind apophyses, before which it is irregularly rugose, basad
of carina polished medially and rugosopunctate laterally, apophyses
rather long and slender; legs rather stout, hind femur hardly reach-
ing apex of abdomen. Abdomen stout, subopaque shagreened beyond
lii-st tergite; petiole depressed, postpetiole much broader than long
wiih two rather prominent elevations between spiracles; second ter-
gite hardly a third longer than broad at base; sheath much shorter
than abdomen beyond first tergite, ovipositor rather stout, subsagittate
at i\[K'x.

Head and thorax black with yellow markings (see description of
color i)attern, p. 141) ; orbital ring interrupted in upper temple; annu-
his em!)racing ilagellar joints (5)6-10(12) ; a single spot in center of
]iifsoscutum; large spot on each side of sternum; propodeal markings
abrui)tly narrowed but not constricted before apophyses, and curving
tr)ward each other just behind the carina. Front and middle legs
yellow, the coxae, trochanters, and femora black behind; hind coxa
yellow below and in middle above, black above on outer and inner sides,
trochanter black basally, reddish apically, femur ferruginous, tibia
and tarsus yellow ; wings yellownsh hyaline, venation blackish. Abdo-
men ferruginous, first segment black with petiole dorsally and apical
margin yellow.

J/,/A. — Much smaller than female, with apical carina complete and
.strongly arched forward medially, the apophyses reduced to highei
elevations in the carina, rugosity of thorax, except in potiolar area,
largely replaced by punclation; abdomen much smaller and narrower,
legs relatively longer and more slender; clypeus entirely yellow
except apical margin; scape beneath; annulus in flagellar joints
(10)11-17(18); propodeal markings confluent anteriorly; all abdo-
minal segments beyond first more or less blaikish basally.

7'j/pe local if y. — Tiinidad Kfo, Panama.

TyjH'.s. — 'I'wo females and one male, the liolot ype an<l all<)lyi)e, taken
March 27 and 20, 1912, by August Busck, ami a i^arntype female by
\V. M. Mann at Ixianias, Bolivia, in Deeembei- 1!>21, on the Mulford
liiological Ivxploration. All are in the U. S. National Museinn (No.
.07070).

20. CRYPTANURA BOLIVIENSIS. new upcritu

Very closely related to coxdfa, frcjm which it can at once be distin-
guished by the entirely red hind coxa and trochanter and first teigile.

Fnnalc. — Length ir» mm., antenna 11.;") mm., ovipositor sheath T) mm.

Differs further fi'om roxdia as follow^: 'I'eniples longei- and less
-harply receding; humeral margins of pronotum subtuberculate

164 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

anteriorly; lateral lobes of mesoscutum nearly impunctate; scutellum
impunctate; thorax laterally much less strongly striate, mesopleuron
mostly punctate rather than striate; abdomen narrower; petiole not
depressed, postpetiole not especially broad, though somewhat broader
than long, second tergite more than a half longer than broad at base.

Antennal annulus embracing flagellar joints (4) 5-11 (14) ; pro-
podeal markings strongly constricted before apophyses and not curving
mesad anteriorly ; middle coxa and trochanter ferruginous behind.

Type locality. — Rosario Lake, Rogagua, Bolivia.

Type. — One female taken in November 1921 by M. R. Lopez, on the
Mulford Biological Exploration. It is in the U. S. National Museum
(No. 57072).

21. CRYPTANURA ISTHMUS, new species

Female. — ^Length 15 mm., antenna 12 mm., ovipositor sheath 5 mm.

Structurally almost identical with coxata., but diflFering in color as
follows : Mesosternum yellow only along sternaulices ; propodeal mark-
ings constricted basad of apophyses and not curving toward each other
anteriorly; hind coxa ferruginous, with an indistinct yellow spot at
base above; petiole stramineous, postpetiole piceous with apical mar-
gin yellow, second tergite obsoletely yellow just before apex.

Type locality. — Trinidad Rio, Panama.

Types. — Two females, the holotype and a paratype, taken by August
Busck at the type locality on March 16 to 19, 1912; and one female
paratype taken by C. T. Greene at Ajicon, Canal Zone, May 17, 1926.
They are in the U. S. National Museum (No. 57073).

22. CRYPTANURA EXCALIBUR, new species

This and the next following two species and probably Mesostenus
platyurus Brulle form a group distinct from the rest of the genus in
the long, stout ovipositor which in side view is almost clavate, becoming
gradually much deeper toward the apex. Brulle's species is not known
to me, but it may be distinguished from the present species by its red
propodeum.

Female. — Length 16 mm., antennae 13 mm., ovipositor sheath 9 mm.

Temples very weakly convex, rather short, occipital carina high,
especially just before joining hypostomal carina; vertex straight
across; frons rather shallowly concave, with rugae radiating from
ocelli, horns short, thick, separate; eyes very weakly convergent
below; face subpolished, sparsely punctatorugose above, somewhat
elevated medially, with a shallow impression on each side of eleva-
tion; clypeus strongly convex, polished and sparsely punctate; malar
space two-thirds basal width of mandible ; cheeks in front view con-
vex; antenna 36-jointed, very slightly thicker and flattened toward
apex. Thorax stout; humeral margin of pronotum transversely stri-
ate, carinately angled anteriorly, scrobe striate; mesoscutum subpol-

ICHXEUMON-FLIES OF GENUS CRYPTANURA — CUSHMAN 165

ished and coarsely punctate, lobes weakly convex; scutellum stron<j;Iy
convex, polished, nearly inipunctate; niesopleuron obliijuely striate
above, rather densely punctate below, sternum similarly punctate;
upper division of metapleuron polished and sparsely punctate, lower
division coarsely, obliquely rugosostriate; propodeum striate behind,
the striations becoming somewhat confused above, basad of carina lat-
erally rugose, medially polished and sparsely punctate; apophyses
rather stout; legs stout, fenmr hardly reaching apex of abdomen.
Abdomen very finely shagreened and subopaque beyond first tergite;
petiole slightly depressed, postpetiole distinctly broader than long,
a shallow impression on each side above spiracle; second tergite more
than a half longer than broad at base; ovipositor sheath nearly as long
as abdomen, ovipositor very stout, nearly twice as deep near apex as at
base.

Head and thorax black with yellow markings (see description of
color pattern, p. 141); occipital ring nearly or quite interrupted on
temple; annulus on flagellar joints (5)6-11(13) ; scape entirely black;
mesoscutum with a single median spot ; mesosternum yellow only along
sternaulices; propodeal markings extending broadly forward from
apophyses but not reaching carina. Front and middle legs yellow,
the coxae, trochanters, and femora black behind ; hind leg ferruginous,
a small spot at base of coxa above and the tibia and tarsus yellow.
Abdomen ferruginous, postpetiole apically and laterally yellow;
sheath black.

Type locality. — Trinidad Rio, Panama.

Types. — Three females, all collected by August Busck, the holotype
on March 29, one paratype on March 23, 1912, at the type locality, and
a paratyjje on June 3, 1007, at Tabernilla, Canal Zone. They are in
the U. S. National Museum (No. 57074).

The paratypes are slightly smaller than the type but otherwise
much the same.

13. CRYPTANURA ACINACES, new ipecies

Female. — Length lo mm., antennae (broken), ovipositor sheath
8 mm.

Structure very similar to that of excalibiir, l)ut abdomen blaclc with
yellow markings. From the above descript ion of fxra/J/fur the pi'esent
species differs principally as follows: Temples very distinctly convex;
horns on a common base; sculpture throughout thorax denser and
coarser; apophyses short and stout.

Head and thorax black and yellow (see description of coloi- p:ittcrn,
p. 141); mandibles entirely black; scape yellow below; annulus on
flagellar joints (5) 6 12 (13) ; orbital ring entire; mesoscutum with a
Finglc median yellf)w spot; scutellum entirely yellow; mesosternum
yellow only along sternaulices; propodeal markings extending broadly

166 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

forward from apophyses, but not reaching carina. Legs ferruginous,
tibiae and tarsi yellowish ; front coxa yellow in front, black behind ;
middle coxa yellow in front, piceous to ferruginous behind ; hind coxa
ferruginous, with a large yellow spot above and a small piceous spot
in apical middle ; wings yellowish, venation brownish ; abdomen black
and yellow, petiole entirely black.

Type locality. — Colima Volcano, Mexico.

Type.— OwQ female in the U. S. National Museum (No. 57075),
collected by L. Conrad.

24. CRYPTANURA PROPINQUA (Cresscn), new combination
Mesostenus propinquus Ceesson, Proc. Acad. Nat. Sci. Philadelphia, 1873, p. 152.

Female. — Length 13-16 mm.

Structurally very similar to aclnaces but with temples weakly con-
vex; apophyses rather slender.

Body color almost exactly as in aclnaces except that the mandibles
apparently are always yellow at the base and the orbital rings are
very narrow or even interrupted in the temples, and the scutellum
is more or less black at the base ; but the legs are yellow, with the front
and middle coxae, trochanters, and femora piceous behind, the hind
coxa with a broad piceous stripe on the outer side of the upper sur-
face and a shorter one on the inner side, the hind trochanter piceous
with a trace of yellow on the upper and lower surfaces of the basal
joint, and the hind femur piceous above and on the inner surface.

Before me are a paratype and another female from Mexico (Fron-
tera, Tabasco) ; a pair from San Jose, Costa Rica (M. Valerio, No.
101) ; two females from Costa Rica (Paul Serre, 1920) ; and one female
from Nicaragua (Mniszech, 1871), the last three received from the
Paris Museum.

The male has the malar sj^ace shorter, the frontal horns more slen-
der and separated, the apophyses reduced to high elevations in the
apical carina, which is distinct and sharply angulated medially, the
petiolar area defined laterally by curved carinae, the apical slope of
propodeum not distinctly transversely striate and the space between
the carinae longitudinally striate, and the abdomen small and slender
and subpolished.

25. CRYPTANURA PRETIOSA (Viereck), new combination

Polyacnidia pretiosa Viereck, Proc. U. S. Nat. Mus., vol. 46, p. 382, 1913.

This species differs from all the other species with red abdomen
which follow it in the present arrangement in its entirely red front
and middle femora and in the form of the yellow marking of the
lower division of the metapleuron. The latter is in the form of an oval
spot extending forward from the hind coxa, whereas in the other
species it either extends the entire length of the sclerite or has an
angular forward extension above.

ICHNEUiMON-FLIES OF GENUS CRYPTAXURA — CUSIIMAN 1G7

The type and all()(yi)e are in tlie K("ini<;liclip Zooloj^jisclie !Nru?ouni,
Berlin, the material examined in this study consist in«i; of a male para-
type from Villa Mora, Parajjuay, and a female from Georgetown,
British Guiana, identified by myself. The latter specimen agrees
perfectly with the original tlescription.

This is probably synonymous with spUonota (Cameron).

26. CRYPTANURA GRACILIPES. new species

Notable principally for its very slender legs.

FernaJe. — Length 14 mm., antenna 12 mm., ovipositor sheath 5.5 mm.

Temples distinctly concave; occipital carina high, especially at lower
end, where it curves sharply mesad to join the hypostomal carina
far back from the base of the mandible; vertex in front view slightly
concave; frons with a few short striae radiating from ocelli; horns
short and stout; eyes bulging, weakly convergent below; face some-
what elevated medially, polished and very sparsely punctate, somewhat
rugose above and in a shallow impression on each side of middle;
clypeus polished and sparsely punctate, subnasute in profile; malar
space nearly as long as basal width of mandible ; cheeks in front view
nearly straight; antenna 35-jointed, slender, distinctly though not
strongly thickened and flattened toward apex. Thorax rather unusally
smooth, pronotum with humeral margin carinately angled anteriorly,
upper portion, including humeral welt, polished and sparsely punc-
tate, scrobe striate; mesoscutum polished, sparsely punctate, lobes
weakly convex; scutellum narrow, convex, polished; mesopleuron
])olisht'(l and punctate below, obliquely striate above; sternum and
upper division of metapleuron polished and sparsely punctate; lower
division of metapleuron coarsely obliquely rugosostriate; pro])odcuni
basad of carina polish(Nl medially, rugose laterally, behind carina with
striae radiating from between apophyses, posterior face transversely
striate, bas;il median ai'ca defined, apophyses long and slender; legs
verj' slender, hind femur about eight times as long as deep, more than
three-fourths as long as tibia, reaching beyond apex of abdomen.
Abdomen rather slender, subpolished; fiist tergite polished, j^etiole not
depressed, postpetiole broader than long but not abruptly widened at
spiracles, rather weakly convex; second tergite jdiout three-fourths
longer than broad at base; ovipositor sheath distinctly shorter than
abdomen beyond fii'st segment; ovipositor slendei- juid of unifunn
depth to the distinctly subsagittate apex.

Head and thorax black and yellow, abdomen largely ferruginous
(.see description of color pattern, p. Ill) ; orbital ring broadly inter-
rupted in temple and narrowly so in malar space; scape entirely black;
anmilus on flugellar joints (5)0 10(12) ; mesoscutum with a single
discal spot; .scutellum black basally; tegula with a broad stramineous
and piceous margin; mesosternum yellow only along .sternaulices;

168 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m

metapleuron broadly black along lower and anterior margins; pro-
podeal markings barely including apophyses; front and middle legs
yellow with femora and front coxa black behind, middle coxa ferru-
ginous behind, hind leg ferruginous, the tibia and tarsus and a small
spot at base of coxa above yellow, femur with a narrow dark streak
above; wings hyaline, venation brownish; petiole yellow, postpetiole
piceous with apical margin narrowly yellow.

Male. — Like female but smaller and more slender; frontal horns
longer and more slender; antenna 38- join ted; propodeum with
apophyses short and thick, apical carina distinct and bent abruptly
forward medially; abdomen much more slender, second tergite more
than twice as long as broad at base. Annulus on flagellar joints
(9)10-16(18) ; postpetiole only very narrowly yellow at apex.

Type locality. — Trinidad Rio, Panama,

Types. — Two of each sex, a holotype female, allotype male, and a
male and a female paratype, all collected at the above locality by
August Busck, March 17-30, 1912. They are in the U. S. National
Museum (No. 57076).

The paratypes are somewhat smaller than the type and allotype.

27. CRYPTANURA GRACILIS, new species

Female. — Length 14 mm., antenna 12 mm., ovipositor sheath 5 mm.

Temples very weakly convex, more than half as broad as short diam-
eter of eye ; occipital carina moderately high, curving slightly inward
at lower extermity, in side view very nearly parallel to posterior
margin of eye ; vertex straight across ; f rons with striae radiating from
ocelli, horns short and thick, separated ; eyes weakly convergent below ;
face polished, sparsely punctate, transversely striate above and in a
shallow impression on each side of a low median elevation; clypeus
polished, sparsely punctate, very strongly convex ; malar space nearly
as long as basal width of mandible; cheek distinctly less than twice
as broad as malar space; antenna 38-jointed, slender, barely thickened
and weakly flattened toward apex, the joints in the thickened portion
fully as long as thick. Thorax distinctly compressed ; humeral margin
of pronotum transversely striate and with a strong carinate angle
anteriorly, scrobe striate; mesoscutum coarsely and rather densely
punctate, scutellum narrow; mesopleuron obliquely striate above,
finely punctate below; sternum finely punctate; upper division of
metapleuron punctate, lower division coarsely obliquely rugosostriate ;
propodeum with median basal area defined, lateral areas largely rugose
but polished and punctate in middle; area behind carina irregularly
rugose, posterior face coarsely transversely striate, apophyses long
and slender ; legs slender, hind femur fully six times as long as deep,
and fully three-fourths as long as tibia, but not quite reaching apex of
abdomen. Abdomen subopaque, finely shagreened beyond first tergite,

ICHXEUMOX-ILIES OF GENUS CRYPTANUHA — CUSHMAN 169

narrow, petiole not distinctly depressed, post petiole as lonp^ as broad,
weakly convex; second tergite about three-fourths longer than broad
at base, and hardly a half broader at apex than at base; sheath dis-
tinctly shorter than abdomen beyond first segment, ovipositor slender,
of nearly uniform depth to the subsagittate apex.

Head and thorax black with yellow markings (see description of
color pattern, p. 141) ; orbital ring interrupted in temple; clypeus yel-
low only in middle, the black of the suture extending upward in the
facial impressions; scape entirely black; annulus on flagellar joints
(6) 7-19 (12) ; mesoscutum with a single median spot ; scutellum black
in basal middle; mesosternum with a small trace of yellow next to
sternaulices; metapleuron broadly black below and anteriorly; meta-
pleuron posteriorly and metasternum ferruginous; propodeal mark-
ings extending only a short distance before apophyses; front and
middle legs yellow, black or piceous behind to apices of femora, middle
coxa ferruginous behind, hind leg ferruginous with tibia and tarsus
yellow; wings hyaline, venation blackish; abdomen entirely ferru-
ginous.

Type locality. — Villa Lutecin, near San Ignacio, Misiones, Argen-
tina.

Type. — One female in the U. S. National Museum (No. 57083) taken
by E. R. Wagner in 1910.

28. CRYPTANURA GENALIS, new species

Female. — Length 13 mm., antennae 11.5 mm., ovipositor sheath
5.5 nmi.

Very similar to gracilis, differing ])i-incipally as follows: Occipital
carina very high, in sitle view di\eigiiig below from postei'ior margin
of eye; frontal horns on a low common base; eyes parallel; clypeus
f)nly moderately convex; clieeks sli-ongly convex and fully twice as
broad as malar space; antenna 32-jointed, stouter and more distinctly
thickened toward apex, the joints in the ihicktned portion distinctly
transverse; mesosculnm very sparsely punctate, prescntum not striate
along notaulice.s; lobes not flatteneil; scutellum broader; lower divi-
sion of metapleuron punctatorugose; i)roj)o(leuiii without a basal me-
dian area, basal legion polished, sj)arsely punctate behind, more
densely and coarsely j)unctate behind caiina, rugosopunctate laterally,
posterior face closely transversely stii.ile. ;ij)ophyses very short and
thiek; legs stouter, hind femur not nearly six times as long as deep
aiifl barely two-thirds as long as tibia. Alxlonien stouter, postix'tiole
much bioader than long, second tergitc barely n Imlf longer than broad
at base; sheath as long as aixlomen beyond first tergito.

Orbital ring not interrupted in temjile; black of clypeal suture not
meeting in middle and not extending upward into facial impressions;
anindns on flugellar joints (5) 0-11 (13) ; mesostei-num cons|)icuously
yellow along sternaulices; propodeid maikings triangularly produced

170 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

anteriorly ; petiole tinged with yellowish and postpetiole with apical
margin yellow.

Type locality. — Venodio, Sinaloa, Mexico.

Type. — One specimen in the U. S. National Museum (No. 57077)
received from B. Preston Clark.

29. CRYPTANURA VARIEGATA (Brnlle), new combination

Mesostenus variegatus BruixI!;, Histoire naturelle des insectes, Hymenoptera,
vol. 4, 1846, p. 277, pi. 40, fig. 3.

Closely related to the two preceding species and differing from the
above description of gracilis only as follows : Temple flat, hardly half
as broad as short diameter of eye; clypeus only moderately convex;
antenna 32-jointed, distinctly thickened and flattened toward apex,
the joints there strongly transverse; scntellum rather broad and more
weakly convex; propodenm without defined median basal area, lateral
areas largely polished and sparsely punctate, rugose only laterally.
Abdomen rather broad, postpetiole broader than long, second tergite
more than a half broader at apex than at base.

Clypeus black only apically and laterally, the black not extending
upward into the facial impressions; metapleuron and sternum not
at all red; annulus on flagellar joints (4) 5-10 (12) ; propodeal mark-
ings extending triangularly nearly to basal carina.

One female in the U. S. National Museum, taken in October 1922 by
H. Lang at Kamakusa, British Guiana.

30. CRYPTANURA PARANENSIS, new species

Female. — Length 13 mm., antennae (tips broken off), ovipositor
sheath 6 mm.

Very closely related to variegata^ from which it is distinguishable
by its distinctly longer ovipositor sheath and by the fact that the
occipital carina diverges more strongly below from the posterior
margin of the eye. From variegata it differs further as follows:
Frontal horns on a common base ; thorax laterally less strongly sculp-
tured, the striation of mesopleuron confined to the upper anterior
portion, the punctation of the lower portion and of the sternum finer,
and the lower anterior portion of metapleuron smooth and polished ;
scutellum yellow only at apex and in basal angles; mesosternum en-
tirely black; propodeal markings extending only very sightly for-
ward from apophyses ; abdomen somewhat more slender.

Type locality. — Peixe Boi, Para, Brazil.

Type. — One female in the IT. S. National Museum (No. 57078) taken
November 27, 1907, by Miss H. B. Merrill.

What may be the male is represented by three specimens received
from the Paris Museum and collected by Manger in Brazil. The labels
also bear the figures 11-53. In these specimens the propodeum has
a distinctly defined petiolar area from the anterior end of which the
apical carina extends laterally and forward to the bases of the

ICHNEUMON-FLIES OF GENUS CRYPTANUIL\ — CUSHMAN 171

apf»i)hyses. The only complete aiitennii is 4;3-jointe(l with the aniuiliis
on tla<xellar points (10) Il-IG (lb). The postpetiole is piceous, the
niicldle coxae are very lar<>;ely ferruginous, while two of the
sjiecimens have the mesosterninn largely yellowish.

SI. CRYPTANURA TENUITEREBRATA, new species

Fcinale. — Length 17 mm., antennae 15 mm., ovipositor sheath 9 mm.

Conspicuous for its large size and long slender ovipositor, this
species is, nevertheless, very closely allied to pai'anensis and vaHegata^
tlilFering from the former principally by the characters employed in
the key. In structure and sculpture it agrees very well with variegata^
while in detail of color, especially of scutellum and propodeum, it is
more like parancn-sis. The flagellum is somewhat stouter basally than
in either of the other two species, the first joint of the flagellum being
distinctly less than six times as long as thick, and the antenna is
38-jointed, with the annulus embracing flagellar joints (5) 6-19 (14).
The ovipositor sheath is nearly as long as the abdomen and the ovi-
positor very slender, and of uniform depth to the distinctly subsagit-
tate apex.

Type locality. — Rurrenabaque River, Beni, Bolivia.

Typef<. — Two females, the holotype captured in November 1921
by "W. M. Mann on the Mulford Biological Exploration; the paratype
fr<jm Sapucay, Paraguay. Oct<;ber 19, 1902. They are in the U. S.
National Museum (No. 57079).

32. CRYPTANURA INCERTA (Cresson), new combination

Mcftostnuix incertus Cbesson, Proc. Acad. Nat. Sci. rbiladeliihia, 1873, p. 161,
ft'iiiale.

This and the species next following differ from the preceding six
species in the much shorter and stouter hind femur, which is barely
two-thirds as long as the tibia. The antenna in this group is some-
what more slender and less strongly thickened and flattened toward
the apex. Otherwise these s|X'cies are, in structure and color i)attern,
very similar to the preceding group.

Female. — Length 11 mm., antennae U mm., ovipositor sheath 4 mm.

Temple flat, very strongly receding, distinctly less than half as
broad as .short diameter of eye; vertex very weakly convex; frons with
striae radiating from ocelli, horns very small, on a common base; eyes
weakly convergent below; face ])olished and sparsely j)unctate,
striate above and in longitiidinal imi)ressions; cly|)eus very strongly
convex, polished and sparsely punctate; malar space nearly as long as
basal width of niainlible: mueh more than half as long as cheek; an-
tenna slender, weakly thickened and flattened toward apex, the joints
in the thickened portion weakly transverse. Thorax distinctly com-
pressed; humeral margin of pronotum transversely rugose and cari-
nately prominent anteriorly, .scrobes striate; me.so.scutum with lobes
flattened, polished, sparsely and coarsely punctate; scutellum narrow,

172 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

moderately convex, polished, with scattered punctures; mesopleuron
striate above, polished and sparsely punctate below, as are also the
sternum and the upper division of the metapleuron ; lower division of
metapleuron coarsely, obliquely striatorugose ; propodeum with basal
median area defined, lateral areas polished and coarsely, sparsely
punctate to rugose, area behind carina irregularly rugose, posterior
face transversely striate, apophyses rather long and slender. Abdo-
men rather stout, minutely alutaceous subopaque, petiole not depressed,
postpetiole broader than long; second tergite little more than a half
longer than broad at base, its sides broadly divergent; ovipositor
sheath hardly as long as abdomen beyond first segment, ovipositor
slender, of uniform depth to the subsagittate apex.

Head and thorax black and yellow (see description of color pattern,
p. 141) ; mesoscutum with a single median spot; propodeal markings
obliquely truncate shortly anterior to apophyses. Front and middle
legs yellow, with coxae, trochanters, and femora black posteriorly;
hind leg ferruginous with femur more or less piceous apically, tibia
entirely and tarsus except the more or less blackish apex yellow. Wings
slightly infumate, especially around apical margin. Abdomen ferru-
ginous, usually with apex of tergite 1 narrowly yellow and postpetiole
piceous, though sometimes entirely ferruginous.

Of this species I have examined six females and one male as fol-
lows: The National Museum female cotype from Mexico; a female
and a male taken by August Busck, March 23 and 27, 1912, at Trinidad
Rio, Panama; a female from Alhajuelo, Panama, April 7, 1911, A.
Busck ; and two females from an altitude of 2,000-3,000 feet, "Pinches
& Perene Vs," Peru, received from the Geographical Society of Lima.

The Peruvian specimens have the white markings somewhat less
extensive, with mandibles and sternum entirely black, and the first
tergite entirely red.

33. CRYPTANURA MACULIFRONS, new species

Female. — Length 11.5 mm., antennae (gone), ovipositor sheath 4
mm.

Similar to incerta, from which it differs virtually only as follows:
Mesopleuron obliquely striate over most of its surface; face white
only medially and in orbits, mandibles entirely black; propodeum dor-
sally, metapleuron below, and metasternum entirely reddish piceous to
ferruginous; white markings of thorax as in incerta except that the
mesosternum is entirely black; legs as in incerta except that the front
coxa is very largely piceous and the middle coxa is entirely ferrugi-
nous, as is also the abdomen.

Type locality. — Misiones, "Env. de San Ignacio, Villa Lutecia,"
Argentina.

Type.— One female in the U. S. National Museum (No. 57082) la-
beled "E. R. Wagner, 1900."

ICHNEUMOX-FLIES OF GENUS CRYPTANUIL\ — CUSHMAN 173
KEY TO ALL SPECIES OF CRYPTANURA*

This key is based partly on specimens and is partly compiled from
characters gleaned from published descriptions. Of necessity it de-
pends very largely on color characters and is purely artificial.

1. Thorax red and yellow, rarely partly black above 2

Thorax blaek and yellow, rarely propodeuiu and metasternum more

or less reddish 9

2. Head red and yellow ruficeps Cushman

Head black and yellow 3

3. Abdomen black and j-ellow 4

Abdomen more or less, usually largely, red 5

4. Mesoscutum immaculate black : mesopleuron red :

Propodeum immaculate red albomarginata (SzC'pligeti)

Propodeum with a black cruciform mark.

albomarginata var. (Sz^pligeti)

Mesoscutum red or piceous with a yellow median spot ; mesopleuron largely

yellow piceothorax Cushman

5. Mesoscutum with two small marlis discally 6

Mesoscutum with a single median spot 8

€. Mesopleuron white 7

Mesopleuron red bipartita (Brulle)

7. Apical carina not developed similis (Sz^pligeti)

Apical carina developed medially dicostata Cushman

8. Mesoscutum black variegata (Brull6) var. 4 (Sz«5pligeti)

Meso.scutum red rufa Cushman

9. Abdomen red 10

Abdomen bla/k and yellow 36

10. Scutellum conical 11

Scutelium not conical 12

11. Metapleuron subconically elevated tuberculata Cushman

Metapleuron not elevated scutellaris (Sz^pligeti)

12. Hind femur black above, red or yellow below 13

Hind femur red, at n)ost black apically 15

111 Hind femur yellow below lucida (Szc^pligeti)

Hind femur red below 14

11. Scutellum entirely yellow basimacula (Cameron)

Scutellum yellow only at apex volcanica (Cameron)

1'). Hind C'lXa black and white 16

Hind coxa red 17

10. Humeral margin of pronotum conically elevated on each side.

politigaster Cushman

Humeral margin of pronotum not conically elevated coxata Cushinan

17. Proi)odeuni with a pale spot in each anterior angle 18

Propodeum immaculate ba.sally 19

I'trrohtris ( Szejiligef i ) , lioplcuris (Sz«'pligetl), and longipes (Sz^'pligeti)
and its male variety run to 17 but no farther by the descriptions.]

' Spool flc nnmoH whoHc authors nro Riven In parcnthrHos nrp here for the flfRt time nneS
In conitilnnf Ion with titp kpiktIc nniiic Cryplniiurn, oxcfpt In tlio rnno of rrli/pn (CrrsHon)
and npinnria (Unill*'-). Tlip nnnic-H Intnrntnria Ciinioron, mrxlrana ("roHHon. jirniilnqua
CrcHson. nn<l vcrarpacin CnmfTon wpr« formorly uiifli-r MctoKtrnua. The re«t of the nnnica
here transferred have formerly been under I'lilydinua.

174 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 95

IS. Propodeum with yellow spots basally uniformis Brues

Propodeum with reddish spots basally rugosa Brull4

19. Middle femur entirely black fusciventris (Cameron)

Middle femur not entirely black 20

20. Front femur and usually middle femur black or piceous behind or below 21

Front and middle femora entirely red 32

21. Propodeai markings each with a narrow, tonguelike forward extensioa

before apophyses 1 22

Propodeai markings truncate basally 23

22. Postpetiole red ; upper margin of pronotum tuberculate on each side anteriorly.

boliviensis Cushman
Postpetiole piceous; upper margin of pronotum merely carinate anteriorly.

isthmus Cushman

23. Mesosternum white at least along sternaulices 24

Mesosternum not at all white 2&

24. Hind legs very slender, femur apparently nearly 8 times as long as deep;

temples in dorsal view very sharply receding and slightly concave.

gracilipes Cushman
Hind legs stouter, femur apparently not more than 6 times as long as deep ;

temples flat or weakly convex 25

[variegata (Brulle) varieties, nigripes Brull6, and striata Brulle run
to 24 but not farther by the descriptions.]

25. Hind femur short, distinctly less than two-thirds as long as tibia ; ovipositor

sheath barely twice as long as first abdominal segment incerta (Cresson)

Hind femur at least two-thirds as long as tibia ; ovipositor sheath distinctly
more than twice as long as first segment 2ft

26. Cheeks fully twice as broad as malar space; temple rather broad, its angle

with the longitudinal axis less than 45 degrees ; ovipositor sheath much

shorter than abdomen : 27

Cheeks much narrower ; temple narrower, its angle with the longitudinal axis
moi'e than 45 degrees ; ovipositor sheath nearly or quite as long as
abdomen 28

27. Apophyses short conical ; lobes of mesoscutum strongly convex and sparsely

punctate; postpetiole broader than long, margined with yellow,

genalis Cushman
Apophyses long; lobes of mesoscutum flattened and densely punctate; post-
petiole as long as broad, not margined with yellow gracilis Cushman

28. Hypostomal carina distad of occipital carina shorter than malar space ; ovi-

positor slender and of nearly uniform depth except at apex.

tenuiterebrata Cushman
Hypostomal carina distad of occipital carina as long as malar space, very
high ; ovipositor stout, much deeper near apex than at base.

excalibur Cushman

29. Head in dorsal view with temples concave and much less than half as long

as short diameter of eye ; scutellum black medially paranensis Cushman

Temples flat and nearly half as long as short diameter of eye ; scutellum
entirely yellow , 30

30. Middle coxa black and yellow ; face entirely yellow atripectus Cushman

Middle coxa red ; face with a black stripe on each side of middle 31

31. Ovipositor as long as abdomen and much deeper apically than basally.

platyurus (Brulle)
Ovipositor much shorter than abdomen and of unifoi'm depth.

maculifrons Cushman

ICHXEUMOX-FLIES OF GENUS CKYPTAXURA — CUSHMAX 175

32. PropcMh'al spots with ton^io-shapcil oxtonsions nntoriorly_ vnriegata ( I?iull«5)
I'ropiMk'al siKjts witliout such extoiisidiis anteriorly or, if extciuliiif; forward

from apophyses, the extensions truncate or tapering 33

33. Mesosternum hhick 34

Mesostenunn at least partly yellow 35

34. Posti>etiole black hyalina Brull6

PostiK>tiole red conica Cushuian

3-~». Probably synonymous spilonota (Cameron)

pretiosa (Viereck)

36. Propodeum with two yellow spots basally 37

Propodeuni immaculate basally 42

37. Me.^oscutum immaculate discally though sometimes with yellow marginal

markings 38

Mei^o.scutum with two small markings discally 41

38. Mes().scutum immaculate ectypa (Cresson)

Mesoscutum with yellow marginal markings 39

39. Mesosfutum with yellow lines laterally opposite tegnlae.

quadrimaculata Cushman
Mesoscutum with cuneiform markings on anterior lateral margin 40

40. Prescutum yellow on each side nitidiuscula (Cameron)

Presi'utuin immaculate mediostrigosa Cushman

41. Hind femur black posteriorly bicarinata Cushman

Hind femur with a black line above championi (Cameron)

42. Me.so.scutum with a .single median spot or immaculate, rarely (spinaria) the

median spot flanked on each side by a small mark on inner margin of

lateral lobe 43

Mesoscutum with two discal marks 51

•13. Meso.scutum immaculate ablata (Cresson)

Me.so.scutum maculate 44

41. Prop( ideal spots with tongue-shaped anterior extensions, which rarely become

confluent across median line 45

Propodeal markings without such extensions 49

4'). Hind coxa red ( 9 ) or black with a yellow spot above ( $ ).

spinaria (Brulle)

Hin<l coxa yellow with black markings above 4l>

4";. Hind femur red septentrionalis Cushman

Hind femur black and yellow 47

47. Hind fennir bhuk both dorsally and ventrally planiscutellata Cushman

Hind femur black only dor.sally 48

48. Hind femur with a narrow median yellow line dorsally.

lineatifemur Cushman
Hind femur entirely black dorsally veraepacis (Cameron)

49. Hind coxa and femur red acinaces Cusliman

Hind coxa and fenrur blark and yellow 50

50. Mcsosterimm yellow; r»vip<».sltor sbrath shorter than abdomen.

lamentaria (Cameron)

Meso.Hternuin yellow only lalfrally and on each side (if iniddle [xistfriorly ;

ovli)o.sllor sheath subeqiuil to abdomen In It-nglh propinqua (CresHon)

51. Me.sopleuron with a large yellow s|HJt anteriorly and a small one p«isteriorly ;

sternum, p4'tioIe, and hind femur entirely black apophysis Cushman

Mesopleuron with n single large ol)li(|ue mark, Joined jMisleriorly to a yellow
line on sternum; petiole partly yellow; hind fennir black only above 52

176 PROCEEDINGS OF THE NATIONAL MUSEUM voi-. 9s

52. Petiole black above, yellow below; hind femur with a median yellow line

above orizabensis (Cameron)

Petiole black below, yellow above; hind femur entirely black above 5S

53. Mesosternum largely yellow ; propodeal spots broadly truncate before

apophyses, the latter long mexicana (Cresson)

Mesosternum yellow only along sternaulices ; propodeal spots narrowed and
tonguelike before apophyses, the latter short bilineata Cushmaa

ADDENDUM
CREMNOCRYPTUS, new genus

Genotype. — Polyaenus spiniferus Cameron.

This genus will run in all existing keys to {Polyaenus Cresson) =
Oryptanura Brulle, but the frontal horns are of quite different form,
being flattened below and compressed above and separated by a deep
groove. It also differs by the following characters : Ocelli situated on
the sides of a distinct, sometimes very high, elevation; antenna in
female only slightly thickened, slightly flattened on the outer upper
side but not below; occipital carina strongly sinuate at lower ex-
tremity; mandible distinctly tumid at upper basal angle; upper
margins of pronotum tumid, but not angulated anteriorly by the
epomia, which are weak or obsolete; notaulices shallow and extend-
ing only about halfway to scutellum ; sternaulices short and shallow ;
scutellum very broad, very weakly convex, and with sparse, coarse
punctures; nervulus interstitial or very nearly so; abdomen coarsely
and deeply punctate on basal three tergites, male abdomen fusiform;
ovipositor neither distinctly subsagittate nor swordlike, but rather
tapering and distinctly flattened above at apex.

CREMNOCRYPTUS SPINIFERUS (Cameron), new combination

Polyaenus spiniferus Cameeon, Journ. Straits Branch Roy. Asiatic Soc, No. 46,
p. 117, 1906.

Five specimens of each sex from Borneo, one female from Mindanao,
and two females and one male from Singapore are before me. These
agree exactly with Cameron's description.

CREMNOCRYPTUS CINGULATUS (Tosquinet), new combination

Polyaenus cingulatus Tosquinett, Mem. Soc. Ent. Belgique, vol. 10, p. 45, 1903.

There can be no doubt that this New Guinea species is congeneric
with the genotype. A considerable series, including both sexes, from
Mindanao, Basilan, Samar, and Luzon belong, I suspect, to the species,
though I hesitate definitely to identify them as such. They differ
from spiniferus in having the stemmaticum strongly elevated above
the ocelli ; the thorax, especially the posterior face of the propodeum,
much more sparsely sculptured; the basal tergites more coarsely and
less densely punctate; the antennal annulus constantly shorter; and
from all but the Mindanao specimen in the blackish color of the entire
inner side of the hind tibia.

U. S. SOVERHMENT PRINTING OFFICE: 1948

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued i'-f^jNoA, VJiJ^ l>y '^

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSELIM

Vol. % Wuhington: 1945 No. 3194

NEOTROPICAL LANTERNFLIES OF THE GENUS PHRIC-
TUS IN THE UNITED STATES NATIONAL MUSEUM,
WITH DESCRIPTIONS OF FOUR NEW SPECIES

F)V John S. Caldweix

The Neotropical genus P/irlcfy^<i (Homoptcra: Fulgoridae^) was
established by Spinola in 1839- for the unique species Fulgora dia-
dema Linnaeus. Since that time various species have been described
by Signoret, Distant, Schmidt, Lallomand, and Metcalf; in 1905
Schmidt presented a key to the known species, which was modified
by Metcalf in 1938.

For such large and truly spectacular insects the species are relatively
little known and their classification is in a confused state, probably
because their descriptions have Ix^en extremely inadequate and very
few have been illustrated. Specific identification has been based large-
1\' on color and marking, characters that are often variable in intensity
and exactness of pattern, wliile the structural characters generally
emphasized concern the fmni of ihc giDtesque cephalic process, which
in most cases cannot he adequately described. The length of this
structure relative to the length of the pronotum has, in the past,
served to segregate llu' species into gr(»Mi)s, which in tiiiii have been
broken down into the respective species on the basis of color. This
methofl (»f identification i-' almost a neressary evil, because the most

' Kvon Hioiit'h Fulfjorn I.lnnnciih' jm n Hynonyni "f Lntrrnarin IJnnnoiiH (Fonnnh. Vrnr.
Ulol. Sor. W.-iHhlni:tr«n. vol. .17, pp. 43-44, 1044), In the Hplfcllon of ii family nnmo I follow
the* prlnrlplc proiuiHod by C. W. SnIiroHky (V<tIi. VII Hit. KonK. Knt., vol. 1. pp. fldU 0(13.
1939).

' Ann. Sor. Knf. Frnnre. vol. 8. pp. 218-221. 18.30.

177

«2M»r)7 1.')

178 PROCEEDINGS OF THE NATIONAL MUSEUM vol.88

pertinent structural characters of both the cephalic process and the
genitalia defy description. The median notch in the caudal margin
of the pronotum and the furcation of the median carina around this
notch also furnish constant specific characters. In final analysis the
male aedeagus, first illustrated by Metcalf, is of excellent value for
identification purposes, but it must be remembered that the aedeagi
in the Fulgoridae are inflatable and that they present a very different
appearance when inflated than when deflated. The females possess
good genital characters in that parts of the first valvulae are heavily
sclerotized and ornate with spurs and ridges. Although these valvulae
are not radically differentiated among the species, they are relatively
constant wfthin each species, and a comparison of the accompanying
drawings will readily demonstrate their specific value.

The purpose of this paper is to present the more pertinent structiu'al
characters of the genus, together with characters of color and pattern,
in the hope that some of the present confusion may be cleared up and
that recognition of the species may be made a much easier task.

Because four undescribed species are added here, and because
notatus Lallemand is not included in any existing key, a revision of
the key is presented here. Unfortunately, xanthoptei^A Schmidt^
and notatus Lallemand * are known to the writer by their descriptions
only. The order in which the species are discussed follows a tentative
phylogenetic arrangement.

ARTIFICIAL KEY TO THE KNOWN SPECIES OF PHRICTUS

1. Cephalic process flattened apically, transversely arcuate, lacking definite

apical teeth auromaculatus Distant

Cephalic process with 5 apical teeth quinquepartitus Distant

Cephalic process with 3 apical teeth 2

2. Hind wings with large hyaline apical spots ocellatus Signoret

Hind wings without hyaline apical spots, but small pruinose areas some-
times present 3

3. Elytra scarlet, continuous broad yellow transverse fasciae present.

tripartitus Metcalf
Elytra some shade of green or brown, transverse fasciae if present inter-
rupted medianly 4

4. Basal area of hind wing yellow, golden, or orange 5

Basal area of hind wings red to scarlet 8

5. Black or fuscous area in hind wings covering apical three-fourths ; cephalic

process longer than pronotum regalis, new species

Black or fuscous area in hind wings covering apical one-third or less ; cephalic
process shorter than pronotum 6

6. Basal area of hind wings golden yellow; elytra flecked with yellow and

brown xanthopterus Schmidt

Basal area of hind wings orange 7

3Bnt. Zeit. Stettin, vol. 71. pp. 144-14R. 1910 (Ecuador).
<Ent. Tidskr., vol. 52, pp. 188, 1931 (Ecuador).

L.\NTERNFLIES OF GENUS PlIHICTUS — CALDWKLL 179

7. Klyini jellowisli, irregularly Uiiisculale uver all with brdU n ; liiiul wings not

uiaoulate sordidus, new species

Elytra brown, inaeulute in basal two-tbirds with small round orange sjiots;

hlud wings maculate with brown notatus Lallemaud

8 (4). Elytni green in basal two-thirds, with a few small round red or orange

maculae; apical third with large brown maculae 9

Elytra brown or reddish brown 10

It. Tet'tli in ajHi ot' cephalic process obtuse, somewhat deflected caudad ; median

carina of pronotuni delinitely forke<l caudad inoebiusi Schmidt

Teeth in aix'X slender, not dellected caudad; pronotal carina not delinitely
furcate hoffmannsi Schmidt

10. Elytra some shade of reddish brown, with pink calloused areas present, es-

I)ecially basally; cephalic process toothed on ventral surface.

diadema (Linnaeus)
Elytra brown, maculate with y«'llow ; cephalic process smooth beneath__ 11

11. Combined length of c«*phalic pincess and head as long as pronotum; expanse

of trifurcate aiM'X equal to distance between (xular si)ines.

minutacanthis, new species
Combined lengrth of cephalic process and head longer than pronotum; ex-
panse of apex much greater than distance between ocular spines.

punctatus, new sp<H'ies

PHRICTUS AUROMACULATUS Distant

Pi-VTK 7. Kk.ukks C. L'4: F\j>.tf. S. Fu.ukks 1, IL' ; Plate 0; Pl.\te 10

Phrk-tu.s uuromaruhitu.s I)isrANr, Aim. Mag. Nat. Hist., vt»l. Hi. ser. 7. pp. 072-673,
l".m.').

'riieie may Ije .some doubt regiirdiiij^ the identity of this species,
because Di.staiit makes no definite statenuMit as to whethef the apex
of the ceplialic pi'ocess is trifurcate or flattened. He <loes mention tlie
carinate aiileiior marojiii and states tliat the process is sliorter than in
previously (h'.scfibed species. In the specimens studied not only is
the pfocess veiy short liut also the apt'X is flattened and tiansverse.
The cohjr of the hind wings is golden and not bi'ight yt'ih)w ; otherwise
the description matches the specimens. Length oxer all, male .■*.."' mm.,
female oG mm.; elytra. male2G nun., female i'!) mm.

Male bearing the data: "Tiimiiifasa, Bolivia. Dtc. Mill ford Biol.
Expl.. I'.t'-M I'-ii'-J (W. M. Mann)"; female, "Ixiamas. Bolivia. .Miilfoid
Biol. Expl., lU-2\ -I ( M. K. I>)pez)."

'I'ype locality : Pxili via.

I'MKK TIS OCKI.I.ATrS Siiinont
I'l.ATi-: 7, Ki<ii iiKs I», 21 ; Pi_\tk S. FniUiiK W
I'hrittuK urrlluluH SK.NoKi'rr, null. Soc. Km. i'niiice, vol. :',, ,ser. .3. p. v, 1S.'">5.

As yet this is the only known .species with large hyaline areas williin
the dark area of the hind w ings. The (•( plialic process is htnger than
the pronotum. The caudal margin of the pronotum is dee[)ly not<he<l

180 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

at the middle with the lateral margins of the notch sharply elevated
and acute caudad. Length over all approximately 41 mm.; elytra
32 mm.

One badly damaged female from "Colom." [Colombia?], Baker col-
lection.

Type locality : Venezuela.

PHRICTUS SORDIDUS. new species

Plate 7, Figukes 3, 20 ; Plate 8, Figures 1, 13 ; Plate 9

Allied to ocellatus in general pattern of elytra but lacking ocellate
spots in the hind wings. Length, male 40 mm., female 46 mm. ; elytra,
male 30 mm., female 37 mm.

Dorsum of cephalic process, vertex, clypeus, and median stripe of
pronotum light gray. Lateral and ventral margins of process dull
cinnamon. Lateral areas of pronotum chocolate-brown. Elytra brown
on basal two-thirds, irregularly maculate with dull yellow; broad
transverse fascia present at base of reticulate area, broadly interrupted
in center by a conspicuous brown dash. Apical third light yellow,
maculate with large, irregular brown spots. Hind wings dull faded
orange in basal two-thirds, apical third fumate.

Cephalic process shorter than pronotum ; trif urcate apex with very
obtuse teeth, especially laterally; median tooth somewhat deflexed
caudad. Posterior margin of pronotum broadly notched; median
carina ending at base of notch. First valvulae in female not trif urcate
apically.

Male holotype, U.S.N.M. No. 57224, Ecuador (Goodfellow), Goding
collection. Female allotype, Quevedo, Ecuador (F. Campos R.), and
one headless female, Ecuador, Goding collection.

These specimens appear very faded, and it is unknown whether the
color is natural or the result of immersion in preserving fluid. With
regard to color they resemble the species figured and described by Met-
calf as diadema; however, the cephalic process is distinctly shorter
than the pronotum and therefore they cannot be that species.

PHRICTUS MINUTACANTHIS. new species

Plate 7, Figures 2, 16 ; Plate 8, Figure 2 ; Plate 9

Resembles ocellatus in color and marking but differs from it in lack-
ing the ocellate areas in the hind wings and having darker brown
elytra. It is much smaller and more brightly colored than sordidus.
Length, female 37 mm., elytra 30 mm.

Entire venter, clypeus and face, lateral area of cephalic process, and
dorsum of prothorax except median stripe dark brown. Dorsum of
head, cephalic process, and median stripe of pronotum dark gray with
apex of cephalic process rose to red. Elytra brown, fairly evenly

LANTERN FLIES OF GENUS PHRICTUS — CALDWELL 181

maculate with dull yellow, transverse fascia at base of reticulations
irre<;ular and imperfect. Hind wings light red in basal two-thirds,
apical third fuscous.

Cephalic process slemler, shorter than pronotum; tritid apex very
small, especially the median tooth. Caudal margin of pronotum with
broad median notch, outer angles of notch abruptly acute and greatly
elevated; median carina of pronotum apparently not forked around
notch. First valvulae in female short, broad; apices more sharply
angled than in oceUatus.

Female holotype, U.S.N. M. No. 57225, fium Chaciuimayo, Peru,
February 15, IDlS (C. H. T. Townsend).

PHKICTUS PUNCTATUS, new upecieb
Pi^TE 7, FiiiUBES 12, ii3 ; Platk 8, Figire 6; Pi-iTK I)

Siuiilar to ininutacanthu but much larger and with more brightly
colored elytra. The cephalic process is longer, with the apical teeth
more acute. Length, female 44 mm., elytra 32 mm.

Clypeus very light brown. Face and lateral margins of cephalic
process cinnamon-brown. Vertex, dorsum of cephalic process, and
median stripe on pronotum dull yellow. Elytra brown w^ith bright-
yellow maculae groupt'tl toward the transverse fascia; claval area dull
yellow; costal margin with two conspicuous yellow spots before
leticulate area; transverse fascia bright yellow, narrow, interrupted in
nnddle by a dark-brown dash more or less surrounded by yellow
maculae. Maculae in right elytron tending to form an oblique strip
U'tween the dark-brown ditsh and center of clavus. Hind wings
hyaline-carmine in basal two-thirds, apical third light fuscous.

Cephalic process equal in length to pronotum; trifid apex slender,
with broader expan.se than width of head including eyes. Caudal
margin of pronotum broadly concave, with a small notch in middle of
concavity. Median carina definitely forked caudally around notch,
with surface of pron(;tum on each side elevated above the forked
carina. First valvulae in female with apices broadly concave on outer
margins.

F'emale holoty|)e, I'.S.N.M. No. 5722G, bearing the following data:
"? IJugaba, Panama, collection Wm. Schaiis.'' One female paralype
fiom El Volfdn, Panama, March 17, 11)43, is in the collection of the
American .Museum of Natural Plistory.

PHRICTUS HOFf'MANNSI Srhmidt

1'i.An-; 7, FtoUKKH 10. 18; I'i.ait-: 8, Ji'k.ukk J> ; Tlatk 10

I'hrictUH hoffmantiHi S< hmiht. Knt. Z<-1|. Sit-tiin, vol. iV,, pp. ,'{,'i8 .'140, l!M>r>.

This s|)ecies most clearly resembles moebiu.si Schmidt but is dis-
tinguisiusJ frouj it by a narrower cephalic process, with (he apical

182 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

teeth slender and lying in the same place. It also resembles notatus
Lallemand in having the elytra sometimes brownish and maculate,
with small, round orange spots, but is separated from notatus by hav-
ing the basal areas in the hind wings red and immaculate instead of
orange and maculate with brown.

Length, male 37 mm., female 46 mm. ; elytra, male 28 mm., female
37 mm. Unfortunately, pests have destroyed the internal male
genitalia.

Represented in the collection by one male with no accompanying
data other than "Goding Collection,-' one female from below Macas,
Ecuador (E. W. Rorer) , and one female from Baiios, Ecuador, altitude
1,800 meters (F. Campos R.).

Type locality: Peru.

PHRICTUS MOEBIUSI Schmidt

Plate 7, Figures 11, 15 ; Plate 8, Figube 5 ; Plate 10
Phrictus mofbiusi Schmidt, Ent. Zeit. Stettin, vol. 66, pp. 340-342, 1905.

Resembles hoffmannsi in general appearance but differs in having
apical teeth of the cephalic process more obtuse, with the median tooth
deflexed caudad, elytra possibly lighter green, and with maculae fewer
in number but brighter, and apices of first valvulae of female very
acute. Hind wings differ from those of notatus in that the colored
areas are red and scarcely maculate.

One female bearing the following data : "Medellin, Vy. and Porce"
(F. L. Gallego M.). Probable locality : Medellin, Colombia.

Type locality : Colombia.

PHRICTUS DIADEMA (Linnaeus)

Plate 7, Figures 1, 4, 13. 14 : Pi^te 8, Ficiures 11, 14 ; Piatb 10
Fulgora diaderna Linnaeus, Systeuia naturae, ed. 12, vol 1, p. 703, 1767.

Although dkulenia is one of the more common species in the genus,
its exact status is evidently still confusing. It was first figured by
kStoU,^ but either the figure is erroneous or diaderrm is unknown today.
The hind wing in the figure is unlike that of any known species, and
the black color includes a much larger area than in typical diaderna.
The figure by Drury ° (as annata) shows a much more typical pat-
tern, while the photographic reproduction by Costa Lima ^ is dmderna.
as accepted by most workers today.

The general color of the elytra varies from light buff to dark shades
of brown and sometimes even appears greenish ; however, in all this

" Representation exaetement colorize d'apres nature des Cigales et des Punaises
(Cigales), pi. 5, fig. 22, 1780.

* Illustrations of natural history, vol. 3, pi. 50, fig. 4, 1782.
' Insetos do Brasil, vol. 3, p. 45, fig. 40, 1942.

LANTERNFLIES OK GENU8 IMllUCTUS CALDWELL 183

variation pinkish oalloiisod areas aiv always |)n'S('Mt in the basal por-
tions. The tiifuirato a|i(\\ of the ccpiialic i)roccss exhibits much
variation as to size, deHeclion, antl form of the teeth. Tlie shape of
the internal male fronitalia is constant and best shown by tlic illustra-
tion (pi. s, tier. 14). The apices of the Hrst valvulae in the female are
bluntly tiifurcate. Five females ranc^e in len<»;th from 4S to 52 mm.
and four males from 41 to 46 mm.

Records indicate that this spi'cies ranjjes throu<::h the (iiiianas into
Brazil; also it has been recorded as a minor pest of cacao in Bahia,
Brazil. In a letter to the writer, Pedrito Silva states that his reference
to quinqiu'pnrfitiin Distant appearing in Tropical Agriculture^ is in
error, as the pest on the cacao tree is diadenia (Linnaeus) and not
quinqiu' part it ?/.*? .

Type l(x.'ality: '"Indien" (probably Brazil).

PHRICTUS REGALIS, new species
Pl.\te 7, rn.UKKs 7. lit; Pi..\TE 8, F^atTRE 8; Piatf, 10

Greatly resembling dwdema when the elytra are closed but with only
the basal fourth of the liind wings yellow. Length M mm., elytra 36
nmi.

Median stripe on pronotum, vertex, and dorsum of cephalic process
gray; apical teeth red. Venter of cephalic process brown, becoming
fu.-cous towaivl clypeus; clypeus light yellowish. Elytra deep oliva-
ceous, with calloused ai'eas and transverse fascia light red; costal and
apical mai-gin lightly washed with black. Hind wings with basal
fourth yellow, remainder black, with two or three large red spots
present halfway to apex and three or four large yellow spots present
in the black area neai" the basal fourth.

Cephalic process stout, longer than pionotuni, l(Migtli and xcrtical
height about equal; trifurcaic apex with I(»ng acute teeth. Caudal
inai-gin of pidiiotnm scarcely in<lentc<l medianly; median carina
appearing deeply bifiwcate. (Unfoi'tunately. most of this ai-ea is
obliterated by a large pinhole.) 'J'rifurcale ajjices of Hrst female val-
vulae with apical aiul outer teeth acute, inner teeth bliinl in latei-al
aspect .

Female holotype. U.S.X.M. No. r)7227, from Maroni Kiver. Ficnch
(iiiiaiia. vicinity of I)Ms('i-rc ((i. Molicrg).

IMIKKTUS UriN(JUKPAUTITUS Dintnnl
I'l.AIT. 7, l-'KifRKS K, 17; I*T,ATK K, F-'KilKKS 10, ].'»; ri.AIK'.l

I'hii'iuK iiuintfiu p'lrliluM Disiant. Itiuldiriji < 'ctil raii-AiiM'ricatia. IIi)innptrra. vnl. 1.
P 2t. pi t. flir. 8. 18.S:?.

This unii'^ua! '-pecies closely resembles triparlitux Mel calf in color
and marking but is ilistinguisherl from ii bv the apex df the cej)halic

•Trop, Acr. vol. 21. p. 12, 1044.

184 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

process bearing five teeth. In the 15 specimens examined the apical
teeth vary from acute to obtuse, and sometimes the intermediate teeth
are much reduced in size. The longitudinal veins in the elytra are not
so green as in Distant's illustration, and the bluish-white pruinose
spots in the dark apical area on the hind wings are sometimes absent.
In addition, the red areas in the hind wings are often less maculate
apically with black. The shape of the internal male genitalia appears
to be veiy close to that figured by Metcalf for tripartitus ; however,
the expansion of the inflatable sacs is much less, and the writer be-
lieves that these two species are distinct. As in diadema the two sexes
vary considerably in length, the males ranging between 43 and 45 mm.
and the females between 47 and 49 mm.

All specimens are from Panama and the Canal Zone.

Type locality : Panama.

PHRICTUS TRIPARTITUS Metcalf

Plate 7, Figures 5, 22 ; Plate 8, Figxjre 7 ; Plate 9

Phrictiis tripartitus Metcalf, Bull. Mus. Comp. Zool., vol. 82, p. 365, pis. 20, 21,
1938.

Metcalf believes that this species may be the unnamed variety of
diade/ma described by Walker ^ and figured in the Biologia.^" In
general color, pattern, and size it approximates specimens of male
quinquepartitus in the collection. Although the aedeagus of the male
is of tlie same general form in both species, it is much more inflatable
in quinquepartitiis ; in the female there are differences in the first
valvulae.

In the specimen believed to be tripartitus the teeth in the trifurcate
apex of the cephalic process are very obtuse but between the median
and right-hand tooth is a definite bump that may be construed as a
vCvStigial tooth; on the other side of the median tooth the margin is
crenulate. This specimen, measuring 51 mm., is longer than any speci-
men of quinquepaTtitus. The difference in total length is probably
accounted for by the longer cephalic process possessed by tripartitus.

Female plesiotype from Virginia, Guatemala, November 1915 (Wm.
Schaus) .

Type locality : British Honduras.

" List of homopterous insects in tlie collection of the British Museum, p. 264, 1851.
'" Biologia Centrali-Americana, Honioptera, vol. 1, pi. 4, fig. 5, 1883.

U. S. GOVERNMENT PRINTING OFFICE. 1945

U S NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 7

CEPHALIC PROCESS IN PHRICTUS.

Frontal profile of riylil half: 1. diaJema (fcmaU-); 2, minularanllns (male); 3, jorduhu
(male); 4, diadrma (male); 5, triparlitus (entire profile, female); (u auromacualus ( <;niaU-):
7 rfgalis (female); 8, quinquepartitw: (male); 9. nrrllalus (female, imperfect); lO.V'o/-
mflnn/i (female); 11, mo/-tiM.rt (female) 12, punctalui (female) , ^ ,^ * •

Lateral profile: 13, diadniu, fmale); 14, diadrmu (female); Is, ».oW;;»u (female); 16 m,«u/fl-
canlhis (male); 17, quinqiupartilus (male); IS, huffmannn (male); P. ra'fl/i.f (f<-maie);
20, jnrdidus (female); 21, ncfllaius (female, imperfect); 22, irtpartilui (female); 23, puncta-
lus (female); 24, auromarulnius (fenjale). .

Owing to their larv'c si/.e, rrRali^, diadnna, nui'u,nrt>,ir!ttu<. and iriparlUus are drawn approx-
imately one-half scale.

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE

Valvula and Genitalia in Phrictus.

Lateral and ventral profile of first female valvula (left): 1, sordidus; 2, minutacanlhis; 3,
ocellatus; 4. auromaculatus; 5, moebiusi; 6, punctatus; 7, tripartitus; 8, regalis; 9, hoffmannsi:
10, quinqueparlitus; 11, diadema.

Ventral aspect of male aedeagus: 12, auromaculatus; 13, sordidus; 14, diadema; 15, quinque-
parlitus.

U S NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 9

AUROhlACULATUS

PUNCTATUS

MINUTACANTHI S

SORDIDUS

^- •:* V

■^

TRI PARTITUS

QUINQUEPARTITUS

SPECIES OF PHRICTUS DORSAL ASPECT
auromaculalus, female; punclalus, female lioldtypc; minulacanthuf, female licl-nyiH-
sordiduj, female allcjtype; Iripartitus, female; <iuinqufparlUu<, male.

U. S. NATIONAL MUSEUM

PROCEEDINGS, VOL. 96 PLATE 10

•\

REGAL/ S

DIADEM A

HOFFMANNSI

MOEBIUSI

AUROMACULATUS

DIADEM A

SPECIES OF PHRICTUS: DORSAL ASPECT.

egalis, female holotvpe; diadema, female; hofmannsi, female; moebiiisi, female; auro-
maculatus, male; diadema, male.

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issited 5' J5(V '^■^ 0?'l^i by the
SMITHSONIAN INSTITUTION

L'. S. NATIONAL MUSEUM

Vol. %

Wuhington : I94S

No. 3195

HYPOKIIAMPIirs l^VTRIS. A NEW SPECIES OF HEMI-
KAMPIIID FISH FUO^r SIXALOA. MEXICO, WITH AN
AXALYSIS OF THE OEXEKIC CHARACTERS OF HYPO-
RHAMPH US AND HEMIRAMPHUS

Hv RoilIIfT R. MlI.LKR

Till. |i.iii<iiy of oui- kiiowU'tl^^c of tlu' fresh-Av titer fish fauna of
MortliWfstiTii M('.\i<-o is cNident from the novelties which Ralph G.
Milh-r has c'<tnected in that re*;ion in recent years. In addition to
the distinctive I)oroi<omn fon'ithl Hubbs and Miller (19-H) and a new
(riht being described by nie in Copeia, a new species of hallbeak of tlie
;zenus IlyporhamphuJi is now made known.

About GO years a^^o Meek and Goss (1885, p. 221) wrote that the
American halfbt-aks rcfeired to /// mh-t/mp/n/s^ were "in a condition
of gieat confusion."' Althoii^'h a niniibci- of papci's dcalin<^ with tluf
New World .-pecics have appeared since liiat time, the systemat ic status
and particnlaily the distriitut ion of the American forms are still far
from clear.

The discovery of the new halfbeak, described below, biin^^s nj) the*
(juesiiou of the <reneric validity of //i/por/ttt//i/>/i>/s and has prompted
u ciitical study of hifisilirtis'/s and un'ifaxciaiiis, the ^feiiotypes, I'espec-
tively, of //rri4iromp/iu.s Cuvier and IIi/p(>rhain]>JiUi< (lill. This study
has j)roved to be most pr"odiict ive. for a number of t rencliant and easily
observed cliaraiters, lieretofnrr appaicntlv o\erlooked, weic found.
The presence or ab-ejice of scales on the upjiei- jaw also was noted by
Smith ( ll).'i:5, p. VM)). In |)reparin;j; table 1, in which the «^enotype.s
of IlrmiramphuH and Uyporhamplivx are compared, I examined 135
s|)ecimens of ^inifayciatns and 05 specimens of lir<i-'<irnnxis in the col-
lections of the W S. Xational .MuH-um. These specimens represent

' SiK-Urd llrmirhnmphun liy tlirni niiil liy i\ lioHt of otluT nuttHirH. Tlio rtrlk'iniil HpoMltifc
bjr Cuvlor (1817. p. \HC) Ih Himi Unrnphun.

r.n.'ir.ao— 4.'»

IK.--,

186 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

material from the known American range of both species : ■mufasciatus,
from Cape Cod to Uruguay in the Atlantic and from San Diego -
to Peru in the Pacific ; and hrasiliensis, from New York to Brazil.

Gill (1859, p. 131) based Hyporhamphus principally on the tricuspid
teeth (whence the name of the type species, H. trieuspidatus, a syno-
nym of unifasciatus) , but he later (1863) found, and Poey (1860,
p. 298) previously had noted, that Eemiramphus likewise has tricus-
pid teeth. Poey's and Gill's observations on the nature of the teeth
were correct, and hence I do not agree with Weed (1933, pp. 47, 57)
and others who stated that the teeth are simple in Hemiramphus. As
Smith (1933) has shown, and as I have also observed, the form and ar-
rangement of the teeth vary with age and with different species. The
jaws of a single individual may have unicuspid, bicuspid, and tri-
cuspid teeth, and, in at least one American species, Hyporhamphus
rosae (Jordan and Gilbert), only the largest individuals appear to
have tricuspid teeth — hence the frequent statement that H. rosae has
only unicuspid teeth.

The fundamental characters distinguishing the American species of
Heiniramphus and Hyporhamphus^ such as the presence or absence
of scales on the upper jaw, the presence or absence of a bony rim along
the side of the nasal fossa, and the arrangement of the sensory canal
and pores on the preorbital (fig. 9), may be features that will separate
world half beaks of this type. This is suggested to me by Smith (1933,
pp. 130-131), who made a primary division in his key on the basis of
a naked versus a scaled upper jaw, and b}^ the very few Old World
half beaks I have examined. In Euleptorhamphus Gill, however, the
upper jaw is scaled as in Hyporha7nj)hus, whereas the rim and the
form of the nasal fossa and the sensory canal of the preorbital are
essentially as in Hemiramphus.

The pattern of the scales on the uj^per jaw, the shape of the pre-
orbital, and the arrangement of the teeth maj^ be found to have generic
or only specific value. The solution of these problems will necessitate
a comprehensive review of the halfbeaks of the world.

The form of the sensory canal and the pore on the preorbital are
usually visible in Hyporhamphus^ but the overlying scales and skin
must be dissected from this bone in Heniiramphus before the canal
and pores can be clearly seen. The two pores shown near the upper
end of the posterior margin of the preopercle in Hyporhamphus (fig.
9, A) are apparently absent in Hemiramphus^ but this character was
checked only on a comparatively few individuals of each genus.

In table 1 I have abandoned the "key" characters — air bladder cel-
lular or simple, sides of body vertical or convex, position and shape of

2 In material from San Diego, Calif. (Stanford Nat. Hist. Mua. No. 9912) I found one
specimen of this species, Which, to my linowledge, represents a northward extension of
known range on the Paciflc coast.

A NEW IlLMlHAMrillD IISIL — MlLLEIi

187

dorsal lin, ami position of pel vies — used by many writers to separate
Ueininunjihu.s from Ilf/porfunnphiu^. The nature of the air bladder
is diflicult to discern but may be of considerable phylogenetic impor-
tance; the form of the sides of the body is an untrustworthy character
because it is frequently rendered impractical by preservation; the posi-
tion and shape of the dorsjil lin is not so distinctive a feature as is the
ditVerence in the basal lenjiths of the dorsal anil anal fins; and the
position of the pelvic fins is useful largely for specific or subspecific
separations.

Table 1. — Diaguustic differences bettceen Hyporhamphus unifaBciatue and Hemi-

ramphus brasiliensis '

Character

unifasciaiui

brasiUensis

Upper Jaw

Sc-altti

Naked.

MaTKin of nasal

fossa

Surmounted by a prominent bony

Lacking a bony rim in this position.

(see flg. 9).

rim along posterolateral border.

Sensory canal

pre-

Unbranched; with an exposed pore on

Branched; witli a jiore at end of pos

orbital (see flg. 9)

side and another pore at terminus

terior branch (wliich terminates in a

of canal near anterior margin of

bony rid^'c near front of orbit) and a

nasal fofxsa.

pore at end of anterior branch near
antcroventral nisiri^'in of nasal fossa.

I)'>r««l fiT!

Over or nearly over origin of anal, its
base and that of anal equal or sub-
equal.

In advance of anal oripin, its base 1.5
to 2.1 times that of anal fln (1.3 or 1.4
in young).

Caudal Hu

Moderately forked, the distance be-
tween caudal ba.se and shortest

Deeply forfceii, the dLsfancc between

caudal base and shortest caudal rays

caudal rays 7.-1 to 9.0 in standard

12.5 to IC.3 in standard length.

length.

Sh:'.]!*.- cjf iiasiil

fossa in

Broad, and little dcpressc*!. Its great-

Narrow and greatly depresised, Its

a'JuIt.

est inner diameter more tlian one-
half that of orbit.

greatest dianietor oni-fourth to one-
third that of orbit.

> Characters of Ilvjwrhamphvi confirmed on the type specimen fll.P.N.At. No. 8407) of the genus, H
trtciupid(Uus(,=-unifafdiiliu), and of Jhniiramphun on "lopotyie.s" (sjieeimens from Jamaica, U.S.N.M.
No. 30077) of //. briuUientiM.

The new species described below is the lirst to be definitely recorded
from fresh vater in the New \\'oild. It apjieais to be restricted to
a fluviatile habitat, for a numbei- of collections of halfl)eaks along the
west coast of Mexico in the n-gioii where the new species was dis-
cov«'red ct^ntain no sj)ecies identical with it.

I name this distinctive fish />'/// /.v,gcii it i\e of jxifrr (father), because
my fathej-, lljilj)h (i. Miller, collected the It types hihI only known
spCH.imeii.s.

HVrOIUIAMI'HUS rATKIS, new iprciri
I'LATK 11

Tyin'8. — The holotype (U.S.N.M. N'o. 129950) is a mature aihilt
(presumably a female, sec below), lis iiim. in standard length, and
was collected on May 4, 1912, by IJalpii (J. Miller in Rfo del Fuerte,
one-half mile above the town (»f Kl Fuerte, which is about 2(1 miles
northeast of San lilas, Sinola, Mexico. Tlie 1.'5 paratypes (U.S.N.M.

188 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m

No. 129957), 107 to 130 mm, long, wore collected with the holotype.
One fish in the lot, a specimen 118 mm. in standard length, is tlie only
individual of the series that has distinctly larger pectoral and pelvic
fins. On examination it was found to be a ripe male. One of the
others, a specimen 109 nun. long with short pectorals and pelvics, was
found to contain eggs in various stages of development, some of them
apparently fully mature. The remainder are ])resumably all females.

Diagnosis. — A Hyporhaiiypku^ with pelvic fins about equidistant
between caudal base and gill opening, 21 to 24 gill rakers on lower
limb of first arch, with a relatively long mandible (3.6 to 4.2 in
standard length), without scales on dorsal or anal fins, and without the
fleshy tip of the mandible red.

Desonption. — Bodj^ rather slender, its depth 8.0 to 9.6 in standai'd
length, little compressed, the sides rounded; w-idth of body in depth
1.05 to 1.4; head 4.5 to 5.0 in standard length; mandible (measured
from tip of upper jaw to end of bony tip) 3.6 to 4.2 in standard
length and 0.7 to 0.9 in head length (broken in one specimen) ; snout

2.8 to 2,9 in head ; orbit 4.0 to 4.3 in head, 1.35 to 1.45 in snout, and 1.65
to 1.85 in postorbital; interorbital 3.8 to 4.1 in head and 1.55 to 1.7
in postorbital; length of preorbital 1.5 to 1.65 in orbit; depth of pre-
orbital 1.5 to 1,75 in orbit; width of nasal fossa 1.85 to 2.15 in orbit;
base of anal fin 1.01 to 1.08 in base of dorsal fin; pectoral short, 8.4 to
9.35 in standard length in females (7.9 in the male) and 1.75 to 1.95
in head (1.65 in male) ; pelvic 2.7 to 3.0 in head in females (2.25 in
male) ; midcaudal rays (measured from midbase of caudal fin to tip
of shortest middle ray or rays) 8.4 to 9.3 in standard length, 1.7 to

1.9 in head, and 2.1 to 2.4 times the length of the orbit.

The fin rays vary in number as follows: Dorsal 13 to 15, usually 14;
anal 15 or 16. usually 16; pectorals 10-10, 10-11, or 11-11, ahnost al-
ways 10-10; pelvics always 6-6. I depart from my usual method in
counting the rays of the dorsal and anal fins and regard every element
as a separate ray, because this procedure has been followed by virtually
all students of this group of fishes. Without exception the first two
rays of the doi*sal fin are unbranched, and the first two rays of the
anal fin are also simple except in two specimens in which the first
three rays are unbranched.

The gill rakers on the lower limb of the first gill arch (counted on
both sides) vary from 21 to 24.

The lateral series scales (counted from upper angle of gill opening
to caudal base) number about 53 to 59, usually 55 to 57; an accurate
count is difficult to obtain because the scales are largely missing from
the sides.

The pelvic fins lie about equidistant between the base of the caudal
fin and the gill opening, varying between the pectoral base and the
middle of the opercle. The dorsal fin varies in position from equi-

A NEW HEMIRAMPPIID FISH — MILLER

189

distant bctwi-en caudal base and pelvic insertions ((. niuch nearer
{lelvic insertions tlian caudal base.

The tei'th of the holotype are unicuspid, bicuspid, and tricuspid and
are arranged in about three to seven irre<!;ular rows in the upper jaw
and two to five rows in the lower jaAv. Tricuspid teeth are present
only posteriorly in each jaw and virtually all the anterior teeth (from
about the middle of each jaw forward) are unicuspid. In the region
where unicuspid and tricuspid teeth intergrade, occasional bicuspid
teeth occur. The tooth rows are conspicuously broader medially on
each side of the upper jaw tlian they are at either end, and teeth are

Figure 9. — Sketch of head regions of Ilyporhamphus and Ifemiramphus to illustrate
certain diagnostic differences (see tabic 1): A, I/yporhamphui unifasciatus, 183 mm. in
standard length, from Key West, Florida (U.S.N.M. No. 34599); B, Hemiramphui
brasilieitsis, \h2 mm. long, from Key West, Florida (U.S.N.M. No. 38684). Drawn by
•Mrs. A. M. Awl, U. S. National Museum.

absent at the tips of both upper and lower jaws. In the lower jaw the
rows of teeth are of nearly uniform width but are somewhat broader
close to the proximal end on each side and theji become narrow gradu-
ally forward and abruptly behind this region. In the largest ])ara-
ty|>e (KiO mm. in standard length) there are more ti'icuspid teeth than
in the holotype (118 mm. long), which agrees with my observation in
//. roiiae that tricuspid teeth api)ear with increasing size of the indi-
\idual (this was also not<'d in other American //rnnramj>hus and
Ih/lforJicinijihuR). Otherwise the teeth of the paratypes have es.sen-
tially the same form and arrangement as in the holotype.

The triangular up{)er jaw is rather bluntly pointed at the apex and
l>road<'r at the base than it is long, \\nien the mouth is closed most of
the outer teeth of the lower jaw are exposed. The scales of the upper
jaw are irregularly arranged, the transverse rows munbering five or
six arross the base, tlion al»oiit four, whereas from about the middh' to
tlio tip of the jaw thoy are bi^erial — with a single scale on each side of

190 PROCEEDINGS OF THE NATIONAL MUSEUM vol. o«

the slight median ridge. Although the scales cross this low ridge pos-
teriorly they do not usually do so anteriorly.

No scales were observed at the tip of the upper jaw, but these may
have dropped off. The sides of the head, including the region of the
mandible below the jaws, are covered with deciduous scales.

The margins of the prolonged mandible or "beak" are nearly par-
allel throughout, diverging little until the posterior end is reached.
The nasal flap is small.

The dorsal and anal fins are low, highest anteriorly, with rays 3 to
5 longest ; these rays in the anal fin are almost three times as long as
the last ray, whereas in the dorsal fin the anterior rays are only about
twice as long as the last ray, which is slightly prolonged and falls
some distance short of reaching the bases of the procurrent caudal
rays. The asymmetrical caudal fin is very weakly forked, less so
tlian in any other American species I have seen except H. rosae. As
in many halfbeaks, the lower caudal lobe is longer.

The air or swim bladder as noted in the single male is simple, with-
out any cellular structure.

Coloration. — The general coloration was noted in the field by the
collector. When taken from the water the body of the new species
was intense blue and green varying in brilliance according to the re-
flection of light from the surface, the blue and green grading into
each other. The fins or the belly are believed to have been yellow or
orange. No bright color was seen anywhere on the beak. This ob-
servation is important, for most, if not all, of the American halfbeaks
have the fleshy tip of the mandible red. According to Herre (1944,
p. 9) the Philippine species of Hemiramphus (including Hyporham-
■pJius) have this tip red, green, or greenish white, depending upon the
species. I therefore interpret the lack of red color on this structure
in patris as a character of specific value.

The color of the preserved specimens (in alcohol) is mostly light
silvery to pale brownish.^ The back, above the lateral band on each
side, is marked with brownish punctulations. which are usually more
concentrated on the posterior borders of the scales. Along the middle
of the back are three narrow longitudinal rows of dark pigment,
broader near the occiput and particularly over the caudal })eduncle ;
the outer rows are more or less continuous past the base of the dorsal
but the middle row is disrupted in this region into a series of U- oi-
V-shaped markings between the bases of the rays. The base of the
anal fin is marked similarly to that of the dorsal base, but the longi-
tudinal rows of pigment are far less conspicuous. On each side of
the body is a dark band, probably silvery in life, which is very narrow
anteriorly and broadest between dorsal and anal fins. The upper
surface of the anterior part of the head, including the upper jaw, and
of the mandible is black; the lower surface of the mandible is finely

A .\K\V HEMIRAMPHII) Fli:H — MILLEK 191

pi<;iiii'iit<'il with black cliroiiuitophorcs ftulinj^ po.steiiorly bo that
both cliiii and tluuat art" hugely coh>rlc>;s. The tips of tlie caudal rays
and those of iho Ioii«i:er dorsal rays arc marked with fine black jiunclu-
lations; tiie other fins are mostly pale. Alon<i: the underside of tho
caudal peduncle are three rather irregular lontritudinal rows of dark
l>iLrinent. The silvery peritoneuni is ovei-lain by foppi'iy Ijiowii and
by fine, black punctulations.

Habitat and associates. — Rfo del Fuerte, near El Fuerte, Sinaloa,
is a deep river with sand and mud bottom and abrupt rocky banks.
On -May 4, r.»42, when the types were collected, the current was fairly
swift, and hauls with a 2o-foot bag seine were made in water generally
1 to .") feet deej) but more than G feet in places. At noon the air was
37*^ C. and the water 32 C No vegetation was seen, and the shore
was sandy, with trees along the bank. Collecting was confined largely
to the backwaters. The point wlierc the fish were secured is fully
KK) miles upstream from the Pacific.

In the large colle«'tion made here, the following fishes, tentatively
identified, wer«' also seined: A species of cyprinid fish of the genus
Gila; two specimens of a catfish of the gemis Ictaluius; cyprinodont
fi.shes of several genera including MoUienisia sphenops; six mullets,
Af/onostonuis montieola: a large niunber of the fresh-water atherine
Mclaniris cryataUinus ; and two gobies. Ait'aou,s (or Clumopkonis)
transandeanus and Gohiomonis 7>wntlatv-s\ Most of these species are
confined to fresh water.

Ranfi< . — The new s[)ecies was collected only in the Kio del Fuerte.
Ralph G. Miller -aw halfbeaks in the Rfo Culiacan at Culiacan, Sina-
loa. Mexico, alxiiit l.">0 miles south of El Fuerte and about 40 miles
ui)Stream fr(»m tin* Pacific, but the identity of this species is unknown.

Relationships. — Tlyporhamphus patris appears to be the southern
repi-esentative of //. ros,ic (.Jordan and Gilbert) (1880). which is
known frojii San Pedro, Calif., south to the tip of Paja California,
then up the west side of the Gulf of California and southward along
(lie maiidand of Mexico to Guaymas. Sonora (Evermann and Jenkins,
1801, p. 13.'); record confirmed by examination of the five specimens
from (iuaymas in the Stanford Natural History M\iseum, No. 437).
Ki'f) del Fiier-te, tlie liabital of jxitris, is about 170 miles soiilli of
tiuaynias.

'ilie two s|tecie^ agree in most measuremenl> and coimts and in the
following important charact<'rs: (1) P(»st«'rior position of the pelvic
fins; {'!) gill rakers: 21 to 2;') on the lower limb of the first gill arch
in my counts for rosne^ 21 to 21 for patris; {W) long mandible, which
api)ears to Ix' slightly longer in roKar^ but a series of comparable sizes
would probably eliminate this difference; (4) no scales on i\\*.' dorsal
or anal fins; (.'>) dentition. The tw(> species differ as slutwn in the
(oiiiparison presente<j in table 2. Soine oi- all of these difrerenc<»s

192

PROCEEDINGS OF THE NATIONAL MUSEUM

may vanish wlien larger series of both species from more localities
are available, but it seems best at this time to regard them as distinct
species.

Table 2. — Comparison of Hyporhamphus patris and H. rosae

Character

Tosaf

patria

Color of mandible

Dark red

Black.

Fleshy tip of mandible

Red

Not red.

Scales in lateral series

58-65

54-59.

Bony interorbital into length of

head.
Diameter of orbit Into head

3.4-3.8.

3.4-1.0

3.8-4.1.
4.0-4.3.

length.
Size -

Smaller; largest specimen 107 mm.
in standard length, usually
much less than 100 mm. long.

Larger; largest specimen 130 mm.

long and smallest 107 mm.

The posterior position of the pelvic fins and the few gill rakers
readily separate patns from EyporhampTius unifasciatus (Ranzani),
H. roherti (Valenciennes) {=hildebrandi Jordan and Evermann),
H. snyderi Meek and Hildebrand, and JI. gilli Meek and Hildebrand,
the other species reported from Middle America (Meek and Hilde-
brand, 1923, pp. 236-241, pis. 16-17) .

Acknowledgments. — Material of Eyporham/phus rosae was bor-
rowed from the Stanford Natural History Museum, through the kind-
ness of Miss Margaret Storey, and from the Chicago Natural History
Museum, through the courtesy of K. P. Schmidt aiid Mrs. Marion
Grey. I am grateful for this cooperation.

LITERATURE CITED

C'l'vu B. Gkoiii.k8 L. t". F. D.

1.S17. Li' ri'pne aiiiiniil distribue d'apros son «irg;misiiti<Mi, \ynur scivir dc base
ft lanatitmio coniparie, vol. 2. xviii + 532 pp., lllus. Pails.
K\i:i{MAN.N. Bahton W.\rbkn, and Jknkins, Ouvkr Pkebi.es.

ISStl. HeiMirt upon a (•«>llei-tion of flsh«'.s made at Guayiuas, Sonora, Mexico,
with di'scriptioiis of new species. Pnic. U. S. Nat. Mas., vol. 14, pp.
121-lt;.'^. 2 pis.
GiM, Tii>:oi)oi;k Nicholas.

\H'>9. Deseription of HiipoihitiiHihux. a new ^'enns of fislnss allied to Ilfttii-

rhdniphu-s Cuv. Proc. Acad. Nat. Sci. Phil:idelphia, lSr>[), p. 131.
18Js{. Note on the genera of Ileinirhaniphiri.tc Pi(.c. Acad. N.il. Sd. Pliila-
delphia, ]8«3, pp. 272-273.
IIi-RKK. Albert W. C. T.

I!t44. A review of (In- lialfbeaks or Heniiramphidae of the Philippines and
adjacent waters. Stanford Univ. Pnltl.. univ. ser.. binl. .sci., v«>l. 9,
No. 2. pp. .S!^-8»5, 1 map.
Hrmis. CaR[. Lkavitt, and Millkr. RoitKitr Rush.

IIMI. DoroKOiita smUhi, the fiisl kmnvn gizzard shad from the Pacific drain-
age of Middle America. Copeia, 11)41, No. 4. pi). 232-238, 1 fig.
JoKDAX. Damd Stark, and GiiJirarr, Charles Henky.

188<J. Description of a new species ot ncmirhamphwi (Hcmirhamphus rosae),
from the coast of California. Proc. U. S. N.it. Mus., vol. 3, pp.
33.v:«6.
Meek, Seiu Eugknk. and G<)ss. Damd Kore.

IXSii. A revi«-w of the American .spwies of (he genus n< niii lnunplitis. Proc.
Acad. Nat. Sci. IMiiladelphia, 1S84. pp. 221-220.
Mekk, Seth Eugene, and IIili)ebr.\m), Samuel Fheueruk.

ir>23. The irfurine fishes of Panama : Part 1. Publ. Field Mus. Nat. Hist. No.
21.";. zo.il. .ser.. vf.l. 1.'. xi + '■^^'^^ PP . 24 pis.

PoKV. FkI.II'E.

ista). Meniorias soImc I.i liistoria do i.i isi:i i\t ('\ih:i. vol. 2. f.-isc. 'J., pp. '.t7-33»>.
S.Miin. .1. L. B.

P»:'..''.. Tin' South African sixcies of the genus II< mil luiuiiiliiis <'uv. Trans.
Roy. S(M-. South Africa, vol. 21, pt. 2. pp. 12!» l.'.d. 1 fig., 3 pis.
Wk>.i>. Aii-rkd <'i.evela.m>.

VXVA. Notes on fishes of the f.-millv I lemirhaiiii>hid.ic. Piihl. I'ifid Mas. Nat.
Ilisl., zr.ol. s«'r.. vol. •_'2. itii. II <■><;.

103

u t •ovtmiiidiT ruiNTiNo orrict r it«i

U S NATIONAL MUSEUM

PROCEEDINGS VOL 9C PLATE 11

u C_

;->

PROCEEDINGS 0.^ THE UNITED STATES NATIONAL MUSEUM

issued I^^vA-sjIJ^b h '^«

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM

Vol. % Waihington: 1946 No. 3196

NOTES Oy. RECENTLY MOUNTED REPTILE FOSSIL SKELE-
TONS 'IN THE UNITED STATES NATIONAL MUSEUM

By Charles W. Gilmore*

Skeleton of a nearly complete Eocene crocodile and a partial skele-
ton of Corythosaurus , a crested dinosaur of the Upper Cretaceous, have
recently been added to the exhibition collection of fossil vertebrates in
the United States National Museum. In the brief notes presented
here, attention is called to some of the more interesting anatomical
features of these specimens, and measurements arc given of all the
more important bones in order to make these data available to stu-
dents of the fossil Reptilia. The crocodile specimen appears to be
unique in being the first complete skeleton to be mounted for exhibi-
tion in this country.

SKELETON OF CROCODILUS CLAVIS COPE

Platks 12-1 f)

Among the specimens collected by the 1930 Smithsonian Paleon-
tologieal Expedition to the Bridger Basin in southwestern Wyoming
was an unusually complete skeleton of a crocodile, U.S.N.M. No.
127l'.». It was found by George B. Pearce in the badlands between
lievett and Little Dry Creeks in horizon B of the liridger fornuition.
The skeleton as it lay in the ground was only i)artially articulated, but
inasmuch as it is an isolated specimen the few bones found detached
and sf-attered can surely l)e regarded as peitaining to a single individ-
ual. The skeletal parts preserved are as follows: Skull, lower jaws,
23 presacral vertebrae, 2 sacral vertebrae, 33 caudal vcrtcbiac 17
chevrons, complete pectoral and pelvic girdles, If) eer\ ical ribs. 17

•Mr. Ollmorp died on ."^cptcmbor 27, 194'>.— Ki<.

6W7.14— 16 1 Jtr)

196 PROCEEDINGS OF THE NATIONAL MUSEUM ^^^ gg

thoracic ribs whole or in part, right humerus, both radii, right ulnare,
both radiaha, parts of 2 metacarpals, both femoro , both tibiae, both
fibulae, both tarsi, all metatarsals, 1 1 phalangials, 2 inguals, and num-
erous dermal scutes.

On account of the rarity of good crocodilian skeL'tons in paleonto-
logical collections, it seemed desirable to articulate this specimen for
public exhibition. After several months' work this w;as accomplished
by Norman H. Boss, chief preparator, who is to be higtdy commended
on the excellent results achieved.

The individual bones are thoroughly mineralized an d practically
free from postmortem distortion. A few elements, ho'>vever, either
through injuries or disease are abnormally deformed. The right
scapula is a most interesting example of a badly healed fra cture. In
life the scapula was cleanly broken through the narrowesu" part of
the blade. This upper portion dropped down on the inside of the
proximal half for fully an mch below the point of fracture, and inhere
the two parts were securely knitted together by extraneous bvOny
growth. Although this fracture must have been exceedingly painijil
at the time, after healing the limb undoubtedly continued to function.

The second metatarsal of the left hind foot exhibits a pathologic
condition that has enlarged the shaft of the bone to nearly twice its
normal size. This lesion can probably be attributed to an injury.
Other lesions are found on the left coracoid, anterior thoracic ribs,
caudal vertebrae, chevrons, and skull. That this animal was a pug-
nacious individual and often engaged in combat, probably with
others of its kind, is clearly indicated by the considerable number
of healed wounds.

There are 23 presacral vertebrae preserved, but a restored lumbar
was introduced between the first and second, the only point showing
evidence of a break in the series, in order to make the presacrals corre-
spond in total number to the vertebral formula of Crocodilus americanus
as determined by Mook.^ This introduction makes four lumbar
vertebrae, whereas Mook recognizes only three in C. americanus;
but as a mounted skeleton of this species (U.S.N.M. No. 14874) has
four, it seems reasonable to assume that a similar variation may
occur in the fossil species.

The caudal series consists of 37 vertebrae of which 33 are original
bones. Four vertebrae at the tip of the tail are fully restored.
According to authorities the total number of caudal vertebrae is
subject to considerable variation among living individuals of the same
species.

The skeleton, over all from tip to tip, measured a^long the curves
of the spmal column has a length of about 9 feet 10 inches.

Pending the publication of the monographic study of the Croco-

1 Mook, C. C, Bull. Amor. RTus. Nat. Ffist., vol. 44, pp. 70-78, 1921.

REPTILE FOSSIL SKELETONS — GILMORB 197

(lilia by Dr. C. C. Mook, this specimen is provisionally identified as
portaiuin*]: to the species Croc(x!ih!.<t claris Cope. ITay * recoo7iizes 10
species of (\ocodibt'i from the Bridger formation alone, but a thorough
revision would doubtless greatly reduce this list.

The mounted skeleton has been given a defiant attitude (see pi. 13)
with the jaws agape in order better to display the mouthful of teeth.
The pose and style of mount adopted were largely determined by the
character of the skeleton. Because of the extreme hardness and
brittleness of the fossilized bone, it was found impractical to drill
the bones for secm'hig them to metal supports, and so Mr. Boss
worked out a scheme of half relief and half free mount that overcame
this difficulty an<l gives pleasing results, as is clearly shown in plates
12 and lo.

The skidl and lower jaws are unusualty complete and only slightly
distorted by crushing.

In the upper and lower mandibles there are alveoli for 76 teeth,
premaxillaries 10, maxillaries 30, and dentaries 36. Of this dental
series 29 teeth were found in place; enumerated from the front these
were distributed as follows: Second of the right premaxillary, a germ
tooth, and fourth of the left premaxillary; hi the right maxillary the
fifth, nintli, tenth, and eleventh, none in the left maxillary; in the
left dentaiy the first, fourth, fifth, sixth, seventh, (germ tooth), eighth
(gemi tooth), ninth (germ tooth), tenth, and eleventh; in the right
ramus the first, second, thii'd, fourth, fifth, sixth, seventh, eighth
(germ tooth), twelfth, thirteenth, fifteenth, sixteenth, seventeenth,
and eighteenth (the last four being germ teeth).

The other teeth, some 39 in all, were found loose in the matrix
surrounding the skull, and these have been arldtrarily inserted in the
jaws. Thus of the 76 teeth forming the complete dental series,
original tooth crowns of 69 are preserved.

In Crocodilus, according to Mook,^ all the living species of the genus
have 17 to 19 toclh in the upper sei-ios and oidy 15 in the lower, or a
maximum total of 68 teeth in the mouth as contrasted with 76 in the
extinct species. Of 4 other skulls fr<jiii the Bridger in the National
Museum's collections none shows less than 36 in the lower dental
series, and it is evident that the greater number of teelh constitutes an
important feature for distingui.shing the extinct Kocene forms fr(»ni
\]\(' extant member.^ of the genus. It also raises the question of the
propriety of referring these Eocene crocodiles to the genus (Vorodllus.
That , however, is outside the scope of the present paper, and no doubt
will be fully considered by Mook in (he ronrse of his monogrHi)lii(;
study of the order.

> l\ny, f). I'., UiMlogrftphy onil catolfiRiio of ffissll vcrU'l)rnt<'s. Cnni'-rli' Itist. \\'ir liliirti.n V\ih]. sr*),
pp. .■il2-.M3, 1928.
• .Vlwjk. C. C. Hull. Amcr. Mus. Nat. ITIsl., vol. ii. p. I.M. 1021.

198

PROCEEDINGS OF THE NATIONAL MUSEUM

The excellent preservation of the present skeleton offers an unusual
opportunity for recording the important measurements of a single
individual of an Eocene crocodile. In the tables that follow the prin-
cipal measurements of the skeletal parts are given, following Mook's

Table 1. — Comparative measurements {in mm.) of skull, pelvic, and limb bones in

two species of Crocodilus

Measurements

Length of skull, tip of snout to supraoccipital

Length of skull, tip of snout to ends of quadrates

Breadth of skull, cranial table

Breadth of skull, across fifth maxillary tooth

Length of mandible

Lp,ngth of scapula, total -.

A nteroposterior diameter of superior border

Anteroposterior diameter of inferior border

Maximum thickness of distal end

Length of coracoid, total ---

Anteroposterior diameter, superior surface

Anteroposterior diameter, inferior surface

Length of humerus, total ..-

Breadth of proximal end

Breadth of distal end

Circumference of shaft -

Index of circumference over length

Length of radius, totaL -..

Maximum diameter, proximal end

Maximum diameter, distal end

Length of ilium, total oblique - --- --

Length of ilium, total anteroposterior

Distance across both ischiadic processes

Maximum length of ischium, oblique

Anteroposterior diameter, proximal end .-

Maximum diameter, distal end

Length of pubis, total _-_ _.

Maximum diameter, proximal end

Minimum diameter, proximal end

Breadth distal end

Length of femur, total

Breadth , proximal end

Breadth, distal end _

Circumference of shaft

Distance from center of fourth trochanter to proximal end
Distance from center of fourth trochanter to distal end

T J i- (Center of fourth trochanter to proximal end}

Index ratio; L-

l Center of fourth trochanter to distal end J

Length of tibia, total

Maximum diameter, proximal end- __

Maximum diameter, distal end_

Length of fibula, total _ -

Maximum diameter, proximal end

Maximum diameter, distal end

Circumference of shaf t_ _._

- , .. I Circumference of shaft]

Index ratio; >._

I Total length 1

C. clavis
(U.S.N.M.
No. 12719)

418
481
110
126
535
136

77.3

75.4

27
101

60.4

68.5
180

50.5

61
338
107

26.1
125.6
105
145
1.39

93.5

17.4

73.5
223

51.5

81
142

0.570

159

43.1

38
156

C. americanus

(A.M.N.H.

No. 7139)

1 Possibly an error, in view of greater proportions of most of the other measurements.

REPTILE rOSSIL SKELETONS — GILMORE

199

system * used in doscribing: the osteology of the extant Orocodilvs
aniericanvs.

For convenience in reference I have included in parallel columns the
measurements by Mook of a considerably larger (\ amcricanvs.
In view of the antiquity of the extinct skeleton these measurements
show a remarkable similarit)" of proportions between the living and
extinct forms.

Tahle 2. — Comparative measurements {in mm.) of cervical, dorsal, luvihar, and

sacral vertebrae

Length of
centrum '

Breadth of

centrum,

anterior end

Spread of

preiygapo-

pliyses

Spread of
postzyga-
pophyses

Spread of

diapophy-

ses

Total
height

Vertebrae

•z

g
d

s
z

K
Z

d
Z

2
d

w
z

z
z

a
z

i
z

o
d

a
z

z
z

d
Z

w
z

Cervical 2

4g.6

32.5

31.9

32.5

34.9

36.0

39.9

04.5

65.6

25.5

30.7

33.9

32.8

36.2
33.6
44
30

46
40
43
44
44
52
60
54
50
53
50
48
49
49
60
60
61
57
55
65
60
56
65
60
60

32.2

48.5

00.5

GO. 6

68.5

81.5

82.6

26
41

52.5

62.5

59.5

58,5

59.5

55.4

33.8
28
42.6

63.5

62.6

53.5

51.7

73.3

84
101
105
127
149
159
150
160
100
100
UK)
144
130

35.5

64.5

CO
107
150
183
200
213
218
230
242
276
277
250

79
88
06

105

122.9
121.6

118
95
90
88.5
85
80
81
79
80

88
79.8

127

140

156

166

167

175

Dorsal 1 '

180

185

180

181

100

150

148

145

140

136
139

r.iimbar 1

134

133

39
38
42
43.7

120

156
110

235
218
230
202

132

Sacral 1

132
135

142

' Th*> mntKurriiicnts of the vertebrnl centra of Ihn presacral st-rlr* of the fo.ssil specimen do not Include
the ball, and so In using these mea.surenicnts allowance must bo made for this omission. This discrepancy
was dui) to the fact that this prjrtlnn of the backbone was articulated in nxed position before It was decided
to tiko thi? scries of mcfi.<iuremonts.

•Ibid., pp. 71-100.

200

PROCEEDINGS OF THE NATIONAL MUSEUM

Table 3. — Comparative measurements {in mm.) of caudal vertebrae

Length of

Breadth of

Height of

Spread of

Height

Spread of

centrum, an-

prezygapo-

postzygapo-

transverse

terior end

physes

processes

o»

r-"

Caudals

d
A

o
A

d
•A

d
'A

d
"A

o
A,

^^,

:§■

■<

55.9

34.3

58.5

83.9

130

122

196

•2

51.7

59.5

134

132

195

29.4

45.5

31.5

135

126

186

135

116

175

27.4

44.6

28.6

41.5

27.8

46.5

38.5

136

166

55.7

26.9

38.5

75.5

130

112

157

53.7

40.5

132.5

141

53.2

29.5

118

130.5

53.5

34.5

36.6

20.9

67.5

103

120

24.4

35.5

24.1

36.5

68.8

101.5

118

33.5

30.5

21.5

100

120

54.3

22.5

107

22.5

23.5

104

88.5

53.4

21.7

102

44.5

49.8

47.5

44.8

43.6

42.5

40.5

30.5

74.5

70.5

60.5

21.5

19.5

14.5

13.5

6.3

29.5

28.5

20.4

22.5

13.5

10.5

7.5

20.5

21
21
18
18

16.6
16
16

15.2
15
14

13.1
7.5

26
26
24
25
23
22
20
20
18
18
17
15
14
12
11
10
9
8
11

15
14

12.5
11.2
10.5

6.9

14.5

12.5

10.5
9.5
8

6.5
5.5
3.5
2.5
3

•4.5

10
7

4.5

3:9

24.5

21.5

11.5

7.6

3.5

3.6

2.5

1
.5

3
12.5

65.5
65

46
45
38.9

102
106
105
118
109
88
96
87
82
65
50
50
42
35

27
22.5

13.5

1 Estimates

NOTE ON DISEASED CROCODILE VERTEBRAE

Plate 16

A second crocodile specimen, U.S.N.M. No. 12990, from the
Bridger, Eocene, consisting of the greater portion of a skeleton,
shows a pathologic condition of two dorsal vertebrae that is almost
identical with the lesion described by Moodie ^ on the caudal verte-
brae of a sauropodoiis dinosaur from the Jurassic.

It is a spongy growth that surrounds the intervertebral articular
surfaces extending well outward on the sides of both centra (see

5 Moodie, Roy L., Amer. Journ. Sei., vol. 41, pp. 530-531. 19IP.

REPTILE FOSSIL SKELETONS — GILMORK 201

pi. IG). It entirely encircles the centra and has involved two-
thirds of the two bones. All evidence of separate structure is prac-
tically obliterated. The growth is cjuite symmetrical on the two
sides.

Moodie in the case of the lesion on tiie dinosaur caudals says:
"The enlargement is somewhat suggestive of the lesion of chronic
osteomyelitis. It may be a callous growth due possibly to a fracture
of the caudal vertebrae; or it may be a bone tumor." Its true
nature is, of course, uncertain, but mention is made of this specimen
here in order to call it to the attention of students of modern pa-
thology who may be interested in the study of the nature and origin
of disease.

RufTer' has reported typical lesions indicating spondylitis dejor-
mans in the vertebrae of the Miocene crocodile Tomistoma dowsoni,
from Egypt. In this specunen the extraneous osseous tissue, obvi-
ously pathologic, binds the vertebrae together. The new bone,
however, is thicker on one side than on the other, and Moodie ^
observes that "in the crocodile as in man the disease is more marked
on one side." The symmetrical nature of the lesion in the National
Museum crocodile shown in plate 16 would therefore rule out spon-
dylitis dejormans as being responsible for the development of this
altuormality.

ON A SPECIMEN OF CORYTHOSAURUS

Plates 17-19

A partial skeleton of Corythosaurus recently added to the exhibition
scries in the United States National Museum consists of the complete
articulated tail, pelvis, hind limbs, and feet, with several small patches
of skin impressions and o-ssified tendons. This specimen is mounted
in rehef so as to display the right side; the sandstone blocks containing
the bones have been assembled in the same relationships the}' occu-
pied in the ground. The preparation ant! mounting were done by
Norman H. Boss, and it was due to liis skillful manipulation that so
much of the epidermal impressions were preserved.

The specimen (L'.S.X.M. No. 1549:^) was acquired by purchase
from the Royal Paleontological Museum of the University of Toronto
for use in connection with the Smithsonian exhibit at the Texas
Centennial Exposition at Dallas in \\)'M . It was collected l)y Levi
Sternberg from the Belly River formation, rpjjer Cretaceous, 2 miles
south of St(!veville, on the Red Deer River, Alberta, Canada, in VX.V.\.

From an exhibition standpoint this si)eeimen is of interest in having
several patches of skin impressions preserved. When found the

• HufT( r, i^\t Marc Arnian<l, A pftlhoIoRlcnl sih-cIiiu'ii diillriK from tho Lower Mio«'iio imtIik!. ]n Appon-
(llx to Foiirtati's "C"onlrl»>iillon k \'t,i\ii\n Vfrt6br*s .Mloc^kmit do rfegypto," pp. 101-109, illiis. Survey
Dopartnient, MInhtry of FInanro, ralro, 1030.

' Moo'llo, U. L., ralropatliolo?) , p. 17.^, 1OT3.

202 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

skeleton was lying on its left side, but the definition of the skin pat-
tern of that side had been dulled or wholly destroyed by the presence
of a considerable amount of vegetal matter on which it lay. The
bedding plane beneath the skeleton was unusually in-egular, indicat-
ing that the cross-bedded planes were laid down by currents acting
from different directions.

The caudal series consists of 75 vertebrae and appears to be com-
plete. This information, together with that furnished by Brown,^
shows the complete vertebral formula of Corythosaurus to be 15 cer-
vicals, 19 dorsals, 8 sacrals, and 75 caudals. It is presumed, how-
ever, that as in many other reptiles the caudal series will be subject
to some individual variation in number. The first 16 caudal verte-
brae have transverse processes as in the type. The first chevron on
the tail of Corythosaurus is carried between the fifth and sixth caudal
vertebrae; thus the total number of caudal vertebrae can be accu-
rately determined and a close estimate of their combined length can
be obtained from this specimen which has the first two caudals com-
pletely hidden by the overlying ihum and a patch of skin impressions.
Measured along the curve this tail has a complete length of about 455
cm. (14 feet 11 inches).

A complete description of the ossified tendons of Corythosaurus has
already been given by Brown, and thus it is only necessary to men-
tion that the present specimen fully corroborates his determination
that they are disposed in two layers.

Small patches of skin impressions are present on the midsection of
the tail, on the pelvis, and on the feet. Those pieces of the integu-
ment best preserved cover the thirty-first to the thirty-fourth caudal
vertebrae, respectively. The detailed mosaic pattern of the flat,
polygonal scales is clearly and beautifully shown in plate 18. Origi-
nally the whole midsection of the tail beginning with the sixteenth
caudal was covered by skin impressions, but most of the center of this
patch was so friable that it could not be preserved. The outside
portions, however, outline the original width of the tail at this
point. The skin on and below the right ilium is composed of scales,
slightly larger than those of the midcaudal region, but otherwise
they seem to be indistinguishable. The pattern of the scales on
the feet is dim and illy defined and adds nothing to oiu- previous
knowledge.

In the course of preparing this specimen many small detached pieces
of skin were found in the matrix. Several of these were folded and
others had been completely reversed. Six species of this genus have
been named, aU from the Belly River formation of midwestern Canada.

I 8 Brown, Barnum, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 710, 1916.

REPTILE FOSSIL SKELETONS — GILMORE

203

Listed in chronological order these are: C. casuarius Brown,' C.
excamtus Gilmore,"^ C. infermeduis Parks," C. bicrisfatus Parks,'^
0. Jronfalis Parks,'* and C. breincristatus Parks.'*

Five of the six species were established on skull characters alono
and are distinguished chiefly by differences found in the shape and
extent of the crest. In the absence of the skull in the specimen under
consideration it appears quite impossible at this time to make a definite
identification of the species. However, on the basis of similarity of
skin pattern and close agreement in proportions to the type specimen
of Con/thosaurus casuarius, as shown in table 4, this specimen is
provisionally identified as pertaining to that species.

Table 4. — Comparative measuremenls (in cm.) of (u'o specimens of Corythosaurus

casuarius

Measurements

U.S.N.M.
No. 15493

A.M.N.n.
No. 5338, typo

Length ot longest che\Ton

Ischhim, greatest length

Ischium, length of terminal foot

Femur, greatest length

Femur, position of fourth trochanter from central point to top of femur

Tibia, length of tibia and astragalus

Fibula, length

40
113.6

20
115.9

58.5
100.7

98.2

38.5
lfr3

22
lOS

58
100

» Brown, Bamum, Bull. Amer. Mus. Nat. Hist., vol. 33, pp. 559-565, pi. 41, 1914.
'• Ollmore, C. W., Can. Field-Nat., vol. 37, pp. 46-52, 1923.

" Parks, W. A., Univ. Toronto Stud., geol. scr.. No. 15, pp. 1-57, 13 Ogs., pis. 1-4, 1923.
n Parks. W. A., Univ. Toronto Stud., gool. scr.. No. 37, pp. 2^45, pis. 4-8, 1935.

u t. aovcmiMCNT miNTiNa orricii it4«

U S NATIONAL MUSEUM

PROCEEDINGS Vni ''f. PLATE 12

r-y

_ »/

f the large palms.
It was met with occasionally in the ground-termite nests where it was
discovered when the test stakes were pulled up.

EPIPERIPATUS EDWARDSII (Blanchard)

Peripatus edwardsii Blanchard, Ann. Sci. Nat., zooL, ser. 3, vol. 8, p. 140, 1847
(Cayenne) ; Rech. Anat. et Zool. faites pendant un Voyage en Sicile, pt. 3,
p. 64, 1849.

Peripatus edwardsi Bouvier, Ann. Sci. Nat., zool., ser. 9, vol. 2, p. 301, pi. 9,
figs. 74-79, text figs. 5, p. 15, and 111, p. 308, 1905 (Panama; Dari6n).—
Fuhrmann, M6m. Soc. Neuchat. Sci. Nat., vol. 5, p. 190, 1912 (Brazil and
Guiana to Panama).

Epiperipatus edwardsi A. H. Clark, Proc. Biol. Soc. Washington, vol. 26, p. 18,
1913 (listed); Smithsonian Misc. Coll., vol. 63, No. 2, p. 2, 1914 (Isthmus
of Panama; from Bouvier); vol. 65, No. 1, p. 23, 1915 (Panama; Dari6n;
from Bouvier). — Fuhrmann, Abh. Senck. naturf. Ges., vol. 36, Heft 2, pp.
277-283, 1915 (listed).

OXYCHOPHORES OF PANAMA — CLARK AND ZETEK 211

New Records. — El Cermcno, Panama; J. Zctek, Jiily 8, 1941.
Female: (O 35 mm. lonp;, 2 mm. broad, 30 pairs of logs.

Balboa; J. Zotek, 1944. Females: (1) 50 mm. long, 34 pairs of legs.
(2) 50 mm. long, 33 pairs of logs. (3) 45 mm. long, 33 pairs of logs.
(4) 35 mm. long, 33 pairs of legs. (5) 25 mm. long, 33 pairs of logs.
Males: (1) 28 mm. long, 29 pairs of legs. (2) 27 mm. long, 29 pairs
of legs. (3) 25 mm. long, 30 pairs of logs. (4) 25 mm. long, 29
pairs of legs. (5) 25 mm. long, 29 pairs of legs.

Range. — Caj'enne; Surinam (Paramaribo); Venezuela (Caracas,
M6rida, Valencia, Haut-Sararc, Bas Sarare); Colombia (Santa
Marta); Panama (El Cermeno, Panama Station, Panama Railway);
Canal Zone (Balboa); Dari6n.

Notes. — At El Cermeno this species was found in situations similar
to tliose frequented by Oroperijyaius corradi, though more often under
boaj-ds resting on the ground.

EPIPERIPATUS BIOLLETI (Boavier)

Peripalus biolleyi Bouvier, Bull. Soc. Ent. France, 1902, p. 258 (San Jos6, Costa
Rica) ; Ann. Sci. Nat., ser. 9, zooi., vol. 2, p. 321, pi. 10, fig. 85, text figs. 115,
116. p. 323, and figs. 117, 118, p. 324, 1905.

Epiperipalus biolleyi A. H. Clark, Proc. Biol. Soc. Washington, vol. 26, p. 18,
1913.— FuHUMANN, Abh. Senck. naturf. Gcs., vol. 36, Heft 2, pp. 277-283,
1915. — A. H. Clark, Proc. U. S. Nat, Mus., vol. 85, p. 3, 1937 (Parismina
and La Caja, San Jos6, Costa Rica).

New record. — El Cermino, Panama; J. Zctek, July 1941. Females:
(1)41 mm. long, 4 mm. broad, 30 pairs of legs. (2) 38 mm. long,
3 mm. broad, 30 pairs of legs.

Range. — Costa Kica (San Jos6, and La Caja, San Jos6, Surubres
near San Mateo, and Parismina); Panama (El Cermeno).

Note. — At El Cermeno this species occurred in much the same
situations as E. edwardsii.

ADDENDUM

Dr. Otto Fuhrmann (1912, p. 190) gives Epiperipatus simoni
(Bouvior) as ranging from Bnizil and Guiana to Panama. There are
no published records of this species from anywhere west of Veno/iu la.
He also included "Brazil" in the range of Epiperipatus edwardsii, but
we know of no record of this species from that country.

212 PROCEEDINGS OF THE NATIONAL MUSEUM voL.g»

BIBLIOGRAPHY

Blanchard, Emile.

1847. Malacopodes (Malacopoda de Blainville). Ann. Sci. Nat., ser. 3,

vol. 8, pp. 137-141.
1849. Recherches sur I'organisation des vers. Pt. 3 of Milne-Edwards's

"Recherches Anatomiques et Zoologiques Faites pendaot un Voyage

sur las Cotes de la Sicile. . . ." 353 pp., 26 col. pis.
BouviER, Eugene L.

1899a. Sur les variations et les groupements sp^cifiques des P^ripates

am^ricains. Comptes Rendus Acad. Sci. Paris, vol. 128, pp.

1344-1346.
1899b. Nouvelles observations sur les Peripates americains. Comptes

Rendus Acad. Sci. Paris, vol. 129, pp. 1029-1031.
1900. Observations nouvelles sur les Peripatus [Onych.]. Bull. Soc. Ent.

France, 1900, pp. 394-395.
1902. Peripatus biolleyi, onychophore nouveau de Costa-Rica. Bull. Soc.

Ent. France, 1902, pp. 258-259.

1905. Monographie des on3^chophores. Ann. Sci. Nat., ser. 9, vol. 2, Nos.

1-3, pp. 1-383, 140 figs., 13 pis.

1906. Observation biologique. Le Peripatus edwardsi au Br^sil. Bull. Soc.

Ent. France, 1906, p. 268.

1907. Catalogue des onychophores des collections du Museum d'Histoire

Naturelle de Paris. Bull. Mus. Hist. Nat Paris, 1907, pp. 518-521.

1908. Sur le Peripatus brasiliensis Bouv. BuU. Soc. Philomat., ser. 9, vol.

10, pp. 50-52.
1928. A propos des observations du Fr. Claude-Joseph sur un P^ripate du

Chili. Ann. Sci. Nat., ser. 10, vol. 11, fasc. 2, p. 260.
Brues, Charles Thomas.

1911. A new species of Peripatus from Grenada, with observations on other

species of the genus. Bull. Mus. Comp. Zool., vol. 54, pp. 306-318,

2 pis.

1913. Preliminary descriptions of two new forms of Peripatus from Haiti.

Bull. Mus. Comp. Zool., vol. 54, pp. 519-521.

1914. A new Peripatus from Colombia. Bull. Mus. Comp. Zool., vol. 58,

pp. 375-382, 2 pis.
1917. A new species of Peripatus from the mountains of northern Peru.

Bull. Mus. Comp. Zool., vol. 61, pp. 383-387, 1 pi.
1923. The geographical distribution of the Onychophora. Amer. Nat., vol.

57, pp. 210-217.
1925. Notes on Neotropical Onychophora. Psyche, vol. 32, pp. 159-165.
1935. Varietal forms of Peripatus from Haiti. Psyche, vol. 42, pp. 58-62.
Brues, C. T., and Melander, Axel Leonard.

1932. Classification of insects: A key to the known families of insects and

other terrestrial arthropods. Bull. Mus. Comp. Zool., vol. 73,

672 pp.

(Onychophora, pp. 532-533, figs. 998-1002.)
Camerano, Lorenzo.

1898. Viaggio del Dr. Enrico Festa nella Repubblica dell'Ecuador e region!

vicini, VII: Onicofori. Bull. Mus. Zool. Anat. Comp. Univ. Torino,

vol. 13, No. 316, pp. 1-3.

ONYCHOPHORES OF PANAMA CLARK AND ZETEK 213

Clark, Austin Hobart.

1913a. A revision of the American species of Peripatus. Proc. Biol. Soc.

Washington, vol. 26, pp. 15-20, Jan. 18.
1913b. Notes on American species of Peripatus, with a list of known forms.

Smithsonian Misc. Coll., vol. 60, No. 17, pp. 1-5, Jan. 25.
1913c. Piccolc note su degli Onychophora. Zool. Anz., vol. 42, pp. 253-255,

July 18.
1914a. Notes on some specimens of a species of onvchophore {Oroperipalus

corradoi) new to the fauna of Panama. Smithsonian Misc. Coll.,

vol. 63, No. 2, pp. 1-2. Feb. 21.
1914b. On some onychophores {Peripatus) from the Republic of Panama.

Zool. Anz., vol. 45, pp. 145-146, Dec. 4.
1915a. The present distribution of the Onychophora, a group of terrestrial

invertebrates. Sniithsonian Misc. Coll., vol. 65, No. 1, pp. 1-25,

Jan. 4.
1915b. A note on the occurrence of Epiperipatus imthurmi (Sclater). Proc.

Biol. Soc. Washington, vol. 28, p. 182, Nov. 29.
1929. Peripatus from the Island of Montserrat. Proc. Ent. Soc. Washing-
ton, vol. 31, p. 139.
1937. On some onychophores from the West Indies and Central America.

Proc. U. S. Nat. Mas., vol. 85, pp. 1-3.

COCKERELL, ThEODORE D. A.

1913. A Peripatus from Guatemala. Proc. Biol. Soc. Washington, vol. 26,
pp. 87-88.
FuHRMANN, Otto.

1912. Quelques nouvcaux Pdripates am^ricains. M6m. Soc. Neuchdt. Sci.

Nat., vol. 5, pp. 176-192, 16 figs.

1913. t)ber einige neotropische Peripatus- Arten. Zool. Anz., vol. 42, pp.

241-248.
1915. Uber eine neue Peripatus- A.Tt vom Oberlauf des Amazonas. Abh.

Senck. naturf. Ges., vol. 36, pp. 277-283, 1 fig., 1 pi.
Mabcus, Ernst.

1937. Un onychophoro novo, Peripatus (Epiperipatus) cvrlinae, sp. nov., de

Goyaz. Rev. Mus. Paulista, vol. 21, pp. 903-910, 2 pis.

PiCADO, C.

1911. Sur un habitat nouveau des Peripatus. Bull. Mus. Hist. Nat. Paris,
1911, pp. 415-416.
Wheei.er, William Morton.

1898. A new Peripatus from Mexico. Journ. Mon^h., vol. 15, pp. 1-8, 1 pi.

■. t. •ovimaiNT mrRTiii* orricit if4«

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM

Vol. % Wathington: 1946 Nq. 3198

ECHiraOID AVORMS OF THE NORTH PACIFIC OCEAN

By Walter Kenrick Fisher

The echiuroids, sometimes called spoon worms from the shape of the
contracted prostomium or proboscis, are cigar-shaped or sausage-
shaped creatures, essentially highly muscular sacs filled with fluid in
which the long alimentary canal and other organs have great freedom
of movement. The mouth is anterior, usually at the base of a snout
or a long proboscis used for gathering food. The skin is highly gland-
ular and is covered by a very thin cuticle. Typically there are two
hooked setae behind the mouth, and two genera have either one or two
circles of setae at the posterior end of the body. The alimentary canal,
in contrast to that of most annelids, is several times the length of body
and consists of a long foregut, differc'ntiated into pharyiLX, esoi)hagus,
giz/.ard, and stomacii, a still longer midgut or intestine, accompanied
for a considerable part of its length by a collateral intestine, or siphon,
)ind finally a short hind-gut or cloaca, into which empty two usually
\<)hiniinc)us, sometimes branched, vesicles, the walls of which are
studded with minute ciliated funnels. The anterior nephridia,
tyj)ically elongate, thin-walled sacs, varying from one to many, but
usually from one to four pairs, hav(! a basal or terminal nephrostome,
the lips of which may be greatly prolonged and spirally twisted. The
nephridia vary greatly in size and when filbnl with eggs or sperm ar(^
(tftcn very large. The vascular system consists of a ventral vessel
f(»llowing the nerve cord to form a looj) in the proboscis from the tip of
u hich a median vessel pa.sses backward in tiie proboscis and along the
dorsal side of foregut to beginning of midgut, where a neurointestiiuil
vessel joins it to the ventral vessel. In Urechis there are no blood
vessels.

215
•""oaa*— *fl — 1

216 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Echiuroids are burrowers in mud or sand, where they fashion more
or less permanent tunnels. Sometimes they live under rocks; some-
times in mud-filled mollusk shells or sand-dollar tests, which afford
some protection; or they inhabit the rock galleries excavated by boring
clams. Their food consists of organic material contained in the mud
which they swallow in large quantities, or of lighter organic detritus
selected by the usually long proboscis. In the same species from dif-
ferent locaUties the intestinal pellets vary with the character of the
bottom. One genus (Urechis) has very specialized feeding habits and
uses only finely divided material, including bacteria. It is probable
that any small organisms living in the surface film of mud will be
eaten by echiuroids. Gislen (1940, p. 30) found the intestinal pellets
of Echiurus echiurus "to consist of the same stuff as that which is
formed by the detritus-film growing on the aquarium bottom. There
are thus plenty of sand grains; further the pellets consist of diatoms,
algal threads, debris of leaves of phanerogams (Zostera et al.),
Infusoria, Bacteria, occasional Nematodes and Rotatoria and, to a
large extent, of amorphous brown stuff which emanates from
decomposed organic substance."

The smallest sexually mature echiuroid I have ever seen is a
Listriolobus pelodes 7 mm. long (0.275 of an inch), and the largest is
Urechis caupo, 470 mm. long preserved, or 18.5 inches. I have seen
a relaxed living Urechis caupo 19.75 inches long. The Japanese
Ikeda taenioides, a remarkable and isolated form, attains a body
length of 16 inches with a proboscis of 58 inches, or a total length of
6 feet 2 inches (Ikeda, 1907, p. 20).

More helpless, unprotected animals can scarcely be imagined. The
immature stages are prey for every predaceous inhabitant of the
sea bottom. The adults are regularly eaten by fishes, especially
flatfishes and rays, as well as by the Indians of Chiloe Island, Chile
(Gay, 1854, p. 475). In Japan and Korea Urechis unicinctus is
extensively used as bait. Sato (1939, p. 319) states that in Korea
the natives catch it by means of iron hooks and dry it for food.

Systematics of the Echiuroidea present the usual problems in
addition to others inherent to the group. The principal diSiculty is
the lack of structures having a permanent form. The setae are of
very limited use; everything else is soft and capable of distortion.
The practical difficulties encountered are those which would confront
the student of holothurians if these creatures did not carry embedded
in the skin a species label in the guise of characteristic calcareous
deposits. Most of the generic and specific characters of echiuroids
must be sought by careful dissection of the internal organs, which
arc susceptible to variation arising from accidents of fixation. Never-
theless, a fairly satisfactory system of genera can be constructed.
But it is obvious that closely related species may not be recognized,

ECHIUnOID WORMS OF NORTH PACIFIC — FISHER 217

or if recognized they may be impossible to describe in the absence
of trenchant characters. Descriptions of echiuroids based on external
charactei-s or on a very summary enumeration of a few internal
features have made it difhcult to determine the generic position of a
number of described forms.

It has been a time-honored procedure to classify the echiuroids,
sometimes in combination with the sipunculoids and priapuloids,
under the name Gephyrea, as a class of the Annelida. In 1898 Prof.
Adam Sedgwick, in his "Students' Te.xtbook of Zoology," set up
separate phyla for the Sipunculoidea and Priapuloidea but retained
the Echiuroidea as a class of the Annelida. Since the development
of I rcchis caupo has been thoroughly elucidated (Xewby, 1940),
it is now known that the echiuroids are not more closely related to
annelids than to moUusks. Dr. Newby writes at length on a com-
parison of echiuroid development with that of the other invertebrates
and on the phylogenetic position of the Echiuroidea. In conclusion
he says (p. 209):

There are many echmroid characteristics which indicate that this group is
separate from the annelids: (a) The mode of development of the first somatoblast
is different, (b) The anus is not homologous in the two groups and no procto-
daeum is formed in echiuroids. (c) The mesodermal bands do not develop
teloblastically in echiuroids. (d) The elongation of the larva is not teloblastic
in echiuroids. (e) Three laj'ers of body muscles are formed in echiuroids. (f)
The ectomesoderm contributes to the body musculature in echiuroids. (g) A
ciliated intestinal groove is formed in echiuroids and this becomes the primordium
of the siphon. These structures are not found in the annelids, (h) Anal vesicles,
probably of endodermal origin, are found in echiuroids. (i) The coecuin of the
larval digestive tract becomes a linear part of the adult tract in echiuroids. (j)
The mesodermal bands of echiuroids show no evidence of segmentation, (k)
The lack of segmentation in the mesoderm considered with the questionable
nature of the segmentation of the nervous system and mucous glands and further
considered with the "segmentation" of the shell glands of the chitons (molluscs)
makirs it appear probable that the echiuroids have a i^riinary lack of metamerism.

.\gainst these numerous difTercnccs there are only three clear-cut characteristics
in common between the echiuroid.s and annelids which are not also po.ssessed by
the mollusks. (a) The annelidan cro.ss develops in both groups, (b) Both
groups possess setae. (However setae are to lie found in another group of animals
(Brachiopoda) which do not belong to the Annelida.) (c) The lateral halves of tho
nervous system become merged into single, unpaired structures.

With the above facts in mind it is evident that the echiuroids are only distantly
related to the annelids. When numerous differences which appear in their
fievelopment are con.sidcrerl, it seems improbable that the inclusion of the echiu-
roids with the annelids as a sub-phylum or cla.ss, is justified. It is probably more
accurate to consider the Echiuroidea as forming a separate phylum, di.stinct from
the f)hylum Annelida, and I herewith i)roi)()se that they be so Cftusidered.

In the keys no nicntion is made of Kpithetosoma Danielssen and
Koren, 1881. Th6cl (1900, p. 9) has demonstrated satisfactorily
that the animal is not an echiuroid, but most likely a nemertean.

218 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

Neither is Poeobius meseres Heath (1930) included. This remarkable
pelagic transparent worm was first taken in 350 meters, Monterey
Bay, Calif., and was later found to be abundant off southern Alaska.
Its anatomy has been fully described by Professor Heath. Sub-
sequently the writer observed and sketched a living animal. The
blood vessels are clearly visible and contain a dull green fluid, but the
enlargement of the dorsal vessel is dull red. Blood vessels extend to
tip of the 2 prostomial palps and the 10 (possibly peristomial) cirri.

The creature has no paired appendages, no somatic segmentation,
and no setae. "The nervous system conforms to the usual annelidan
type, with supra-oesophageal ganglion, circum-oesophageal connec-
tives, and a ventral nerve chain comprising 11 pairs of ganglia with
the usual commissures and connectives. The somatic musculature
comprises four great longitudinal bands, extending throughout the
length of the body, and a more delicate external sheath of circular
fibres." This is the annelid pattern and distinctly not the echiuroid.
The head is unlike that of any known echiuroid but resembles
that of some polychaete annelids. The alimentary canal and
nephridia seem to the writer to be specialized m much the same way as
in the case of Sternaspis, which in one species {S. spinosus Sluiter) has
the prostomium prolonged outward on each side to form a grooved
palplike organ. The Scoleciformia, however, have definite mesoder-
mal segmentation.

The difficulty in finding a place for Poeobius may well mean that it
is not an annelid or a echiuroid or a link between the two. Although
nothing whatever is known of the development of Poeobius, we have
to assume that mesodermal segmentation is absent; therefore it is not
an annelid. Its nerve cord is segmented (implying pseudometamer-
ism). The nerve cord of larval echiuroids is segmented, but this is
lost in the adult, suggesting that the ancestors, while deprived of
mesodermal metamerism, still had a pseudometamerism of the nerve
cord. A tenable hypothesis is that the echiuroids and Poeobius
stemmed from a common group that was as fundamentally un-
segmented as the Amphineura among mollusks. According to this
view Poeobius is the survivor of a lesser phylum, comparable to the
Phoronidea and Priapuloidea. As the genus now floats in a sort of
taxonomic limbo, it may be provisionally assigned to a new phylum,

POEOBIOIDEA.

The region covered by this report includes all the water north of a
line drawn from Cape San Lucas, Baja California, to the southern
end of Sakhalin Island on the east Asiatic coast. The Gulf of Cali-
fornia has been included, and a species long ago dredged by the
Albatross in Japanese waters has been added, as it modifies the
concept of Acanthohamingia, which I wished to include in the key.

ECllILKOlU WUK.MS OF NUHllI I'AflFlC — FISHER 219

Tlio spcciinons upon svhirh tliis ])apfr is based have been accuinu-
lati'il slowly over a considerable perioti of 3'ears.' In addition, the
material belonjrinf: to the United States National Museum was
placed at my disposal, and an iiu])ortaTit collection belon<rin<2: to the
Allan Hancock Foundation of the I'niviTsit}' of Southern California
was tcndereil by Dr. C)l<;a Ilartman. The types of all the new species
are in the collection of the National Museum.

The followino; ni'W <^eiuis, based on an extralimital species, will be
found in the text: Lissomyema, tyj)e Thalassema mellita Conn (under
Listriolobus).

Phylum Echiuroidea'

Ik'hiuroidea Sedgwick, 1898, p. 527 (class of Annelida).

Unsegmented. bilateral, fusiform or saccuUform animals with
anterior mouth and posterior anus, but no proctodacum; a long con-
voluted alimentary canal lying in a spacious coelom of scliizocoelous
type; a muscular body wall composed of three layers, of which the
middle (with one I'xception) is composed of longitudinal fibers; with
one to very numerous anterior nephridia functioning as gonothecae;
with typically two anal vesicles having numerous ciliated fuimels
and functioning as excretor}' organs; alimentary canal tyjiically with
collateral intestine or siphon; usually with a ])rostomial proboscis,
which nuiy exceed length of boily but which is sometimes absent;
usually with ectodermal setae, of which two, ventrally situated be-
hind mouth, are most constantly present, together with sometimes
one or two circles at posterior end of body; but setae absent in a few
genera; ventral nerve cord unsegmented forming, around the mouth,
a loop which follows border of proboscis; gonad, where known, in
mesentery above nerve cord, or hi the mesenteries surrounding cloaca.

KEY TO CLASSES

o'. Body wall with inncniiowt circular or oljliijiic layer uf inu«ck's well dcvoloped;
anal vcHicles present; collateral intestine or siphon well developed; pro-
bosciH and anterior setae present in nearly all species Echiurida (p. 220)

• I am Mix-cifilly ln-holflon to niy foriiuT colloiKiic I'rof. fii'<)rKe E. Mncniiilllc for iiiiitcriMi of I'rrrhit
caupo, LiMriolobu* prlodfM, nrul Ochdottoma ocloinvotum; to Eilwnrd F. Kickctts for tlu' type of l-xhiurua
eehiurui ataikanut nn<l n sriinll colli-ctlon miiric by lilni and Join) Stcliilx'rk In the (liilfof C'liliforniii: to Dr.
Olitn llHrtiiinn (or n spccltncii of Limnmvrmn mrlliln: to I'rof. S. K. Lluht for ft [M-rfcct sihtImicii of lAntrMnlitu
ptlodtt; Ui tlm Mii.wum of Companitivo '/ool'>Ky for a .^in'i-lmoii of L'reckin chtlentin; to Dr. \V. I,. I.loyd,
Cnbrillo .Vlarlni^ Miiacum, San I'wlro, Calif., for the loan of a spiclnii'ii of Ochiloitoma octorni/ntum; to I'rof.
John n. (iprnuM for the loan of scvcml rare rcprlnU; and to Dr. Waldo L. Schinitt, U. S. National Museum,
for numerous favors.

• Kchluroldra wa.i Introdutwl m a subphylum by A. II. riark (Hull. Inst. OcAannpr. Monaco, No. 400,
p. 24, 1021) and a.s a [iliyluni by W. W. Newby (IWO. p. 210) Bn<l I.lbblc II. Ilyinan (HMO, pp. 34, M). As a
matter of record, F.dwaril K. Hlcketls wiwt the first to u.ifl Kcliluroldi-a aa n i>liyluin name. In an exwllont
semlpopulnr text "Betweon Pacific Tl«|t«" (UlcketLs and C'ajvln, 19.10, p. 272), i>ertiap.s incited thereto by
tho present writer who has a>l%'ocnted this proc«<luro to his classes for 20 ycnrs.

220 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

0^. Body wall with innermost circular layer missing or degenerated to a net
of fibers; no anal vesicles; apparently no siphon; no proboscis and no
setae Sactosomatida »

Class ECHIURIDA

KEY TO ORDERS

o'. In body wall longitudinal muscle layer lying between outer circular layer
and inner oblique layer; nephridia, normally paired, not excessively nu-
merous.

6*. A closed blood- vascular system; no specialization of intestine for anal

respiration Echiuroinea Bock (p. 220)

62. No vascular system, coelomic fluid being heavily charged with large blood
corpuscles containing hemoglobin or hemoglobin plus hematin; in-
testine with terminal portion enlarged, thin-walled, to receive water

from cloacal pump Xenopneusta, new order (p. 262)

a?. Longitudinal layer of body wall lying outside of both the circular layer and
inner obHque layer; nephridia excessively numerous, unpaired (and with
terminal nephrostome) ; proboscis excessively long.

Heteromyota,* new order

Order Echiuroinea Bock, emended

A closed blood-vascular system; no specialization of intestine for
anal respiration.

KEY TO FAMILIES

ci. Dimorphic; male degenerate, planarianlike, parasitic in or on female; female
resembling Thalassema but with bifid proboscis in some genera; anal vesicles
consisting of branched tubules ending in numerous cihated cups; anterior
setae sometimes present; posterior setae absent Bonelliidae (p. 249)

a?. Not dimorphic; proboscis usually conspicuous, sometimes several times length
of body, but never bifid; absent in one genus; anal vesicles not branched
but in form of elongate sacs, surface of which is covered with minute cilated
funnels; anterior paired setae present in all genera, posterior setae in
Echiurus only Echiuridae (p. 221)

3 New name for Saccosomatida Thfiel (1906, p. 14). Th§el instituted the group as a suborder for Sacco-
soma vitreum Danielssen and Koren (1881, p. 34, pi. 6, figs. 1-8). This species is based on a single small
example dredged in 1,215 fathoms north of the Faroe Islands. It is a female and the species may prove to
be dimorphic, as there is a single nephridium filled with eggs and opening near the mouth. The proboscis
may have been lost. It is aberrant from all other echiuroids and may not be an echiuroid. Saccosoma
Danielssen and Koren is preoccupied by Saccosoma Motschoulsify, 1859, in Coleoptera (Bull. Acad. St.
Pfetersbourg, vol. 1, column 304). The new name "Sactosoma" (with identical meaning) is proposed to
replace Saccosoma Danielssen and Koren.

* Based on the remarkable genus Ikeda Wharton, 1913, pp. 243-270. Type, Thalassema taenioides Ikeda,
1904, p. 63; 1907, p. 16, pi. 1, fig. 3; pi. 2, figs. 18-22; pi. 3, figs 23-36; pi. 4, figs. 37-47. This large echiuroid,
with a proboscis upward of a meter or more in length and nephridia from 200 to 400 in number without
indication of paired arrangement, is so different from the general run of the phylum that it deserves to be
set apart as the type of at least a distinct order. The arrangement of muscle layers is different from that of
all other echiuroids and indicates a long separation from typical stock. See Sato, 1931. p. 179.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 221

Family ECHIURIDAE (de Blainville, 1827, restricted)

KEY TO GENERA

a'. Two circles of posterior setae Echiurus Gu6rin-M6ncville (p. 225)

a*. No posterior setae present.

b^. Proboscis absent Arhynchite Sjito (p. 247)

6*. Proboscis present.

c*. No differentiated thicker bands in longitudinal muscle layer,
d*. Nephrostome of nephridia without elongated, si)irally coiled lips.

Thalassema Lamarck (p. 230)
(P. Nephrostome with elongated, spirally coiled lii)s.

Anelassorhjmchus Annandale (p. 221)

c*. Longitudinal muscle layer with very slight to pronounced differentiation

into longitudinal bands, 8 or more in number.

d'. Nephrostome of nei)hridia without spirally coiled lips; inner layer of

muscles not differentiated into separate transverse fascicles between

longitudinal bands Lissomyema, new genus (p. 224)

(P. Nephrostome with elongated spiral lips.

e'. Differentiated longitudinal muscle bands weak, zones between not
showing a fasciculate arrangement of inner oblique muscles; in
small specimens longitudinal bands very faint or visible only in

posterior region Listriolobus W. Fischer (p. 233)

e'. Longitudinal muscle bands strongly developed, zones between
crossed by separated fascicles of innermost, oblique layer.
p. Nephridia in 1 to 5 pairs; vascular ring vessel at beginning of

midgut Ochetostoma Lcuckart and Ruppell (p. 240)

p. Nephridia, at least in male, in to 14 groups of 1 to 4, the groups
arranged in pairs; vascular ring vessel at posterior end of
pharynx Ikedosoma Bock (p. 224)

Remarks. — In the foregoing synopsis all the generic divisions,
with the exception of Echiurus, are the result of subdividing the old
genus Thalassema. In a very real sense these groups are provisional
because adequate descriptions and figures of the internal structure
of many species have not been published.

Thalassema Lamarck.^ — The geiuis has been restricted to a few
species grouped around the type, Thalassema thalassema (Pallas),
generally known as lli. neptuni Gaertner. The middle, longitudinal
layer of muscle fibers of body wall shows no sign of differentiation
into thicker bands. The internal opening of the nephridia is very
simple, without prolongation into si)irally coiled lips.

ANELASsoRHYNrHis Aiuiandale (1922, p. 148). — It may not be of
any practical value to recognize this group. The species dilfer from
Thalassema in having the nephrostome lips j)rolonged and spirally
coiled, but little is known of other details of the internal anatomy.
Amiundalc based the genua on the structure of the |)r()boscis of four
cstuarinc species occiiniiig in lnackish water of India and Siam.
He says:

The genus con.sists of Kchiuridae allied to ThnlnHmma Gaertner, but differing
in the structure, function, and phy.siology of tlic proboscis. Thi.s organ is rela-
tively stout and short, incapable of great prolongation or autotomy. The ciliated

222 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se

groove on its ventral surface is feebly developed and the lateral margins of the
ventral surface bear (except in A. microrhynchus) gill-like outgrowths. The
longitudinal muscle-fibres of the body form a single sheath and the musculature
bears a close resemblance to that of some species of Thalassema. There are two
pairs of nephridia. The anal funnels are simple and thin- walled; their ciliated
funnels are minute.

The type-species is A. branchiorhynchus (Annandale & Kemp). The other
species are A. dendrorhynchus (Annandale & Kemp), A. sabinum (Lanchester)
and A. microrhynchus (Prashad).

It seems to me that the modifications of the proboscis, which
exhibit a number of gradations in complexity, are adaptations to an
ecology in various ways abnormal, a parallel development being
found in Ochetostoma arkati (Prashad). But these species agree with
certain others in having a more specialized nephi-ostome than is
found in Thalassema thalassema and close allies.

1. With two pairs of nephridia (behind the setae): sabinum Lanchester,
branchiorhynchus Annandale and Kemp, dendrorhynchus Annandale and Kemp,
microrhynchus Prashad, semoni Fischer.

2. With three pairs of nephridia, all three opening behind the setae: mucosa
Ikeda, vegrande Lampert (no proboscis). First pair opening in front of setae:
inanense Ikeda, moebii Greef.

Unless some definite character other than the nephrostome is dis-
covered, there will be a practical difficulty in distinguishing young
Listriolobus , in which the differentiation of longitudinal muscle
bands is very weak.

LissoMYEMA. — Through the kindness of Dr. Olga Hartman I
have received a specimen of Thalassema mellita Conn collected by
her at the type locality, Beaufort, N. C, in June 1940. It is 36
mm. in length, with proboscis 16 mm. additional. From the outside
the eight longitudinal muscle bands are clearly visible. Figure 10
represents a dissection of the anterior portion. The muscle bands
are much more sharply delimited than in Listriolobus by having an
incipient fasciculation of the muscles of the oblique layer, possibly
representing the first stage in the differentiation of the strong trans-
verse bundles characteristic of Ochetostoma. The species has simple
fan-shaped nephrostomes and very heavy interbasal and radiating
seta muscles. The gizzard is relatively short and the stomach (C)
is relatively long. An individual variation is the presence of three
nephridia on one side and two on the other. The species is described

Figure 10.^ — Lissomyema mellita (Conn): Dissection of anterior region of a specimen from
Beaufort, N. C, X 12. Six of the eight muscle bands are diagrammatically indicated by
dots. The alimentary canal is drawn to the right to disclose the organs beneath it.
{B^, &, B*, dorsal, neurointestinal, and ventral blood vessels; C, stomach; G, gizzard;
/, intestine; MD, middorsal muscle band; MI, interbasal muscle; MF, midventral muscle
band; N, nephridia; NC, nerve cord; O, esophagus; P, pharynx; S, seta; Sil, beginning of
siphon.)

ECHIUHOII) W01{MS OF XOHTH PACIFIC — FISHER 223

FiGURK 10.— (Sec opposite page for legend).

224 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

as having two pairs. The anal vesicles are voluminous with numerous
conspicuous ciliated funnels. The ventral blood vessel sends an
important branch to the pharynx and esophagus.

For this species, therefore, I propose the new genus Lissomyema,
which differs from Thalassema in having eight well-differentiated longi-
tudinal muscle bands and incipient fasciculation of the oblique layer;
from Listriolobus and Ochetostoma in having simple fan-shaped nephro-
stome without trace of spiral extensions. Type, Thalassema mellita
Conn. (Fig. 10.)

Ikedosoma Bock. — Thalassema elegans Ikeda^ does not belong in
Thalassema. Ikeda (1907, p. 50) writes: "All the longitudinal lines
visible on the outside, excepting the one which runs in the mid-ventral
line and is superposed by the nerve-cord, appear on the inner surface of
the body-wall as slightly elevated narrow ridges or thickenings oj the
longitudinal muscular layer. In the ten zones separated from one
another by the above lines, the circular muscle fibres form more or
less regularly arranged transverse bundles." This structure of the
body wall closely approximates that of Ochetostoma, but elegans is
peculiar in having numerous (13 to 27) nephridia in six or seven
pairs of groups comprising one to three nephridia each. "The internal
opening present at base is provided with 2 relatively short spiral
lobes." The dorsal blood vessel ends with the "heart" on the hind
end of the pharynx and is therefore shorter than in typical Echiuridae.
"The neuro-intestinal vessel arises from the ventral median point of
the ring-sinus, which surrounds the extreme hind end of pharynx"
{ibid., p. 52). There is no interbasal muscle and no intestinal coecum.

Thalassema gogoshimense Ikeda (1904, p. 66, pi. 1, fig. 19) is ap-
parently congeneric with elegans. The excellent colored figure shows
the same white longitudinal stripes as elegans, reflecting the muscular
structure of body wall. Ikeda says: "It shows an essential agreement
with Tlialassema elegans. Indeed, the agreement may be said to be
complete, the only difference being that all the visceral organs in the
present species are developed on a smaller scale in proportion to the
smaller size of its body." In the females, however, the nephridia
are present in three pairs all situated behind the setae, while in the
male they are in six to eight groups, arranged in pairs, each group
with one to four nephridia, which are like those of elegans in structure.

The above paragraphs were written and a name was assigned to
the genus before I saw Bock's paper. They are retained since we
independently arrived at the same conclusion.

« Ikeda, 1904, p. 66; 1907, p. 47, pi. 1, fig. 4; pi. 4, figs. 48, 49; Sato, 1939, p. 366; Bock, 1942, p. 18.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 225

Genus ECHIURUS Gu^rin-iMeneville

Echiurus GifRiN-MfiNEViLLK, 1831, p. 9, pi. 6, fig. 3 (ex Shipley, 1899, p. 342)
(type, Lumbricus echiurus TaUas, 1700= Echiurus pallasii CUi<5rin-M<5nevilIe,
1831). — Skorikov, 1909, p. 80. — Spengkl, 1912b, p. 173.

Echiuriclae with two rings of posterior bristles, a well-developed
proboscis, two or four nepliridiii (without spirally coiled lips), and a
postpharj-ncjeal tliaphragni, which separates incompletely the small
liead coelom from the perivisceral cavity.

ECHIURUS ECHIURUS ALASKANUS. new subspecies

Plate 20

Echiurus Pallasii C. B. Wilsov, 1900, p. 174.
Echiurus echiurus Spengel, 1912b, p. 183.

Diagnosis. — Differing from typical E. echiurus (Pallas) of the north
Atlantic and neighboring Arctic Ocean in having the proboscis strongly
attached to the body and in having the posterior setae definitely
curved lather than nearly straight. Length of type, 230 mm. plus
much contracted proboscis, 20 mm.

Description. — ^Length of body upward of 230 mm., commonly 100
mm., stout; proboscis adherent, fleshy, convex above, the edge in-
curved ventrally, su})truncate distally, usually 15 to 20 mm. long in
contracted state. On ventral suHace of the proboscis a difTcrentiated
thickening extends as a low ridge from the mouth for about one-
fourth length of proboscis but sometimes considerably farther. The
integument is roughened by rings of prominent vermcae most crowded
at ends of body. In the middle region, where they are generally
loss crowded, rings of more prominent verrucac alternate with zones
of three to five nngs in which the verrucac are smaller or more widely
spaced, or both. The appearance depends largely upon the degree
of contraction of the body muscles.

The anterior setae are stout, strongly curved, and situated back of
the base of proboscis a distance e([ual to about its greatest width. In
each circle of anal bristles there may be variations of G to 8, as: 8-8
(posterior ring); 8-7; 8-G; 7-7; 7-G. In some specimens where the
number is less, inequality of spacing indicates loss of setae. These
pf)stcrior setae \t\ry in degree of curvatun* but are slightly more
curved than in typical A', ecldurus. The anterior setae have a strong
interbasal muscle connecting their inner ends. Occasionally a second
seta, in process of formation, a(;comi)anies one of the primary. The
principal posterior muscle from the setae attaches to body wall just
behind the anterior nephridium.

The inner circular layr of body muscle shows n division into closely
placed fascicles at each end of body, where the animal is nonnally
most contracted.

226 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Nephridia 4, the anterior pair close behind the setae and a little
farther from nerve cord. The funnel is conspicuous, with an undu-
lating or frilled border, but is not prolonged into spiral lips. The
nephridia of all specimens examined (taken in summer months) were
contracted. In some cases they were very small, and the anterior
pair very inconspicuous.

Anal vesicles are simple, elongate, thin-walled sacs attached to
ventrolateral wall of the cloaca and closely beset with minute ciliated
funnels.

Diaphragm (pi. 20, figs. 1, 4). The diaphragm is a curious, thin-
walled, funnel-shaped septum incompletely separating the peripharyn-
geal coelom from the general body cavity. Its general form is best
appreciated from the figure in which it is shown in a semidiagrammatic
fashion. The anterior, rouglily circular edge is completely attached to
the body wall, while ventrally it is attached to body wall on each side
of the nerve cord (which here lies within the ventral mesentery of
pharynx and esophagus). A large oblique posterodorsal opening of the
diaphragm (with complete free edge) allows the esophagus (with its
strong ventral mesentery) to pass backward into the general coelom,
sometimes above and sometimes below the interbasal muscle. The
rim of aperture apparently has a sphincter. The two halves of the
double ventral mesentery of esophagus merge with diaphragm along
its paraneural part and a short distance above the nerve (pi. 20, fig. 5).

Alimentary canal. The pharynx remains always in the peri-
pharyngeal coelom. It is attached to the body wall by numerous strong
muscular strands having an annular arrangement. There is a con-
tinuation forward of the double ventral and dorsal mesenteries sep-
arated into frenula. The dorsal blood vessel lies in this mesenterial
complex. The head cavity is therefore much occluded by tissue.
The lining of pharynx is anteriorly thrown into coarse folds.

The esophagus begins just behind the region of the radiating frenula
of pharynx. It has, in the anterior portion, a dorsal mesentery of
slender separate strands, but there is a double membranous ventral
mesentery throughout its whole extent. This mesentery is anchored
in the peripharyngeal chamber on each side of the nerve cord, where, a
short distance above body wall, it merges with the diaphi-agm. By
means of its muscular mesenteries, all of the esophagus can be with-
drawn mto the head cavity.

The esophagus, on passing through the right side of the diaphragm
close to posterior border, becomes a long gizzard, marked by rings,
which are prominent amiular ridges of the lining. Beguming with the
gizzard the alimentary canal is moored only by dorsal mesenterial
ribbons as far as the cloaca, which has radiating muscular frenula.
Along the dorsal side of the gizzard held by a perforated mesentery

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 227

is the voluminous dorsal blood vess(>l with numerous papilliform
l)ranchos, at least anteriorly.

A very short rudimentary stomach or crop lies between the gizzard
and liesritming of intestine (indicated by the ventral ciliated p-oove).
The lining of stomach is thrown into 12 strong longitudinal folds,
contrasting sharply with the annular folds of gizzard. Where tiie
stomach becomes intestine, the dorsal blood vessel splits to form tiie
ring vessel.

The intestine has the usual three parts: presiphonal, siphonal, and
postsiphonal. The first is about as long as the gizzard, or a little
longer if relaxed.

The siphonal part, rougldy 20 to 25 times length of presiphonal part,
is marked by longitudinal folds of the lining which arc evident super-
ficially. The siphon is about one-fourth the diameter of the intestines.

The postsiphonal intestine has thinner walls and is about 10 times
the length of presiphonal segment. The ciliated groove forms a
ridge along its ventral side, and ends at a coecum (not always inflated)
just in front of the cloaca. The fecal pellets which fill this part of the
intestine are elongate ellipsoids and sometimes contain coarse material.
I have found leaves of the hemlock (Tsuga).

Vascular system. This consists of a dorsal and ventral blood
vessel and neurointestinal connective. These vary in caliber in dif-
ferent specimens. The dorsal vessel is likely to be considerably in-
flated over part or the entire length of gizzard, vnih irregular lobose
swellings anteriorly. The ventral vessel, attached to middorsal line
of nerve cord, ends posteriorly as a solid cord just in front of the in-
testinal coecum. The neurointestinal connective results from the
brandling of the dorsal vessel at the beginning of intestine by which
tlie neurointestinal ring (B^) is formed. The connective branches
again (5^), to form the muscle ring, before merging broadly with
the ventral vessel (B*).

Type.— v. S.N M. No. 20G09.

Type locality. — Auk Bay, Juneau, Alaska, collected by E. F. Rick-
etts, August 14, 19:n.

Specimens examined. — One hundred and twenty -four as follows:

Collection of Stanford Univkk.sity •

Kiikak Bay, Shelikof Strait, Alaska, 12 apccimcns, uruicr rocks, in iimd; McMillan,

1924.
\\ rangell, Ala.ska, 37 .si)ccimen.s; A. \V. Greely and R. K. Sno(lnra.ss, 1897.
Auk Bay, near Juneau, Ala.ska, 3 specimens, slate beach, under rocks, in murldy

.sand, lowest intertidal zone, July 17, 1931; K. V. Hickclts (also typo from

this locality, August 14, 1931).
Huston Inlet, Queen Charlotte Islands, Briti.sh ('(jlunihia, 42 specimen.^; W. F.

Thornp.son, July 1, 1913.
Alaska, pos.sibly Dutch Harbor, t'nalaska, 5 specimens.

228 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Collection of U. S. National Museum

Cape Smyth, Alaska, 3 fathoms. No. 850, Point Barrow Expedition; 2 specimens

in very bad condition, without proboscis ; possibly these are intermediate with

true echiurus.
Off Cape Strogonof, Alaska, Albatross station 3291, lat. 56°58'30" N., long.

159° 11' W., 26 fathoms, black sand, gravel, 1 specimen from stomach of cod,

in poor condition.
Bristol Bay, Alaska, No. 4597, 2 specimens, one lacking proboscis.
Unalaska, Alaska, No. 16314, 2 specimens.
Bering Island, Nos. 4151, 16315, 2 specimens, typical.
Alitak Bay (head of Lazy Bay), Kodiak Island, gravelly sand, January 22, 1941,

5 specimens.
Stepovak Bay, Alaska, 15-90 fathoms, October 24, 1940, 1 specimen.
Dolgoi Harbor, Alaska, October 6, 1940, 1 specimen.
Wrangell, Alaska, No. 4538, August 28, 1882, under stones, 2 specimens.
Chasina Bay, Alaska, No. 4601, 1 specimen.
Saginaw Bay, Alaska, No. 4117, 1 specimen.
Without stated locality (probably Unalaska), 5 specimens.

Remarks. — The type specimen was taken from its burrow, a tube
in sandy clay beach, 24 inches below the surface, along with a com-
mensal polynoid annelid, Hesperonoe adventor (determined by Dr.
Olga Hartman). The same species is commensal with Urechis
caupo.

This, the common Alaskan Echiurus, differs from the typical form
of Europe and the north Atlantic coast of America in having the
proboscis firmly attached to the body. All writers who have handled
living Echiurus echiurus emphasize its habit of dropping the proboscis
on the slightest provocation.® Most of the 120 specimens of alaskanus
were not handled with care but were simply dropped into alcohol; 106
of these specimens still have the proboscis firmly attached, while at
least 5 lost the proboscis subsequent to fixing, apparently from rough
handling.

Under ordinary circumstances I should have given Brandt's name
sitkaensis to this form, assuming that Mertens would naturally have
picked up at Sitka the common Alaskan species. J. W. Spengel
(1912b), however, succeeded in obtaining one of Mertens's two spec-
imens, upon which Brandt based his description, and found that it
differed fundamentally from Echiurus echiurus in having only two
nephridia, as well as in certain other respects. The name sitkaensis
is therefore definitely associated with a type specimen which has been
redescribed by one of the best zoologists of his time.

The ecology of Echiurus echiurus has been studied by Dr. Torsten
Gislen (1940) chiefly at Kristineberg, Sweden. His very compre-

« Torsten Gislfin says: "As stated before the proboscis is very easily thrown off. In fact probably only
very few men have seen a proboscis in connection with an Echiurus. Forbes and Qoodsir say that it is so
slightly affixed to the body as to break off at the least touch; in only one or two cases did they find it attached,
and then it broke away immediately on the removal of the animal. Only in some exceptional instances
have I been able to secure specimens with the proboscis retained." (Gisl6n, 1940, p. 10.)

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 229

hensive memoir is in the forefront of cxcollonco and will long servo as
a model for this type of work.

ECHIURUS SITKAENSIS (Brandt)

Thalassema (Echiurus) sitkaensis Brandt, 1835, p. 62.
Echiitrus silkaeyisis Spengel, 1912b, pp. 184-189.

Diagnosis. — Corpus circiter tripollicarc oblongum, e subbrunneo
olivaceum, obscurius punctatum et transversim striatum. Proboscis
latiuscula, carnea, transversim purpureo striata, apice emarginata.
Unguiculi anterioris corporis partis et spiculac posterioris lutea.
(Brandt.)

Differing from E. echiurus in having tw^o nephridia, in lacking a
differentiated ridge of tissue along ventral side of proboscis, and in
having skin papillae subequal rather than in rings of larger papillao
alternating with narrow zones of smaller.

Remarks. — This species constitutes one of the major mysteries in
the systematics of the Ecliiuroidea. Ivlertens collected two specimens
at Sitka, both of wliich he dissected. One of these specimens, his
notes on the dissections, and a life sketch in color reached the St.
Petersburg Museum and were used by Brandt. Subsequently all
these became available to Spengel, as he details in his Echiurus paper
(1912b).

Spengel made a thorough examination of what remained of the
internal anatomy and was able to satisfy himself that only two nephrid-
ia were present, in the location of the anterior nephridia of E. echiurus.
The proboscis was very adherent to the body, and it lacked the ridge
of tissue on its concave under surface. As less important differences
he lists: skin papillae subequal, in rather regular and very numerous
rings (not rings of larger pa])illae alternating with zones of smaller);
curvature of anal setae stronger than in echiurus; color, according to
Mertcns's drawing, browiiish olive spotted and cross-stripad with
darker, the proboscis flesh color with i)ur])le transverse stripes.

The tougli, nondcciduous proboscis is characteristic of the Alaskan
I'lchiurus I have examined, but all these have the ventral ridge
I)resent, leaving as the princi])al characters of sitkaensis the two ne-
phridia, absence of proboscis ridge, and the subequal papillae.

Wilson (1900, p. 174) states that he examined Alaskan specimens
of E. echiurus { = ala.skanu.'i) collected by J)r. W. R. Coo in 1899.
"This species was found abundantly at numy different localities
along the Alaskan coast south of the Peninsula and on adjacent
ishuids, nearly always in rich black mud." I have listed 120 speci-
mens from Alaska and British Columbia. None of these is sitkaensis.

If there ii.v a sj)ecies sitkaensis it may nonnally live below low tide
and only occasionally be carried shoreward during heavy storms.

230 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Type. — Formerly in St. Petersburg Museum; collected by H.
Mertens.

Type locality. — Sitka, Alaska.

Genus THALASSEMA Lamarck

Thalassema Lamarck, 1801, p. 328 (type, Lumbricxis thalassema ' Pallas, 1771,
Spicilegia Zoologica, fasc. 10, p. 8, pi. 1, fig. 6).

Diagnosis. — Echiuridae with a well-developed proboscis but \\dthout
anal bristles and without specialized bands in the longitudinal layer
of body muscles; inner oblique layer smooth, except sometimes for a
short distance at anterior end of body ; anterior nephridia (gonothecae)
one or two pairs, the internal ciliated funnel (nephrostome) without
spiral lobes. ^

THALASSEMA STEINBECKI, new species

Figure 11

Diagnosis. — Small, slightly translucent, the proboscis as long as
body, broad proximally, ribbonlike distally; nephridia, two pairs, the
cUiated funnel with simple subcircular opening lacking any trace of
sphal lips; interbasal muscle of setae well developed, strong, passing
through loop of dorsoventral blood vessel; siphon beginning a short
distance from vascular ring; precloacal intestinal coecum; intestinal
mesenteries including conspicuous subfusiform fleshy masses; anal
vesicles as long as contracted body, covered with numerous tiny
ciliated funnels. Length of body 12 mm.; of proboscis, 12 mm.

Description. — The skin is coarsely verrucose for a short distance
back of proboscis and on terminal tliird of body; elsewhere the verrucae

' Thalassema was first used in a generic sense by Lamarck. Although Pallas (Spicilegia, 1771) mentions «
the name Thalassema as used by Joseph Gaertner, he names the animal Lumbricus thalassema, which appears
to be the first valid binomial referring to the species generally known as Thalassema neptuni Gaertner. The
latter is in effect a manuscript name. Neither Shipley (1899, p. 351) nor Wharton (1913, p. 265), who have
offered revisions of Thalassema, gives any reference for the combination Thalassema neptuni. Quatrefages
(1865, vol. 2, p. 595) cites "Thalassema Neptuni Gaertner, citfi par Pallas, Spicilegia Zoologica, fasc. 10, p. 8,
pi. 1, fig. 6." Forbes, 1841, in his "History of British Starfishes and Other Animals of the Class Echino-
dermata," gives a good account of the habits of "Gaertner's spoon-worm, Thalassema Neptuni Gaertner."
The first item in his list of references is Lumbricus Thalassema Pallas.

The derivation of the word seems to be thnlassos (sea) + ema (dart).

8 Shipley (1899, p. 351) in his revision of Thalassema is in error in the statement that the nephridia have
their internal openings gpirally twisted. Lankester (1881, p. 355) writes that they are semicircular and
contrasts them with the spiral sort found in T. moebii, as figured in Greef's "Die Echiuren" (1879, pi. 8, fig. 69).

Figure 11.— Thalassema steinhecki, new species: A, The type, X 14, showing arrangement of
organs; the greater part of the intestine has been omitted and the foregut has been strongly
bent to right in order to uncover the nephridia. B, A seta, X 20, from a specimen taken
in 165 fathoms off San Francisquito Bay, Baja California; above, the hook, further en-
larged. C, Ventral view of the type, X 4. {AV, Anal vesicle; B^-B*, dorsal, ring, neuro-
intestinal, and ventral vessels; C, stomach; CG, ciliated furrow of intestine; CI, cloaca;
G, gizzard; /, intestine; IC, intestinal coecum; Me, mesenterial bodies; MI, interbasal
muscle of setae; N, nephridia; NC, nerve cord; 0, esophagus; P, pharynx; S, seta; Sil,
anterior end of siphon.)

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 231

FicuRK 11. — (See opposite pa^e for legend).

232 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

are obviously smaller, and there is a suggestion of transverse align-
ment. The appearance of the skin will depend a good deal on the
state of contraction of body wall. The middle portion of body is
slightly translucent, not transparent in places as in Listriolobus pelodes.

The setae have a stout interbasal muscle connecting the inner ends,
from which radiate numerous fan-shaped sheets of muscle which
attach broadly to the body wall. A posterior muscle passes over the
first nephridium and merges with the body wall between the first and
second nephridia. Setae 1.8 mm. long with a small hook, the point
of which is at right angles to the shaft. In the type and a number of
other specimens the hook is worn off.

Inner layer of muscles of body wall smooth except in front of
anterior nephridia where there is differentiation into about a dozen
bundles passing around the body. The middle layer is undifferen-
tiated as in other species of the restricted genus Thalassema.

The four nephridia are small in the type, which is probably a male.
In the specimen from station 2828 they are enormously inflated and
full of eggs. In the type the ciliated funnel is very simple, with
smooth lips forming an incomplete circle. In the female the funnel is
flattened and pear-shaped and somewhat distorted by pressure, but
there is no indication of elongation into spirals.

The deflated anal vesicles are as long as the body and have the
appearance of being capable of inflation to a great size.

The alimentary canal is moderately long but conspicuously shorter
than that of Listriolobus pelodes. The contents are not always
formed into pellets. The foregut consists of a pharynx-esophagus,
midway along which is a sharp bend. Both the gizzard and stomach
are unusually short. The former is marked by ring striations. At
the posterior end of the stomach is the ring blood vessel (5^) marking
the beginning of the intestine, on the lower side of which is the ciliated
groove. At a distance back of the ring vessel equal to about length
of gizzard and stomach the siphon begins and runs for about half
length of intestine. The ciliated groove continues to the intestinal
coecum in front of the cloaca. The pharynx has a conspicuous
ventral mesentery, and all along the canal are numerous dorsal
mesenterial strands. Posterior to the region of the siphon the mesen-
teries inclose, or hold, conspicuous yellowish-white masses, sometimes
subfusiform or in irregular sheets, which remind one of the suet found
in mammals. A squeeze reveals several sorts of cells, some of which
may be immature sperm. The cloaca is small and thin-walled.

The vascular system consists of the dorsal vessel ending posteriorly
in the ring vessel, from which the neurointestinal connective passes
to the ventral vessel, forming a loop enclosing the interbasal muscle
of setae. The ventral vessel ends posteriorly at the intestinal coecum.
In the female from station 2828 the vessels are the same.

ECIIIUKOID WORMS OF NORTH PACIFIC — FISHER 233

TVk-— U.S.N.M. No. 20G00.

Type locality. — Ei Mogote, near La Taz, Baja California, low tide,
March 22, 1940, 1 specimen, Steinbeck and Ricketts,
Specimens examined. — Eight as follows:

Albatross station 2S2S, Gulf of California, 2-1° II' 30" N., 109° 55' W., 10 fath-
oms, shells, 1 specimen.
La Plata Island, Ecuador, 7-10 fathoms, rocky with nuUipores, 2 specimens,

Allan Hancock Foundation.
Thurloe Bay, Baja California, S-10 fathoms, rock with gorgonids, 1 specimen,

Allan Hancock Foundation.
Off San Francisquito Bay, Baja California, 1G5 fathoms, shale and gray mud, 1

specimen, Allan Hancock Foi^ndation.
Ensenada de San Francisco, Baja California, 2-6 fathoms, 1 specimen, .\llan

Hancock Foundation.
.\gua Verde Bay, Baja California, 10 fathoms, mud and coral, 1 specimen, Allan

Hancock Foundation.
Dewey Channel, San Eugene Point, Mexico, 21-24 fathoms, coralline, rock, I

specimen, Allan Hancock Foundation.

Distribution. — Baja California to Ecuador, low tide to 1G5 fathoms.

Remarks. — Mr. Ricketts states that the type was associated with
living Denfalivm in sandy mud a short distance below the surface.
The specimen from station 2828 has the intestine filled mth small
fragments of shells.

As this species belongs in the restricted genus Thalassema, it
naturally resembles Th. thalassema. The gizzard, stomach, and
presiphonal intestine are definitely longer in the latter species and the
anal vesicles smaller, although with such extensible structures it is
difficult to make comparisons. It may be recorded that m alcoholic
specimens there is no division of the body into three parts, wliich
Lcigh-Sbarpe (1928, p. 501) reports as a characteristic of thalassema.
The ecology of the two species is very different. Lankester (1881.
J). 350) found thalassema on the south coast of Devonshire "in galleries
excavated in tlie red sandstone (not limestone) which is exjiosed at
spring tides. The galleries appear to be those formed by the Lamel-
Hbranch Gastrochaena which the Thalassema appropriates." Leigh-
Sharpe (1928, {). 499) reports tlu- species from borings in hmestono
made by the mollusk Saxicava, Fl^^mouth Sound.

Named for John Steinbeck, whose expedition to the Gulf of Cali-
fornia collected (lie type.

Genus LISTRIOLOBUS W. Fischer

Lislriolobua Spenokl, 1012c, p. 310 {noitien nudum). — W. Fisciiku, 1920n, p. 210
(no type). (Type, Listriolohux hahamcnsia Fischer.)

Diagnosis. — DifTcring from Thalassema, sensu stricto, in having
elongate, s|)irally (H)iled lips to neplirostome and 8 to 10 narrow merid-
ional thickening.s of the middle, loiigitudinnl muscle layer. Differing
from Ocheiostoma in having the inner oMiciue layer a smooth con-

234 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

tinuous sheet between the longitudmal bands, not divided into
separate fascicles. Nephridia 2 or 3 pairs; interbasal muscle of setae
present.

The character of the longitudinal muscles is not well marked in L.
pelodes until the animal has reached a length of 15 to 20 mm. and
after it is sexually mature. Apparently in L. sorbillans (Lampert) a
similar condition exists (Wharton, 1913). Even so, the structure of
the nephrostome will segregate these species from true Thalassema.
L. riukiuensis (Sato, 1939, p. 359, figs. 10-13) is probably not a
Listriolobus. There is no interbasal muscle to setae; the diagram of
the blood vessels is not the Listriolobus pattern.

Spengel (1912c, p. 316) established Listriolobus for Thalassema
erythrogrammon of Sluiter (1883) and of Wilson (1900). Sluiter's
species came from BHliton in the Java Sea whUe Wilson's was taken
in the Bahamas, Spengel had Sluiter's animal and a duplicate, from
Florida, of Wilson's species, which he characterized as "nearly re-
lated." Unfortunately, as neither of these species had a valid name,
Listriolobus was without a type and was technically a nomen nudum,
Spengel did mention Thalassema mellita Conn as a species inquirenda,
"which species one must include in the above genus I can not at
present decide, nor even whether or not it is justifiable to include
such a form as Th. mellita, which has bundles of longitudinal muscles
that arise from thickenings of the continuous longitudinal muscle
layer." A few lines farther on he says: "The animals described by
Sluiter and Wilson as Thalassema should be put in the same genus
with Th. mellita because of the nature of the sheaths surrounding the
tunic muscles. I propose the generic name Listriolobus for the species
of Sluiter and Wilson."

Wilhelm Fischer (1926a, p. 110) discusses Listriolobus and names
Wilson's and Sluiter's species Listriolobus bahamensis and L. billi-
tonensis. So far as I know this is the first association of Listriolobus
with a species from which a type can be chosen. Since Fischer did
not do this I will so designate Listriolobus bahamensis Fischer, as
being the species more likely to be available for study in the future.
By implication Fischer includes Th. mellita Conn in Listriolobus, but
I have made it the type of a new genus, Lissomyema.

LISTRIOLOBUS PELODES. new species

Figures 12, 13; Plate 21, Figures 1, 2, 4, 4a, 4b; Plate 22

Description. — Largest specimens 40-60 mm. long and 12-25 mm.
thick; proboscis capable of extension to slightly exceeding length of
body, narrow, thin, translucent, so that nerve loop is visible.

Body wall tanslucent, marked by eight narrow longitudinal muscular
thickenings which appear light gray agamst the darker and much
broader areas between them. There are a middorsal and midventral

ECHIUROID WOKMS OF NORTH I'ACllTC — FISHER 235

Figure U— Listriolobus pelodtj, new spoics: Small 8j)ccimcn, X IS, showing arrangement
of organs. The greater part of the intestine has been removed. In this specimen the
ncphridia arc very small. Lettering as in figure II.

236 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

band and three lateral, equidistantly spaced. The bands, 1.5 to
2 mm. broad, represent concentrations of longitudinal muscle fibers,
which gradually thin out on the sides of the bands. The skin is
beset. with small, unequal, subcircular and elliptical glandular thick-
enings arranged in transverse close-set lines. At posterior end there
is an area of greatly enlarged papillae. Coelomic surface of body
wall smooth; innermost layer of slightly oblique muscle fibers very
thin, uniform, not mterrupted by the longitudinal thickenings of
the middle layer.

Setae 2, close together, and close to the mouth. Sometimes two
setae occupy one of the sheaths, with a single normal one in the
other. There is an interbasal muscle uniting top of the sheaths. This
passes through a loop in the dorsoventral blood vessel.

Nephridia 4, variable in size. In the type the anterior pair is
smaller than the posterior, but in another specimen the anterior
pair is the larger and the four are about five times the length of those
of type. They have the same terminal slender portion, which can
undoubtedly be expanded. Both specimens are males. The anterior
pair is situated posterior to setae about the length of the latter. The
ciliated funnel has long coiled lips and is attached to the outer side
of the base of nephridium by a short stalk. In a third specimen the
nephridia are empty and reduced to filaments slightly expanded at
base. In a full-grown female (Tomales Bay) the nephridia are 20
mm. long, slender, and contain numerous eggs (June 7), 0.08 to 0.09
mm. in diameter.

Anal vesicles 2, variable in size, capable of great extension. Each
is fastened to body wall about 5 mm. laterally from the anus by one
or two mesenteries and ventrally by another pair close to nerve.
Into these, which appear hollow, extends a short diverticulum of
the vesicle. There are scattered, very tiny, ciliated funnels. The
well-preserved female from Tomales Bay has vesicles that extend as
far forward as the large posterior nephi'idia and lack the basal diver-
ticulum found in the male.

Alimentary canal. The pharynx extends to the sharp bend shown
in the illustration, followed by the esophagus. The gizzard is short
and the stomach relatively long, but there is bound to be variation in
different specimens o^ving to the accidents of preservation. The
intestine proper starts just back of the ring blood vessel, at the
beginning of the ciliated groove, which soon becomes differentiated
into the siphon. Even in the carefully hardened specimen from To-
males Bay the intestinal wall is delicate and transparent. The
length is difficult to measure on account of unequal contraction. The
siphonal part is on the order of 100 mm. in length and the postsiphonal
125 mm. (length of specimen, 40 mm.). Throughout its length the
intestine is stuff ed with small unequal (1-2 mm.) ellipsoidal mud pel-

ECIIIUHOID WORMS OF NORTH PACIFIC FISHER 237

Figure 13. — Listriolobuj pdodes, new species: A sexually mature but small female, showing
nephridia greatly swollen by contained eggs (not indicated), X 15. Lettering as in
figure 11.

238 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

lets. Just before the liind-gut passes into the small cloaca there is a
spherical ventral coecum, to which extends the ventral blood vessel.

The vascular system can be readily followed on plate 22. The
neurointestinal connective forms a loop around the interbasal muscle
of the setae. The two flaplike expansions of this vessel, shown in
the figure, are not present in the Tomales Bay specimen. The dorsal
vessel varies widely in diameter being sometimes greatly inflated
(Tomales Bay), the inflation extending into the neurointestinal
connective.

Owing to the very thin, translucent body wall the nerves can some-
times be seen under strong illumination such as sunlight. They pass
directly around the body, from the ventral nerve cord, without visible
branches. The translucent proboscis affords an opportunity to trace
the proboscis loop throughout its entire course (pi. 21, fig. 2), This
ganglionic continuation of the ventral cord is near the margin of the
proboscis, to which numerous tiny nerves pass from slight ganglionic
thickenings on the outer side of the cord. No nerves, under favorable
conditions, could be detected on the mesial side of the cord.

Color in hfe: "Proboscis yellow orange, deepest on edge; body dull
grey-violet with greenish raised specks about 0.5 mm. in diameter,
spaced quite regularly about 0.5 mm. apart (the papillae); 8 Ughter
colored muscle bands" (large specimen from Tomales Bay).

Small phase (figs. 12, 13; pi. 21, figs. 4, 4a-b). Small examples
are fairly common on muddy bottom in moderate depths off southern
CaHfornia and in Newport Bay. These preserve badly unless special
care is exercised. They are usually strongly contracted into sub-
spherical form and vary greatly in appearance. Sometimes the body
wall is rather uniformly transparent but more often the posterior
portion is opaque. This region may be smooth or thrown into eight
meridional swellings which occasionally extend the whole length,
giving the appearance of a tiny melon. These swellings are caused
by the contraction of the eight muscle bands and are found in strongly
contracted specimens 7 mm. long. But specimens in which the body
wall is stretched and transparent do not show indications of differ-
entiated muscle bands until much later — at a length of about 15 to
20 mm. In the very small sizes the muscles are Ukely to show first
posteriorly and ventrally. A specimen 20 mm. long from 55 fathoms
off Santa Cruz Island has the 8 bands fully developed. Even in large
specimens (40 mm.) the bands are not always equally conspicuous.

I have found a specimen 7 mm. long sexually mature. Normally
these small examples are transparent in the midregion so that the
nerve cord, intestinal pellets, and egg-laden nephridia can be plainly
seen as yellowish bodies in life. The intestine is characteristically
thin-walled, transparent, and highly convoluted and taxes the capacity
of the body cavity. Its walls are greatly distended with mud peUets

ECHIUROID WORMS OF NOHTH PACIFIC — FISHER 239

about 1 mm. long. A specimen from Los Frailes, Baja California?
has the intestine distended with sand not in pellets.

The specimen shown in plate 21 , figure 4, was 12 mm. long and 9 mm.
liiick when alive, contracting to about 10 by 10 mm. when killed.
The transparent areas of body wall are characteristic and are dotted
with gi'ayish or wliitish papillae. In life the general tone is greenish
or olive, with gi'ay papillae spots, the proboscis bordered with yellow.
In the clear area the lateral nerves can be seen as they leave the ven-
tral nerve cord and in favorable specimens the nerve loop can be
traced around the entire margin of proboscis.

Dr. Olga Hartman describes small specimens from off southern
California as being rich, dark, satiny green in life, wliile those from
Newport Bay Professor MacGinitie found to be decidedly greenish.

Text figures 13 and 14 indicate how widely the neplu-idia vary in
size. In the swollen state the wall is perfectly transparent and
excessively thin and the proximal parts of the four vesicles adhere to
each other, the distal part lying at random, crowded amid the close
coils of the mud-filled intestine. The eggs are suspended in a thin
gel, spaced 1 or 2 diameters apart. They vary from 0.09 to 0.1 mm.
in diameter and are gi-avish in color, surrounded bj^ a clear zone
(about 0.009 mm. thick). The clearly visible nucleus is 0.045 mm. in
diameter.

In text figures 12 and 1 'A the vascular system is shown in a contracted
state. It does not (lifFcr in essentials from that of the fully adult
(pl. 22).

7;v/>^.— U.S.N. M. No. 20008.

Tyj)e locality. — Monterey Bay, Calif., moderate depth, fine sand;
from stomach of flounders.

Specimens examined. — As follows:

Monterey Bay, 10 large and 4 small specimens.

Tomalcs Bay, Marin County, Calif., low tide, soft black sandy mud, about 6

inches below surface, 1 large female, June 7, 1941 (University of California).
Newport Bay, Calif., 7 to 20 fathoms, numerous specimens, 4 to 1 5 mm. ; collected

by Prof G. E. MacCinitio.

The Allan Hancock Foundation tendered for examination an impor-
tant collection from Baja California and off southern California, com-
j)rising the small phase and some of intermediate size (20 to 40 mm.):

JiAJA Camkornia

Lfis p'raih-s, 5 1.') fathoms, sand aixl nlnac, 1 specimen.

I'ppcr end of Culf of ("aiifornia, 21 fathoms, brown mud, 6 specimens.

Camkciuma

Off Newport Beach, ."iO fathoms, mud, 7 specimens.

Off Laguna Beach, 2.'> .'>7 fathoms, sandy mud, 2.') specimens.

240 PROCEEDINGS OF THE NATIONAL MUSEUM vol. sa

Off Bluff Cove (8 lots) 25-100 fathoms, mud, fine sand, 32 specimens.

Off Redondo Beach (15 lots), 10-120 fathoms, mud, fine sand, coarse sand and

mud, 44 specimens.
Off Point Vicente Lighthouse, 17-40 fathoms, coarse sand and mud, 4 specimens.
Off Portuguese Point, 16-20 fathoms, gray sand and seaweed, 2 specimens.
Cortes Bank, 60 fathoms, sand, broken shell, 3 specimens.
Off Point Mugu, 26-30 fathoms, mud, 2 specimens.
Santa Catalina Island, 50-51 fathoms, mud, 3 specimens.
Santa Cruz Island (5 lots), 31-138 fathoms, mud, sand, 15 specimens.
Santa Rosa Island, 28-45 fathoms, 5 specimens.
San Miguel Island, 35 fathoms, mud, 20 specimens.

Remarks. — The type and largest specimens were recovered from
the stomach of flomiders from moderate depths of Monterey Bay.
In nearly all the examples the intestme is disintegrated and the body
cavity filled with pellets of fine sand. This is true also of specimens
from southern California, which were dredged and placed almost at
once into alcohol.

After the drawings and description were completed I received from
Prof. S. F. Light a carefully hardened specimen from Tomales Bay,
collected at low tide, in mud frequented by the clams Schizoihaerus
nuiialli and Macoma secta. This specimen (June 7) was apparently
laying eggs as the nephridia are partly emptied. It is especially valu-
able as it gives the characters of the adult female, is much better pre-
served than the Monterey specimens, and affords opportunity for
recording the life colors of a large example.

In the type specimen, near the anterior left nephridium is a light-
colored lobed mass (pi. 22, x) adherent to body wall. It is possibly a
parasite.

Genus OCHETOSTOMA Leuckart and Riippell

Ochetostoma Leuckart and RtJpPELL, 1828, pp. 7-8 (type, 0. erythrogrammon
Leuckart and Riippell). — Spengel, 1912c, p 316.

Diagnosis. — Greater part of the thickness of the longitudinal muscle
layer segregated into separate longitudinal bands. The intervals
between these bands is crossed by very niunerous separate small
muscle bundles of the inner oblique layer, which remains for the most
part continuous and unbroken over the surface of the longitudinal
bands. Anterior nephridia, 1 to 4 pairs, the coelomic aperture having
spirally coiled lips; with or without coecum at end of intestine; inter-
basal muscle of setae present or absent; two ventral anterior hooked
setae; no anal setae.

Remarks. — The list of species that follows is mostly derived from
literature and is in nowise intended to be a revision, for which speci-
mens and new dissections will be absolutely essential. Most of the
species were described as Thalassema. Th. exilii (Fritz Miiller)
Lampert, which has 8 to 10 muscle bands and two pairs of nephridia,

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 241

with nophrostomc lips merely folded and crinkled, but not elongated,
is possibly a Lissomyema.

a^. Five pairs of nephridia, 3 rather poorly developed, in front of setae; 19 muscle

bands -- O. homelli (Prashad)

Five or four pairs of nephridia, 2 in front of setae; 10 or 11 muscle bands.

O. bombayensis (Prashad and Awati)
a'. Four pairs of nephridia, posterior to setae.

10 muscle bands O. decameron Lanchester

20 muscle bands O. kempi (Prashad)

a}. Three pairs of nephridia, the first pair opening in front of the ventral setae
the second and third pairs posterior to setae.

14 muscle bands O. erythrogrammon LcuoJcart and Ruppcll*

15 or 16 muscle bands O. stuhlmanni (Fischer)'

15 to 17 muscle bands O. leptodermon (Fischer)'

16 to 18 muscle bands O. caudex (Lampert)'

17 or 18 muscle bands O. kokotoniense (Fischer)'

17 or 18 muscle bands O. grifflni Wharton

a*. Two pairs of nephridia.

7 or 8 muscle bands; papillae white, scattered uniformly all over body;
color white in alcohol; proboscis one-third to one-half length of
body; anal vesicles broad and saclike O. formolosum (Lampert)

8 muscle bands.

With incipient gills on edge of proboscis at base; broad smooth zone

around anus arkati (Prashad)

No branchial fringe to proboscis; no smooth zone at posterior end
of body; color in life greenish, in alcohol grayish flesh color;
proboscis one-third to length of body.

octomyotum, new species
10 or 11 muscle bands O. hupferi (Fischer)

13 muscle bands; proboscis longer than body; anal trees one-fourth length
of body O. pellucidum (Fischer)

14 muscle bands; anal trees more than one-half body kiigth.

O. manjuyodense Ikeda

17 to 19 muscle bands; proboscis shorter than body; color dark green

with violet longitudinal stripes and with white papillae scattered all

over the body; anal vesicles long, brown, pointed anteriorly, bearing

short branched outgrowths O. baronii (Greef)

16 or 17 muscle bands; proboscis deciduous, can equal length of body;
setae with interbasal muscle; anal vesicles without branched
outgrowths... _.0. edaz, new species

OCHETOSTOMA OCTOMYOTUM. new Bpcclcs
Plati: 21, Fun;uK 3; Platks 23, 24

ViagnoHu. — Skin transluoent, with ei^ht more opatiiic" lonpitudinnl
strifx'S marking the inuselc bands, and closely sti|>ple(l everywiiere
witii very small, une<|ual, often elliptical glands wliieh increase in
size very nuirkedly over the j)osterior region (where tiie niusele bands
fuse into a continuous sheet); i)roboseis thick and fleshy in preserved

• Sato, 1930, p. Vil considers these flvc, bIopk with Thnlntitma palmie lkr>Iii, Urjl, to I'O nil ono sprcii'M. fdr
which trythronTammon U the oldest name.

242 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

specimens, varying from less than one-third to the full length of body.
Four nephridia, the nephrostome with very long coiled lips. Siphon
begins a short distance behind the ring blood vessel (marking the
boundary between foregut and intestine) ; anal vesicles long, with
pointed apex and with tiny ciliated funnels scattered over surface;
setae without interbasal muscle. Length of full-grown specimen 110
mm. ; proboscis 30 mm. ; diameter 20 mm. A specimen collected at
Cabrillo Beach, San Pedro, Cahf., is 95 mm. long; proboscis 93 mm.

Description. — Longitudinal muscle bands 8, well developed, broader
than the intervals between, rather iridescent and situated as a mid-
dorsal, midventral, and (twice) three laterals. These fuse into a
continuous sheath on the posterior part of body, equal to about one-
third body length of expanded specimens and less of contracted ones.
Anteriorly they remain separated to the base of proboscis. The
innermost or oblique muscles are well developed, and between the
longitudinal muscle bands they form a consecutive series of oblique
partitions alternating with narrow compartments. At the bottom
of these compartments is a thin layer of the longitudinal middle sheet.
In the posterior region the oblique fascicles become less and less
distinct as the anal area is approached. In reality the uninterrupted
layer of longitudinal muscles is covered by a continuous inner layer
of oblique muscles differentiated into thicker and thinner portions.
Thus, forward from the anus the gradual differentiation of the separate
oblique strands can be easily seen. In severely contracted specimens
this posterior region does not show to advantage.

Anterior nephridia, 2 pairs, the ducts piercing the midventral
muscle about midway between its outer margin and the nerve cord.
The anterior pair is situated a short distance behind the setae (rather
less than length of seta). The interval between first and second is a
little more than twdce the distance. They open internally by ciliated
funnels having very long, coiled, extensible lips.

Anal vesicles 2, unbranched, capable of great distension. They open
on the ventral surface of cloaca. Tiny scattered ciliated funnels
may be seen on the surface.

Alimentary canal. There are three general regions: (1) An
anterior division or foregut, in which the longitudinal musculature lies
outside the circular musculature; (2) an intestine proper, in which this
order is reversed, characterized by the presence of the ciliated groove
and along a part of its course by the collateral intestine or siphon;
and (3) a short rectum, or cloaca, posterior to the coecum.

The foregut consists of the pharynx, esophagus, gizzard, and stom-
ach, and the junction with intestine coincides \vith the position of the
ring blood vessel. The pharynx has a tough wall, rather iridescent
with wavy longitudinal lines; the esophagus is thin-walled and con-
voluted, while the gizzard and stomach, both brief, can be differen-

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 243

tinted by texture. The gizzard is nmri-ced by eircular lines, the stomach
by longitudinal representing the longitudinal folds of the lining.
Complicated dorsal and ventral mesenteries attach the pharyiLX and
esophagus to body wall.

The hitestine is, as usual, highly convoluted but when partly un-
raveled is seen to have a principal posterior loop, and then an anterior.
The siphon, or collateral intestine, begins a short distance behind the
ring blood vessel, its course being shown in plate 23, figure 1, where
the dotted line indicates the ciliated groove, which terminates at the
coecum. The walls of the cloaca have thick longitudinal folds. The
anal vesicles empty into it by veiy small pores.

Blood-vascul.\r system. The dorsal vessel, intestinal ring vessel,
doi-soventral comiectives, and ventral vessel are shown in plate 24.
The ventral vessel, which anteriorly follows a part of the free edge of
the ventral mesentery, terminates posteriorly on the wall of the
coecum.

Color in life, greenish; in alcohol grayish flesh color with eight dark
grayish stripes.

fyi^f.— U.S.N. M. No. 20607.

Type locality. — Newport Bay, Orange County, Calif., January-
Februarj^ 1930, G. E. MacGinitie, 11 specimens.

Other material examined. — Cabrillo Beach (San Pedro), Calif., 1
specimen, W. R. Lloyd, Cabrillo Beach Marine Museum.

Remarks. — Newport Bay, where Professor MacGinitie collected
tho tyi)e series, is a rather small tidal inlet south of Long Beach.
The worms were taken from a sandy bar, exposed at low tide, where
they inhabited U-shaped burrows, the mouths of which were 12 to 14
inches apart and the bottom 10 to 12 inches below the surface. A
mutilated specimen was dredged in 25 fathoms.

This sand bar was later removed in order to deepen the harbor, and
tiie species was apparently wiped out at that locality as it has not since
Incn taken in spite of nmch collecting.

Tlie food consists of very fine detritus, which in (he intestine is
fornu'd into ellipsoidal pellets 2 to 2.5 mm. long by 0.75 mm. thick.

The only species with which 0. octomyotum might be confused is 0.
formolosvm (Lampert) from the Pliilippincs and Shanghai. This is a
small form averaging 30 mm. in length, |)r()boscis 8 mm., and diameter
10 mm. The color is white in alcoholic sjx'cimens, the skin very
thin, with white papillae scattered uniformly all over the body. The
anal vesicles are described as broad saclike organs. The species has
seven or eight muscle jjundles, two pairs of nephridia with spirally
coiled openings, and a .spherical diverticulum on the rectum — charac-
tcri-.tics of odorinjotum, except the nunil)(r of nmsclcs which is con-
stantly eight in the California si)ecies.

244

PROCEEDINGS OF THE NATIONAL MUSEUM vol.

Figure 14. — Ochetostoma edax, new species: A, Dissection of anterior portion of body to
show relations of nephridia, setae, vascular system, and anterior part of alimentary canal,
X 6. B, Ventral view of specimen from Puerto Refugio, X 2. (TV, nephridium similar
to the four of type, a male; N^, female nephridia from a smaller, Puerto Refugio, speci-
men. Other lettering as on plate 24.)

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 245

The specimen from the Cabrillo Marine Museum was taken in sand
and is notable for the 93-mm. proboscis, whicli is IG mm. broad near
tne base. The nerve loop, following the margin, is plainly visible.

OCHETOSTOMA EDAX, new species

Figure 14

Diagnosis. — Two pairs of nephridia situated posterior to setae, the
nephrostome with spiral lips; setae with interbasal muscle passing
through a small loop of the neurointestinal blood vessel; longitudinal
muscle bands 16 or 17, with very narrow interspaces; rather long pre-
siphonal intestine; precloacal coecum; two unbranched anal vesicles
capable of great distension. Body wall translucent in middle region,
striped with narrow dark zones; skin papillae numerous, sometimes
whitish, in not well-defined transverse lines, and usually larger on
posterior third or fourth of body, but not hard to the touch; proboscis
deciduous, fleshy, from one-fourth to body length. Length of body
25 to 50 mm.

Description. — The IG or 17 muscle bands merge into a continuous
sheet in ttie posterior fourth of body. The intervals between the
bands are narrower than in octomyotum and transverse oblique fas-
cicles of the inner layer are decidedly weaker. The ventral muscle
band under the nerve cord is broader than the others. Sixteen is
probably the normal number of bundles, the extra band arising from
the incomplete splitting of one of the lateral or dorsal bundles.

Nephridia 2 pairs, opening at outer margin of midventral muscle
and varying greatly in size. In a male the nephridia are very large,
the posterior reaching to posterior end of (contracted) specimen,
while in a smaller female they have the relative size of the three
smaller nephridia of figure 14. The ciliated funnel on the anterior
face of uephridium has long coiled lips. The two anal vesicles, with-
out branches, are covered with ciliated funnels and are capable of
being distended to a large size. They are as long as the body in the
largest specimen, which is, however, strongly contracted.

Ali.ment.\ry canal. Forcgut relatively short, ending with the
ring blood vessel (IP). Oidy two parts are clearly difi'erentinted, the
phar}-nx-esophagus and stomach, with longitudiiud folds. The
gizzard of 0. octomyotum appears to be lacking. The intestine is
long, highly convoluted, witii a very much longer presiphonal segment
than in octomyotum, being lojiger than the foregut. The wall is thin
and in the type is greatly distended with coarse sand; in another
specimen, with sizable fragments of shells and miscellaneous hard
deV>ris. In the type the presiphonal intestijie measmes roughly
30 mm., the siphonal part 120 mm. and postsiphonal 90 mm. The

246 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

latter has a ciliated groove terminating at the coecum just in front
of the cloaca. The walls of the coecum are much thinner than in
octomyotum and lack heavy longitudinal folds.

Blood-vascular system (fig. 14, B^-B^). This is on the same
plan as in octomyotum, but in two specimens the left dorsoventral
connective from the ring vessel was much longer than the right.
Passing through the small loop where these connectives join the
ventral vessel is an interbasal muscle of the setae, not present in
octomyotum. The ventral vessel ends posteriorly in the coecum
without branching.

Color. "Specimens were elongate to grape-shaped, smooth and
thin-skinned, greenish, with obvious and comparatively large spoon-
shaped proboscis" (Steinbeck and Ricketts).

T2/2?e.— U.S.N.M. No. 20606.

Type locality. — Gulf of California: Pichalingue Bay, near La Paz,
Baja California, February 1920, Luis G. Rubio.

Specimens examined. As follows:

Coronado Island, Gulf of California, March 27, 1940, Steinbeck and Ricketts,
3 specimens, under and among slightly subtidal rocks on white sand.

Point Lobos, Espiritu Santo Island, Baja California, March 3, 1940, Steinbeck
and Ricketts, 1 specimen, under boulders of tide flats.

Puerto Refugio, Angel de la Guarda Island, April 2, 1940, Steinbeck and Ricketts,
10 specimens; under boulders on beach. All these are smaller than the type.

Same locality, Allan Hancock Foundation, 1 specimen.

Remarks. — The type is without proboscis. The sketch of the en-
tire animal is from an example taken at Puerto Refugio. As some of
the specimens lack a proboscis and others have only a small one it is
probable that the organ is soon regenerated. The wide difference in
size seems hardly to be due to accidents of fixation.

The great disparity in size of the nephridia in the two specimens
dissected shows that size is of no particular value as a character,
depending as it does on the amount of conta'ined material. There is
a wide difference also in the size of the anal vesicles in the two specimens.

0. edax feeds upon the sand or coarser material in which it lives.
This is not molded into definite fecal pellets. The specimen from
Puerto Refugio had eaten very coarse material, which formed irregular
masses in the intestine. Among the miscellaneous material could be
recognized fragments of pelecypod and gastropod shells and small
whole gastropod shells; serpulid tubes, calcareous bryozoans, barnacle
shells; chelae of small crab; fragments of crab carapace, sea-urchin
spines, and brown algae; many straight sUiceous sponge spicules and
fragments of volcanic rock. Some of the sponge spicules were in
bundles. Others had perforated the intestinal wall and were lying
in the coelomic cavity. A few were in the anal vesicles.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 247

In the list of species, edax is next to bnronii, wliich has 17 to 19
nuiscle banils and anal glands which are described as having short
branching outgrowths.

In July 191S I collected at English Harbor, Antigua, B. W. I., a
small Ochetostoma that has 19 muscle bands, 2 pairs of nepliridia,
a strong interbasal muscle passing tluough a loop of the dorsoventral
blood vessel. The anal vesicles are long, reaching to the anterior end
of the contracted specimen and are covered with ciliated cups, but
there is nothing that can be interpreted as branched outgrowths. The
proboscis is much contracted, fleshy, and very adherent. The
posterior papillae are wliite, conspicuous, crowded, so that they are in
contact, and are hard to the touch. In a section of the body wall they
resemble calcareous nodules. This posterior part of the body is
perfectly opaque while the rest is slightly translucent. This species is
perfectly distinct from edax, and it is noted here because outwardly it
would pass for 0. baronii. The specimen is a mature female with the
four nepliridia enlarged and full of eggs. Length of body, much
contracted, 20 mm.; thickness, 9 mm.; proboscis, 8 mm.

Genus ARHYNCHITE Sato

Arhynchite Sato, 1937, p. 142. (Type, Thalassema arhynchite Ikeda.)

Diagnosis. — Differing from Thalassema in absence of a proboscis;
nephridia 2, with spiral lobes to nephrostome; no intestinal coecum;
no ring vessel around foregut at end of dorsal vessel, the connection
between dorsal vessel and neurointestinal connective being indirect
as in Bonellia; muscles of body wall smooth with no concentration of
fibers in either the middle, longitudinal layer or in the inner, oblique
layer.

ABHYNCMITE INAMOENUS. new spcciea

Platk 25

Diagnosis. — Differing from A. arhynchite (Ikeda) in having a
relatively longer neurointestinal blood vessel, which does not embrace
the interbasal muscle. Intestine vciy long, especially presiphonal
segment; dorsal blood ves.^el slender, not connecting directly with
neurointestinal vessel, which is relatively long and divided into two
before joining ventral vessel; size medium; skin with low papillae.

Description. — Length of type, much contracted, 70 mm.; diameter
25 mm. Skin roughened by low, close-set, pustulate verrucae largest
and most irregular at QWiX^ of body. Setae 2, close to the much
contracted anterior end; 11 mm. long, the inner ends united b}^ an
interbasal muscle which docs not pass through loop of neurointestinal
vessel,

Coelomic surface of body widl perfectly smooth and with satiny
sheen, the muscles continuous, without trace of difTerentiation into
670329 — 1<; :;

248 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9«

longitudinal or oblique bands. Where the seta muscles join body waU
there are a few transverse thickenings of inner layer.

Nephridia 2, inserted close to nerve and posterior to seta about
the length of latter. In the only female dissected these nephridia
are four-fifths length of body and contain numerous eggs. In the
male they are about one-fifth length of body and contain sperm.
The internal opening (nephrostome) is on a short peduncle near base,
with an irregular small lip lacking any trace of spiral structure.

Anal vesicles 2, simple, thin-walled, opening into small cloaca hav-
ing longitudinally pKcate walls. Minute ciliated funnels are scattered
over the surface.

Alimentaiy canal excessively long (600 mm.) with very numerous
coils attached to body wall by a multitude of very delicate frenula
in which is entangled coagulum containing numerous eggs and brown
bodies of unknown nature. The pharynx-esophagus is thin-walled.
Plate 25, figure 3, shows the anterior complex more or less in situ
and figure 4 with the interbasal setae muscle cut and the pharynx-
esophagus pulled to right. The gizzard is about 6 mm. long, but the
length of the stomach cannot be determined on account of condition
of material. The interval between gizzard and beginning of siphon
is the astonishing distance of 170 mm. The siphon accompanies the
following 240 mm. of intestine, while the terminal, postsiphonal por-
tion is 190 mm. There is no intestinal coecum in front of the cloaca.

Vascular system. The dorsal vessel can be traced posteriorly
nearly to the point where the long neurointestinal connective (B^) is
attached to the lower side of the alimentary canal. The relation is
similar to that of Bonellia. Anteriorly the neurointestinal connec-
tion divides into two branches before joining the ventral vessel.
There is therefore no enlarged "heart" at the posterior end of dorsal
vessel, nor a ring vessel embracing the gut at that point.

T^/pe.— U.S.N.M. No. 20615.

Type locality. — Monterey Bay, Calif., 35-40 fathoms, mud, De-
cember 3, 1931, 3 specimens.

Other material examined. — Off southern California (13K miles south
of Seal Beach), 215-225 fathoms, green mud, Allan Hancock Foun-
dation, 1 specimen.

Remarks. — The specimens are in a poor state of preservation, the
alimentary canal being soft.

The outstanding features of inamoenus are the two nephridia, the
internal aperture of which does not have spirally twisted lips; the ex-
traordinarily long presiphonal gut, and the very long neurointestinal
connective not embracing the interbasal muscle of setae. The vas-
cular system lacks a definite intestinal ring vessel — probably a generic
feature. A. inamoenus differs from the only other recognized species,
arhynchite of northern Japan, in respect to the neurointestinal vessel

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 249

already mentionod, and in the probably much longer presiphonal gut.
Details of the gut are lacking in Sato's figure and des('rii)tion.

Family BONELLIIDAE Baird

Bonellidae Baird (name only), 18G8, p. 111. (Includes Thalassema and Bonellia.)

Diagnosis. — Dimorphic echiuroids. The male is degenerate, pla-
narianlike, with ciliated ectoderm, generally one, exceptionally two,
nephritlia '° serving as sperm receptacles, and a vestigial alimentary
canal; it lives semiparasitically or parasitically on or in the female,"
and is sometimes absent from female; females resembling Thalassema
but with bifid proboscis in some genera; two, four, or exceptionally
many anterior ventral setae are sometimes present; no anal setae;
anal vesicles with many branches ending in ciliated cups; one to three
nephridia.

KEY TO GENERA OF BONELLIIDAE

a'. With an elongate proboscis bifid at the end.

6'. With ventral setae " or hooks a short distance behind mouth,
c*. Regularly one nephridium or egg receptacle (either right or left).

d'. Coelomic aperture of nephridium (i. e., the nephrostome) situated
near base of the organ, usually at end of a short lateral tube.

Bonellia Rolando
<P. Large nephrostome at extreme distal end of nephridium and not

facing laterally Bonelliopsis, new genus (p. 252)

c'. Regularly 2 nephridia, having the small nephrostome laterally near
distal end; a small blind tube opening between nephridiopores serving
as a permanent androecium for completely parasitic male; gonad of
female situated on frenula radiating from cloaca; anal vesicles in form
of tubules opening independently into cloaca (see also Acanthoha-
mingia) Pseudobonellia Johnston and Ticgs '*

'• Males of Pteudobontllia have two ncphri'lin; other penera one only.

" The male lives In the foreput. In nephridiuni, on proximal portion of proboscis, in ponital proovc (/lean-
thohamingia) . anrl In a si>cciallzed blind tube or androecium opening between the two nephridiopores {Pteu-
dobontllia ) .

'« In limellia miyniimai Ikeda (lOW, p. 73; 1907, p. 2, pi. 1, rtps. 1, 2; pi. 2, flps. 5-17) there are numerous
(29) very small setae. Ptendobonellia has two to four sctiic; Archiboneltia has four and other pcnora have two
Mtac, but sometimes o com piemen t/il smaller seta Is In process of prowth and liitcr replaces the fimrtlonal ono.

'« John.ston and Tleps, 1910, pp. 21.V229, pis. »-ll. This is a very remarkable penus set ai)art from all other
boncllilds in havlnp a small blind tube, projectinp into coelom and openinp on the ventral body wall between
the two nephridloiwrcs by a narrow ainal whose walls contain stronp sphincter fibers. In this lives per-
manently one very depenernto male. Its posterior end prown fast by onlarped ectodermal colls to the much
smaller epithelial cells of the tubule. The male lacks setae and has two sperm receptacles, dlfTerinp from all
other known males (which have only one). The ovary Is entirely dilTerent from that of other penera. "The
mesenteric strands of mu.sculnr tl.ssue which nmlntain the iK)sterlor portion of the rectum in position arc very
well dcvclopwl and form the basis of the ovary whilst from the ix-ritoneum llnlnp them the ova are developed"
(p. 221). "The anal pinnds or pf»sterior nephridia are ropre,scnt/Ml by two small, tuft-like masses slliiate<l one
on each side of the posterior end of Intestine. Each consists of n mass of very delicate, simple, cylindrical
tubosopenlnpseparately Into the rectum, whose walls In thisreplon are thickened. The tubules arc approxl-
mat/'ly clrailar in section with an Irrepular lumen. They consist of a sinple layer of epithelial cells. Near Its
free end each tubule t)«comes narrowed before o[K>ninR Into the ooclom by a sllphtly dilated funnel fringed
with long cilia" (p. 220).

This species was collected In the Capricorn Group, Oreat Barrier Uoef, due cast from Kcppol Bay, Qaoens-
land. Subsequently Monro (1931, p. 33) reported the species from Low Isloa.

250 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

c^. Three nephridia, an unpaired between a pair; ovary along nerve cord;
anal vesicles: a fascicle of tubules at end of a collecting tube.

Archibonellia Fischer "
6^. No setae present.

c*. Nephrostome situated at end of a short lateral tube near blind distal end

of nephridium Parabonellia Onoda

c^. Distal end of nephridium expanded into plicated rim of large nephro-
stome Eubonellia, new genus (p. 255)

a^. Proboscis when present similar to that of Thalassema and not bifid at extremity.
6'. No sharply marked groove between nephridiopore and mouth.

c*. Female with 2 well-developed ventral setae; anal vesicles 2, elongate,
dendritic; nephrostome near base of the single nephridium; male un-
known Protobonellia Ikeda

c^. Female with 2 ventral setae; nephridia 2, large, with basal 2-lipped
nephrostome; anal glands wide sacs provided with a very large number
of slender excretory tubules with apical funnel; male unknown.

Maxmiilleria Bock
c^. No ventral setae in female.

rfi. Proboscis deciduous, Thalassema-like; 2 external papillae marking
nephridiopores; nephridia 2 or 1 with basal nephrostome; anal
vesicles in 2 thick clusters of tubules opening into a common duct;

male with ventral hooks Hamingia Danielssen and Koren

c*. In place of proboscis a short truncate snout; a proboscis possibly normally
present; 1 nephridium with basal nephrostome; duct of nephridium
passing under nerve cord and opening in median line into a funnel-
shaped depression of skin; anal vesicles; numerous dendritic masses
arising from a basal bladder on each side of large muscular cloaca;
terminal portion of hind-gut, in front of cloaca, greatly enlarged; male

unknown Nellobia, new genus (p. 257)

^2. A narrow, or expanded, slit extending forward from nephridiopore, in 2
species containing 8-10 tiny, integumentary spines; anal vesicles not in
form of 2 dendritic structures or 2 clusters of tubules, but in form of
independent branched tubes or an asymmetrical cluster; 1 or 2 nephridia
with nephrostome near base; males with or without hooks.

Acanthoharaingia Ikeda (p. 260)

n Wilhelm Fischer, 1919, p. 283, figs. 5, 6; 1926b, p. 207, pi. 2, figs. 1-7, text figs. 1, 2. Archibonellia michael-
soni, the type, from a coral reef, Rottnest Island, Fremantle, Australia, is only 12 mm. long; it has a large
median between and above two small nephridia (no nephrostomes were found). The proboscis has two
terminal lappets instead of divisions, and the alimentary canal is short, scarcely over twice length of body.
A. mj6bergi (1926b, p. 208, fig. 1. northwest coast of Australia), however, has a very small unpaired nephridium
(with basal nephrostome) between a pair of very large "uteri" filled with eggs and with terminal nephro-
stome. The alimentary canal is of normal length for a bonelliid, and the proboscis is normally cleft at the
tip. Fischer, in a quandry what to do with this species, places it in Pseudobonellia on the basis of the large
paired nephridia, with terminal nephrostomes, and the normal alimentary canal and proboscis. Even though
the androecium of Pseudobonellia may well have originated in an unpaired nephridium, the fact remains
that in Pseudobonellia the androecium is highly specialized and is no longer a nephridium, while the ovary is
not found along the nerve cord, as in Archibonellia, and the tubes of the anal vesicles open separately into the
cloaca, not into a common duct. The two species of ^rcftiftoneHia may not be congeneric, but the aberrant
species is certainly widely different from Pseudobonellia.

Figure IS. — Bonellia viridis: Dissection (X 5) of anterior part of specimen from Naples
to show particularly the position of nephrostome, CF, at base of nephridium, N, which was
55 mm. long while the animal was only 45 mm. Note the long segment of gut between
gizzard, G, and beginning of siphon, Sil. The anterior end of gonad, Go, Is shown. {B ^,
B^, B*, dorsal, neurointestinal, and ventral blood vessels; C, stomach; G, gizzard; NC,
nerve cord; 0, esophagus; P, pharynx.)

ECHIUHOID WORMS OF NORTH PACIFIC— FISHER 251

Figure IS.— (See opposite page for legend}.

252

PROCEEDINGS OF THE NATIONAL MUSEUM

BONELLIOPSIS, new genus

Diagnosis. — Differing from Bonellia in having the nephrostome at
the distal end of the nephridium instead of near the base, and in the
possession of a much shorter presiphonal foregut; either left or right
nephridium developed; females with two ventral setae; males without
setae.

Type, Bonelliopsis alaskana, new species.

BONELUOPSIS ALASKANA, new species

Figure 16; Plates 26, 27

Description. — Body elongate, subcylindrical, blunt at both ends,
20 to 65 mm. long, and commonly four or five times as long as thick;
proboscis of usual Bonellia form seldom exceeding body length and
usually considerably shorter. Contracted skin verrucose, the verru-
cae squarish, not obviously larger in any particular region; when skin
is stretched the verrucae flatten out into squarish glandular thicken-
ings arranged in irregular longiseries.

Figure 16. — Bonelliopsis alaskana, new species: Two males, the upper 1.27 mm., the lower
1.9 mm. long, X SO. The spermatheca is most heavily shaded; the lightly stippled is
parenchymatous and muscle tissue; sperm duct opens at anterior (right) end.

Body wall thin, translucent. Inner, circular layer of muscles
smooth, but in the region of foregut where the layer is thickest there
is a division into slender fascicles.

Setae 2, small, nearly straight, situated close together a short dis-
tance behind mouth (4 mm. in specimen 44 mm. long). There is a
short but broad interbasal muscle which usually presses upon the
nerve cord and ventral blood vessel.

Nepliridium 1; of six specimens dissected four had the left ne-
phridium developed and two had the right. It is situated close to the
nerve cord directly behind the setae. In some specimens the nephri-
diopore is conspicuous externally. The nephrostome is conspicuous,
terminal, with amply folded lips. When the nephridium is filled

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 253

with eggs its tlistal oiul is usually iuvaginatt'd, coiicoaliiig the iicpliro-
stomo (pi. 27, fig. 1). The mucosa is thrown into shallow longitudinal
folds, most pronounced at proximal end.

The two anal vesicles have the relative size and general form in-
dicated in plate 26, figure 1, where only one is shown. Each opens
by a small pore into the cloaca. The primary branches of the vesicle
vary in number. In plate 2G, figure 2, is shown the tip of one of the
major subdivisions including two of the smaller secondary branches,
each of which bears several fumiels.

Ali.ment.\ky c.\nal. As contrasted with Bonellia viridis the ali-
mentary canal differs in having a much shorter forcgut, especially the
portion between the gizzard and the intestine, corresponding in general
to the "stomach" of Thalassema and allies. The abruptly enlarged
intestine is produced forward into a coecum where the dorsal blood
vessel envelops the intestinal wall. In the specimen of B. viridis that
I dissected this was not differentiated (fig. 15).

The pharynx is connected with the body wall by nmnerous radiating
muscular frenula of which only a few are indicated in the figures.
The mucosa is thick, verrucose, and not very different from that of
the esophagus. The gizzard has stronger ring muscles than the
esophagus (which shows a ringed structure), and the mucosa is
thrown into consecutive ring folds. \Vhen spread out these ring
folds subdivide into eight longitudinal divisions. Behind the gizzard
the mucosa becomes al)ruptly tiiinncr, and between the gizzard and
this elongated stomach there is a sort of pjdoric constriction, with a
very narrow passage. The opening, on the stomach side, is surrounded
by a fiange of tissue. There is no presiphonal ciliated groove, but a
postsiphonal one runs along the intestine to the point where the
ventral blood vessel and genital stolon join the hind-gut. The
fecal pellets are slender blunt ellipsoids 2.5 mm. long. In one I found
an ostracod and a small Balanus.

Mesenteries. The mesenteries are continuous, very thin sheets,
even in the postsiphonal region of the intestine. In plate 27, figure 2,
an attempt has been made to show the principal mesenteries of the
foregut, which are voluminous and folded when the animal is contracted.
In this sernidiagrammatic drawing the left half of the animal only is
slunvn. Wliat is probably the ventral mesentery (VAf) is attachcnl
to body wall over a sinuous and not so smoothly regular course as
shown in figure. It is fastened to the l(>ft sid(> of pharynx-esophagus,
passing gradually to the side of the intestine oj)posite the siphon.
The posterior part of dorsal blood vessel (/?') is attached to, or in-
volved in, this mesentery. The dorsal mesentery (DAf) is attached
to the body wall in a long spiral passing from right siile (dot and dash
line) over to left side. Posteriorly it merges with the ventral mescn-

254 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

tery on its left side, where the foregut enters the intestine. In the
drawing the posterior face of dorsal mesentery is shown and only a
small part of the anterior (or dorsal) side adjacent to pharynx. The
attachment to the gut is along the lateral line, but behind gizzard the
attachment moves over near to that of ventral mesentery. A special
muscular mesentery indicated in plate 27, figure 1, but omitted from
figure 2, joins the gizzard to the stomach. A special transverse dorsal
muscular mesentery supports the dorsal blood vessel anteriorly.
When the animal is extended the effect of these mesenteries is to form
a somewhat spiral anterior cul-de-sac in which the nephridium lies
and in which the eggs probably congregate. But when the neplnidium
is full of eggs, as in plate 27, figure 1, there may still be hundreds of
eggs, seemingly mature, in various parts of the coelom.

Vascular system. The distribution of the principal trunks,
shown in the figures, seems to be almost the same as in Bonellia
viridis (fig. 15). The connection between the dorsal and ventral
vessels is not direct, as in Echiurus, Ochetostoma, and Thalassema,
but by means of lacunae in the intestinal wall, as is the case in
Arhynchite.

The gonad lying along the top of the ventral blood vessel, in the
posterior half of body, seems to be identical with that of B. viridis.

Male. The males are fomid in the foregut, from the pharynx to
the gizzard, in the order of about a dozen to an individual. In one
case I found two or three in the anterior part of the stomach. Dr. H.
Heath, who collected the type series, examined several live specimens
and found no males in the nephridium.

The males vary from elongate-slender to the shortened state shown
in figure 16. When fully extended they are more than twice as long
as figure and only about half as thick. The sperm receptacle is
situated posterior to middle of body and the duct opens at anterior
end, or very close to it. There is very little free coelom, the body
being filled with parenchymatous and muscle tissue, diagrammatically
indicated by the more spaced dots in figure. No setae are present.

Type.— U.S. ^M. No. 20603.

Type locality. — Dutch Harbor, Unalaska, under rocks, at low
tide; Harold Heath, August 1917, 15 specimens.

Specimens examined. — In addition to the above, 25 specimens,
without locality, U. S. National Museum.

Remarks. — The type series was collected by Dr. Harold Heath,
who found them in the intertidal zone under flat rocks. The worms
were arranged around the periphery of the stone with the proboscis
extended to the margin. In life the animals are light green, the
color of "green prunes," which, in the contracted state, they some-
what resemble.

ECHIUKOID WORMS OF NORTH PACIFIC FISHER 255

EUBONELLIA, new genus

Diagnosis. — Bonclliidae with woll-dovolopcd l)ifurcate prol)oscis,
110 setae, and a single (right) nephricUuni, the iHstal end of which is
expanch^d mto the plicated rim of the large nephrostome; anal vesicles
essentially as in Bonellia; male with sperm receptacle reaching nearly
to posterior end of body; body wall thick.

Type, Eubonellia valida, new species.

Rrmarhs. — Tliis genus differs from Bonelliopsis, the only other
known to have a strictly terminal nephrostome, in lackuig setae.
Its foregut has an unusually extensive stomach or crop, between the
gizzard and begnunng of siphon, this segment being short in Bonelli-
opsis. The mesenteries of the mtestine of Eubonellia are m the form
of strands, not a sheet.

Parabonellia Onoda, '* based on Bonellia misakiensis Ikeda,'^ also
lacks setae in both male and female. The nephridium has a small
pedunculate funnel situated, not at the base as in Bonellia, but on the
side of the vesicle near the terminal blind end. Ikcda did not show
this in his figure of a "dissection" of the type, but it is figured by
Onoda (fig. 1) and Sato '^ (fig. 2). None of these writers has figured or
described significant details of the foregut, which are of importance in
classification.

The body wall of Parabonellia is thin whUe that of Eubonellia is
very muscular, but this is a character varying with the contraction of
the specimen and is scarcely of generic significance.

In Eubonellia the male is about half the size of the male of Para-
bonellia, which is described as nematodelike, 3.3 to 3.5 mm. long, and
0.2 to 0.3 mm. in breadth. The sperm vesicle and canal occupy the
anterior fifth to third of the body. In Eubonellia the vesicle extends
nearly to the posterior end of the bodv, which is depressed, planarian-
like.

Profoboncllia Ikcda '* (type, P. mitsukurii, Sagami Bay, Japan, 300
fathoms) has very strong setae with unusually well developed muscles.
The proboscis is similar to that of ThahisKema, and the single (left)
nephridium is similar to that of Bonellia. The nephrostome is a wide
fimbriated funnel at the end of a slender tube opening into the nephrid-
ium near the base of that organ. The anal vesicles aic similar to
those of Bonellia rather than Ilamingia, which the proboscis might
lead one to expect. The vascular system seems to possess a peculiarity
in a rather direct connection between the dorsal vessel and the neuro-
intestinal connective at a point immediately anterior (o the be-
giiming of the siphon. Ikeda treats this very summarily in both

"• On<xln 1«H, p. 418 (PtnidofiontlUa); 1035. p. HI (.Parabonfllia for PteudobontUia, prcocciiple<l).
'• Ik<yU, 1W>I, p. 74, flKS. 24, 103-105.
" ."^alo, 1935, p. 142, Ors. 1,2.
'• Ikcda. 1808*. p. 259. figs. 1-4.

256 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

description and figure, apparently not realizing that it is different
from the relations of the two vessels in Bonellia. Neither his figure
nor description gives any definite details concerning the foregut
beyond the fact that it is very long.

EUBONELLIA VALIDA. new species

Plate 28

Description. — The much-contracted type is oblong-cylindrical, 55
mm. long, and about 20 mm. thick at middle. The conspicuous
nephridiopore is 8 mm. behind mouth. Proboscis unusually broad
and flat, terminally bifurcate, without a ventral sulcus. In its con-
tracted state it is 30 mm. long and 7-9 mm. broad (pi. 28, fig. 3).
The thick skin is thrown into annular welts, the furrows being fre-
quently interrupted. These folds do not carry marked pustulate
thickenings as in the case of Nellobia found at the same station.
From the mouth a short narrow sulcus extends upon the constricted
base of proboscis.

Body wall tough and muscular, 1.5 to 2.5 mm. thick, the middle
longitudinal layer being the thickest. The inner, circular layer is
smooth, but somewhat fasciculated at anterior end of body.

No setae or vestiges of seta sacs or muscles.

The single large (right) nepln-idium has a terminal large nephrostome
with voluminous lips. At the base a simple duct leads to the exterior
approximately in the median line.

The two anal vesicles are of the elongate dendritic type with a
voluminous axial bladder having a few branches proximally. To the
main stem and branches are attached singly or in clusters (pi. 28,
fig. 2) the nephritic elements, which are characteristically very elongate,
ending in pedunculate funnels. The vesicles are attached to each side
of a very small cloacal bulb, the mucosa of which is thrown into longi-
tudinal ridges.

Alimentary canal. The anterior part of pharynx is attached to
body wall by numerous frenula, but there is no peripharyngeal dia-
phragm such as is characteristic of Nellobia. There is a rather delicate
mesentery at the bend of the esophagus (not shown in figure). Food
pellets form in the posterior part of esophagus (0^), as well as in the
gizzard (60, and are collected in the elongate stomach. The wall of
the stomach is almost transparent. Between it and the gizzard there
is a powerful sphincter forming a sort of pylorus as in Bonelliopsis.
There is a weaker sphincter between the esophagus and gizzard.

There is no presiphonal ciliated groove. Much of the intestine is
missing, but enough remains to show that there is an extensive por-
tion, traversed by the siphon, which has thicker walls so that the
pellets are not visible, whereas the postsiphonal intestine (with a

ECHIIHOID WOKMS OF NORTH PACIFIC — FISHER 257

ciliated grroove) has very thin walls and very compact pellets (j)!. 28,
fi^. 4). The ciliated groove entls at entrance to small cloaca! bulb and
a strand of tissue from the ventral blood vessel is attached at this
point.

The vascular system is of the usual bonelliid type and can be fol-
lowed in the fi«rure. The ventral vessel divides into two at about the
point where setae would be if present.

The ^ronad, in the usual bonelliid position, is confined to the median
third of the body and is inactive, there being: no sig:n of egg formation.
In Xdlobia, dretlged at the same time, the gonads were active and the
nephridium full of eggs.

Male (pi. 28, fig. 5). One was found in the pharynx near mouth.
It is possibly not fully matured; length 1.17 mm. The dark body is
the spenn vesicle, the duct opening at the anterior end as in Bonellia
nridis. The small spot back of the gonad is probably the excretory
pore. Small indistinct masses of spermatozoa can be seen in the
coelom, but have not been shown in the drawing. The gonad is situ-
ated more posteriorly than in Bonellia viridis.

Type—V.S.K.M. No. 20604.

Type locality. — Albatross station 5021, Okhotsk Sea, off east coast
of Sakhalin Island, lat. 48° 32' 30'' N., long. 145° 08' 45" E., 73
fathoms, green mud, sand, pebbles, bottom temperature 30.9° F.

NELLOBIA, '» new genus

Diagnosis. — Bonelliidae without setae and possibly without probos-
cis; one nephridium (.left), with a basal ncphrostome and swollen
basal region, opening in the median line but without a genital groove;
two compoand anal vesicles each consisting of numerous trees aris-
ing from a sessile receptacle on either side of the very large muscular
cloaca; terminal portion of hind-gut greatly enlarged; body wall very
thick.

Tvj)e, Nellobia eusoma, new species.

Remarks. — This genus differs from Bonellia, Protobonellia, Para-
bonellia, Eubonellia, and BoncUiopsis in the radically different struc-
tun; of the aral vesicles and from all bonelliids in the extremely large
muscular cloaca and enlarged terminal part of the hind-gut. The
only group which hns fundamentally similar anal vcsi<'les is Acan-
thohamln(}ia Ikeda ^ in which three trees arise indei)(!ndently on each
side of the cloaca {A. ijimai) or more ninnerously as o cluster with
probiibly some coiuiectif)n between the elements {A. shiplei). In
Ilaminf/ia arctica Danielssen and Koren *' there is a very short (1 mm.)

" Anneram of litmf.Uia.

'^ Ikpiln, lOlO, p. l.Tfi, 1)1. 10; !kH> also lOOH, p. 01, pi. I.

•1 Danlflsson an<l Korfn, IhHI, p. '.!n, [il 4 uml .'.. lU'". 1-1H.

258 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

tube on each side of the cloaca to which very numerous small trees
are attached at approximately the same point.

The vascular system of Nellobia is similar to that of Bonellia and
allies in the relation between the dorsal blood vessel and the neuro-
intestinal trunk. No direct connection exists between the two by
obvious anastomosis of terminal branches such as Danielssen and
Koren figure (1881, pi. 5, fig. 14).

NELLOBIA EUSOMA, new species

Plates 29, 30

Descrijption. — The single specimen is contracted to the maximum
extent. The posterior end of the body is invaginated to form a cup-
shaped depression . The intestine had been extruded, after the manner
of holothurians, through a breach in the cloacal wall, and most of it
is missing.

Body of ^OTigZZm form without proboscis, 44 mm. long (allowing for
posterior invagination), and 15-17 mm. thick at middle. Owing to
contraction, the sldn is thrown into irregular transverse folds with
frequent pustulate thickenings, less regular in the anterior ventral
region (shown in pi. 29, fig. 1) than elsewhere. If the very short
truncate snout is the remains of a longer proboscis, it is nevertheless
covered with normal skin. The conspicuous opening of the nephrid-
ium is close to the median line about 4 mm. behind mouth.

The body wall is very muscular and in the contracted state about
2 mm. thick. The middle longitudinal layer is the thickest, the
inner circular layer the thinnest. The latter in the contracted state
of the specimen shows definite fascicles of uneven width which would
probably smooth out when the worm is expanded.

The single neplmdium (pi. 30) is attached on the left of the nerve
cord, but its duct passes under the cord to open in the median line.
The nephrostome has very simple lips and passes into a bulbous and
thick-walled proximal region. The distal compartment, filled with
eggs, has the wall stretched to translucent thimiess but its proxunal
constricted part has glandular walls furrowed longitudinally. There
is a definite opening from the egg chamber into the proximal bulbous
portion.

The anal vesicles are peculiar. Instead of having a roughly treelike
foi-m as in Bonellia and Bonelliopsis, the main vesicle is a sort of
crescent-shaped pouch applied to each side of the large cloacal cavity
and produced on the opposite or free border into numerous (a dozen
or more) unequal dendritic subdivisions. The larger of these have a
few main branches lilve a tree, which in turn are crowded with branch-
lets (pi. 29, fig. 4) carrying many of the bulbous glandular elements
ending each in a ciliated funnel. Around the base of these primary

EClilUKOID WORMS OF NORTH PACIFIC — FISHER 259

iirpliridia aro luiinorous siil)irlol>uIar umuiual yellow bodies arising
from the l)ast' of the iiej)hritie eleiiieiits and from the wall of the collect-
ing tube or branehlet. Xumerous much sni.'dicr brownish-yellow
papillae occur on the walls of the main stems and branches. I
could not find the opening into cloaca.

Alimentary canal. The main features of the foregut are shown
on plate 30. The pharyiLX is attached to body wall by very numerous
crowded ratliating strands. This head cavity is separated from the
rest of coelom by u translucent diaphragm (D) indicated in the drawing
incompletely. Its central l)or(ler encircles the gut behind the pharynx,
and what may be conventionally called the esophagus (extending to
A' in drawing) has the muscular walls marked by prominent ring
folds which cause the mucosa to be thrown into transverse welts.
This muscle layer thins toward end of esophagus and the ring
becomes narrower. The segment X-Y corresponds to the gizzard of
BoncUiopsis. The annulation of muscle is closer. At Y the canal
was broken, and it is possible that something was lost as the segment
between Y and the beginning of siphon is very short. It corresponds
to the so-called stomach of Bonelliopsv?. A tough mesentery unites
loops of the esophagus, whereas in Bonelliopsis the thicker mesentery
joins the gizzard to stomach. The esophagus has a continuous ventral
mesentery attached ventrodextrally but the other mesenteric attach-
ments to body wall are in strands or frenula.

The cloaca is bulbous, with very numerous muscular strands uniting
its rather muscular wall with l)od\' wall. The anterior of these strands
pass between the branches of the anal vesicles. The very expanded
hind-gut seems to be more than an accident of killing, as the condition
of the mucosa indicates that th(> walls have not been unnaturally
distended. The j)rominent ciliated groove continues from the narrow
segment (all the rest of the intestine having been lost) to the beginning
of the cloaea, where a strand from the ventral blood vessel ends.
There is no intestinal coeeum at this point.

The vascular system is of the Bonellia type. The neurointestinal
connective (B^) spreads out fanwise where it joins the ventral vessel
(B*) and its walls a[)[)ear to be glandular as if a i)art of the gonad
complex. Actual ova are found iis far forward as the posterior border
of this fan.

The gonad is of the Bonellia type but extends uiuisually far forward.

Male. Unknown. The foregut was (juite em[)ty, and no males
were found in the nephridiopore.

Typc.—\J.S.N.\[. No. 20005.

Type locality. — Albatross station 5021, Okhotsk Seo, off cast coast
of Sakhalin Island, lat. 48° 32' 30" N., long. 145° 08' 45" E., 73
fathoms, green mud, sand, pebbles, bottom temperature 30.9° F.

260 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

Genus ACANTHOHAMINGIA Ikeda, emended

Acanthohamingia Ikeda, 1910, p. 136. (Type, A. shiplei Ikeda.)

Diagnosis. — Differing from Hamingia in having a well-marked
genital slit extending forward toward mouth from the one or two
nephridiopores, this containing, in two species, 8 or 10 very small
setae imbedded in the skin; anal vesicles numerous, at least not in
two symmetrical clumps which arise from a very short common duct;
body wall thin; proboscis (when known) similar to that of Thalassema;
males with or without ventral setae.

Remarks. — The new species described below has necessitated an
emendation of the original diagnosis since there are no minute setao
in the genital groove. This groove, which extends forward from the
nephridiopore, or pores, occurs in the three known species ^^ and is not
present in any other genus. The general habit of the three species is
much the same, as they are all of delicate build. The body wall is
thin, translucent when expanded, and skin papillae are poorly devel-
oped. The anal trees exhibit differences in the three species, being
most alike in A. ijimai and A. paradola. These are numerous, inde-
pendent, or semi-independent branched tubules, and differ from the
condition in any other genus except Nellobia. But in Nellobia eusoma,
which lacks any trace of a genital groove and is one of the most heavily
built of all bonelliids, the anal trees spring from a bladderlike struc-
ture applied to each side of the very muscular cloaca. The rudi-
mentary bladder figured for A. paradola (left side) may well indicate
the last trace of a similar structure.

In A. shiplei and A. ijimai the male is long and slender and Iscks
setae, whereas in A. paradola the male is lanceolate and planarian-
like and is provided with two curved setae.

ACANTHOHAMINGIA PARADOLA, new species

Plates 31, 32

Diagnosis. — Dijffering from A. shiplei and A. ijimai in the absence
of minute setae from the genital groove of female and in the presence
of a pair of ventral curved setae in the male; nephridia 2 instead of 1;
anal trees numerous, slender, sparsely branched, arising for the most
part independently from the very thin wall of the cloaca. Length of
paratype 90 mm. (pi, 31, fig. 1). Color, pale flesh when seen on a
white background.

Description. — The general habit is much like that of A. ijimai but
proboscis is lacking. The body wall is very thin and translucent,
this thinness being accentuated by inflation. Along the midventral
line the extremely slender nerve cord can be easily seen. The skin

I A. shiplei Ikeda 1910, p. 136, pi. 10; Hamingia ijimai Ikeda, 1908b, p. 62, pi. 1.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 261

is marked by flat circular spaced spots slightly less translucent than
the intervals. Back of the mouth is the characteristic genital groove,
which normally is probably very narrow as in the type (pi. 31, fig. 3)
but in the paratype (pi. 31, figs. 2, 2a) is spread apart owing to
stretching of l)ody wall. In this groove are four males. In the type
there is at least one. At the posterior end of groove are the openings
of the nephridia: two in the type and another specimen (pi. 31, figs.
4, 4(7); one in the paratype (pi. 31, figs. 1, 2). The hemispherical
papilla sho\\Ti in these figures beside the nephridiopore is an artifact.
Although there is but one opening there are two large iiepliridia
containing eggs.

As slated above, the body wall is very thin, on the order of 0.15 to
0.2 mm. thick, and a single thickness is so transparent that printing
can be easily read through it. The fibers of the longitudinal and
circular muscles can be seen under magnification, but there are no
bundles.

All three specimens are in poor condition internally as the midgut
and hind-gut are badly macerated, the contained pellets being adrift
in the coelom. The more essential foregut can be made out with the
associated blood vessels.

The nephridia, in good condition, number two in all three speci-
mens. They have exceedingly thin walls, and the small nephrostome
and its short stalk are situated at the base, close to the swollen ducts
leading to nepliridiopores. In the paratype the nephridia are about
twice as large as those shown on plate 32.

The significant features of the alimentary canal can be seen on
plate 32. The pharynx is much inflated (but probably unnaturally)
and has very thin walls. The esophagus can be traced to 0' where
the fine longitudinal ridges of the mucosa change to equally small
rings. Then follows an exceptionally long segment of the foregut
in which I can find no marked division into gizzard and stomach (or
"crop"). In this the contents are shaped into oblong pellets 1.5 to
2 mm. long. Extremely slender and numerous frcnula connect the
pliarynx to body wall. The continuous ventral mesentery of foregut
is delicate and transparent. The position of the coils of foregut in
figure has no significance as they had mostly broken moorings. All
the foregut is thin-walled.

In connection witli the vascular system the very considerable
length of gut between attachment of dorsal vessel (/?') and ncuroin-
testinal connective {IP) may be noted. The ventral expansion of
IP is considerably farther back in another sj)ecimen. In keejMng with
tiio rest of the animal the blood vessels arc delicate and the ventral
trunk is very inconspicuous. There are numerous opaque noduk'S
on the neurointestinal trunk.

262 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

The nerve cord is the slenderest I have seen in a mature echiuroid,
being only 0.135 mm. in diameter.

The gonad, on the irregular margin of a mesentery dorsal to nerve
cord, is invisible except under high magnification, as the cells are
small and inactive. The whitish eggs in the nephridia are 0.5 to
0.6 mm. in diameter.

Anal vesicles are in the form of numerous slender tubes with spaced
short lateral branches (pi. 31, fig. 6). These tubes are involved in
and fastened to body wall by transparent but strong frenula. In the
type nearly all the tubes of the left side spring from a small irregular
common chamber closely appressed to the transparent wall of the
cloaca, but oh the right side no such common chamber is present.
Here the vesicular tubes arise independently from the cloacal wall.
It is probable that the number of tubes increases with age. The
ultimate subdivisions are not well enough preserved for exact delinea-
tion but they resemble in general those of A. ijimai. Although the
cloaca is not perfect in any of the specimens, that of the type shows a
rather large subspherical chamber with transparent walls joined to the
body wall by a multitude of frenula.

Male. Males occur in the genital groove as shown in plate 31,
figure 2. Probably the groove is normally as in the type, which has
one or two males in it. They are depressed lanceolate in form, 1.2
mm. long, and the tube of the sperm receptacle opens at the middle
of the anterior end, the receptacle itself being about in the middle of
body (pi. 31, fig. 5).

TV^e.— U.S.N.M. No. 20601.

Type locality. — Albatross station 4942, Kagoshima Gulf, Japan, 118
fathoms, brown mud, black specks, bottom temperature 59.8° F.,
2 specimens.

Specimens examined. — The above and 1 specimen (paratype) from
station 4940, same locaHty, 115 fathoms (pi. 31, fig. 1).

Xenopneusta, new order

No blood-vascular system, the coelomic fluid being heavily charged
with large blood corpuscles containing hemoglobin or hemoglobin
and hematin; intestine with terminal portion in front of cloaca en-
larged, thm-walled, functioning as an organ of respiration in connection
with anus and cloaca.

Family URECHIDAE Fisher and MacGinitie, 1928

Diagnosis. — Differing from other Echiuroidea in the absence of a
blood-vascular system, the corpuscles (red or brown in color from
hemoglobin or hemoglobin plus hematin) free in the coelomic
fluid; distal portion of midgut greatly enlarged and in connection
with cloaca serving as a respiratory apparatus; foregut very long,

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 263

includinE: a lonf;: frizzard botwoon an anterior lonp: crop and a posterior
long stoniacli; proboscis reducetl to a scoop-sha})ed ii])per lip.

Genus URECHIS Seitz

Urechis Seitz, 1907, p. 352 (type, Echiiirus chilensis Max Miiller, 1852). — Fisher

and MacGinitik, 1928a, p. 200.
Spiroctetor Skorikov, 1909, p. 77 (typo, Echiurus unicincius von Drasche).

Diagnosis. — Cylindrical or sausage-shaped echiuroids with characters
of family. Body wall is very muscular, consisting of outer and inner
circular layers and middle longitudinal layer, the latter the thickest;
inner layer showing a fasciculate arrangement superficially. In the
region of the posterior pair of nephridia is a zone of compound slime-
net glands lodged in the verrucae of the skin. There are two or three
pairs of nephridia, the basal nephrostome of which has long spirally
coiled ciliated lips for collection of mature germ cells. The two an-
terior setae have a strong interbasal muscle; one ring of curved anal
setae interrupted ventrally. Traversing the coelomic cavity in front
of the anterior setae are paired dorsoventral muscles (pi. 34, fig. 2,
13). The alimentary canal has a definite pattern of attachment to
body wall by muscular mesenteries, difTering in minor details in the
three species (pi. 35, fig. 1). The slender foregut is very extensive,
consisting of pharynx, esophagus, crop (subtended by a powerful
muscular mesentery not attached to body), a long gizzard, and a
stomach attached posteriorly by a strong mesentery. The greater
part of the very long midgut is accompanied by the siphon, which
starts close to distal end of stomach. The ciliated groove of midgut,
which parallels the siphon, extends be3'ond it to the point where the
gut is suddenl}' expanded mto the inflatable respiratory portion. The
external ridge marking the ciliated groove continues along dorsal
side of this expansion, passing distally to the right where it afi'ords
attachment for mesenteries, but there is no groove inside corresj)ond-
ing to it. This inflatable so-called "hind-gut" is equivalent to the
terminal part of the midgut of Kchiurus (wliich is not enlarged).
The only similar abrupt enlargement occurs in Nellobia eusoina. The
true hind-gut or cloacal cavity is separated from the foregoing by a
definite s{)liincter constriction and consists of a thin-walled ant(>rior
liortion and a thicker-walled terminal section willi rugose mucosa.
Verj' numerous frenula attach cloaca to body wall and account for
tlie exj)ansion of cavity l)y which water is inhaled din-ing respiration.
The anal vesicles are voluminous, slender sacs, always deflated, which
open ventrally into terminal portion of cloaca. The glandular walls
are externally rather cauliflowerlike and the entire inner surface is
intricately plicated. The scattered ciliated funnels arc veiy tiny.
The anus is eccentric to the circle of setae, being slightly nearer the
ventral side fpl. 33, fig. 4).

670329 — (6 4

264 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

Remarks. — Three well-known species have a very uniform outer
facies. Although unicinctus of Japan has only two pairs of nephridia,
it is indistinguishable by external features from small examples of
caupo. The details of skin and setae are practically identical. The
alimentary canal, except for minor details of mesenteries, is closely
similar in the two species. Apparently umcindus does not reach the
large size of caupo and chilensis, both of which have a third (posterior)
pair of nephi'idia. The anterior setae of chilensis are blunt and
scarcely tapered while in the other two species they are strongly tapered
and sharp. In chilensis, as compared to caupo, dorsoventral muscle 13
is weaker, the crop and gizzard are longer, and the interval between
end of stomach and beginning of siphon is about three times greater.
The attachment of the anterior end of the respiratory portion of
midgut (pi. 36, figs. 1, 2) presents important differences.

Urechis chilensis (Miiller), synonym U. farcimen (Baird), is found
at Sandy Point, Strait of Magellan. My specimen, taken by the
Hassler Expedition, was kindly donated by the Museum of Com-
parative Zoology.

A fourth species, Urechis novae-zelandiae (Dendy), awaits detailed
investigation (Dendy, 1898; Poche, 1920).

Dr. Carlos E. Porter, of Santiago, Chile, has called attention {in
lift.) to the name Pinuca edulis Claudio Gay (1854, p. 475). Dr. W.
L. Schmitt, to whom Dr. Porter sent the information, had photo-
graphs made of the five pages covering the section on "Sipunculides"
in the "Historia . . . de Chile." Four of these are pages 53-56 of volume
3 published in 1849 and list Sipunculus lagena and S. cylindricus.
Pinuca edulis is described in supplementary volume 8, published in
1854, as follows:

Afiade tomo III, pdg 56. Pinuca edulis. Por haber perdido los ejemplares
que teniamos de este singular Sipunculiano, es precise d lo menos senalarlo & la
atencion de los naturalistas y viajeros. Segun nuestro diario es de un bianco
pardusco siicio y tiene de dos d. tres pulgadas de largo y como una de ancho. Su
cuerpo es subcilfndrico, ligeramente hinchado en el medio y adelgazado en anabas
puntas, siendo la anterior mucho mas obtusa que la posterior. El cuero es grueso,
coridceo, un tanto arrugado en al traves, 16 que proviene de la reunion de una
infinidad de pentitos mas 6 menos prominentes. La boca es pequena, arugada,
rodeada, d poca distancia, de muy pequenos aguijones apenas visibles, subre-
tractiles y dispuestos en circulo. El ano se halla d la otra extremidad y es
bastante grande, liso, circular y un poco hendido. Un diseiio hecho en el lugar,
senala hdcia el medio una reunion de pequenos cuerpos dispuestos en una banda
circular de una linea poco mas 6 menos de ancho.

Este animal que los habitantes comen cocido en la brasa despues de haberle
quitado las dos extremidades, se halla en las arenas de la isla de Chiloe cerca de
Castro, etc. En mi diario hallo notado que hace el pasaje de los Priapos d los
Sipunculos.

Without specimens from Chilo6 Island it is not possible to determine
whether Pinuca edulis is the same animal as Urechis chilensis, which
has not been reported so far north. If Pinuca is an Urechis it is

FXIIIUHOID WORMS OF NORTH PACIFIC — FISHER 265

obvious that Gay has the mouth and anus confused; and the h'ngth of
2 or 3 inches is small for the average size. There are other discrep-
ancies. In a Lrcchis 3 inches long, the anal setae are not "scarcely
visible" but are conspicuous; no mention is made of the prominent
anterior setae; no specimen of Urcchis displays "hdcia el medio inia
reunion de pequefios cuerpos dispuestos en una banda circular de una
linea poco 6 menos de ancho," whether "medio" refers to the body
or to the anus (mouth), which just precedes this sentence in the
description.

The curious respiratory mechanism of Urechis, in connection with
the loss of its blood vessels and the complexity of its blood physiology,
relegates the genus to a very isolated position. Not less important is
the behavior pattern whereby the requirements of respiration and
food are beautifully met and coordinated. At every point specializa-
tion of habit is matched by structural and physiological adjustment of
the most delicate and efficient description. Along with this specializa-
tion and complexity is an amazing viability best expressed by the
term "tough." ^3

All signs point to Urechis as being the last of a very ancient stock,
one that maj' have flowered mto many species during Paleozoic times.
It belongs to the honorable company of Lingula and those other
aristocrats sometimes refeiTcd to as "living fossils."

UnECHIS CAUPO Fteher and MacGlnitie

F1GCHE8 17-19; Plates 33-35; Plate 36, Figures 2, 4; Plate 37

Echiunia sp. Joh.nson and Snook, 1927, p. 178, fig. 153.

Urechis caupo Fisher and MacGinitie, 1928a, p. 200, pi. 9, figs. 1-6; 1928b,

p. 204, figs. 1-3, pi. 10. — Baumbercer and Michaelis, 1931, p. 417. —

Reufield and Florkin, 1931, p. 185.— Hall, 1931, p. 400.— Sato, 1931, p.

178.— Newby, 1932, p. 387; 1940; 1941, p. 303.— MacGinitie, 1935a, p. 341;

1935b, p. 002; 1935c, p. 483; 1938, j). 208.

Description. — The species reaches a large size. One specimen col-
lected by G. E. MacGinitie at Humboldt Bay, Calif., measures 470
nmi. long by 55 mm. thick. The largest specnnen from Elkhorn
Slough, Monterey Bay, Calif., was 500 mm. long when fully relaxed
in anesthesia, but after preservation it shrank to 375 mm. in length
i>y 35 mm. in diameter. These were undoubtedly very old individuals.
Average specimens are 150 to 180 mm. long.

The surface of the body is traversed by fine irregular channels giv-
ing a rugose api)earance, which is most pronounced in the head region
anterior to the zone of slime glands. The latter, a sort of clitellum,
is usually distinguishable by the circular trend of its fine furrows.
Its anterior border coijicides roughly with the second j)air of nephiidia,

n Sc« Rc'indfl and Florkin, 1031; nniimNrttcr nnd MIcharlLt, 1011; Hall, 1931; MncOlriitip, 103.15, 1935c;
Newhy, IWO. IWl; Fisher and MncOlnltlo, 192«a.

266

PROCEEDINGS OF THE NATIONAL MUSEUM

while the posterior border is spaced behind the third pair a distance
about equal to interval between second and third pairs. This zone
contains the slime-net glands. ''Three to 10 or even more glands
are located on the outer surface and in the outer part of the sides of
the papillae. These glands develop from the surface epithelium
and are formed by an invagination of this layer of cells and the out-
lying cuticle. The gland cells develop numerous terminal cones
which penetrate into the cuticle of the duct (formed by invagination) .
During the secretion of any one slime-net only about one-eighth of
the gland cells are active. In these cells the terminal cones perforate
the cuticle and open into the duct as minute tubules from which the
fibrous secretion which forms the slime-net is discharged. The net

Figure 17. — Urechis caupo Fisher and MacGintie: A vertical section of a mature slime-
net gland taken while actively secreting; drawing by W. W. Newby (1941).

ECHIUKOID WUKMS 01" XOKTH PACIFIC — FISHER

267

itself is i)robal)ly fibrous in iiaturo, althoutrli this lias not been demon-
strated. (Figs. 17. IS.l

"The body wall consists of the cutich*, the surface epitheliinn or
epidermis, and the underlyin<; connective tissue called the cutis
(Jameson, 1S99. p. 572) or corium (Seitz. 1907, p. 326). Tliese three
layers constitute the skin. Beneath tlicni are the outer circular,
the longitudinal, the inner circular muscle layers and the parietal
peritoneum." -*

inactive cell-/-

active

cell • ■ /

Figure 18. — Urechis caupo Fisher and MacGinitic: A vertical section of the middle part
of a slimc-nct gland taken while actively secreting; drawing by W. W. Ncwby (1V41).

Anterior setae terminally tapered, sharp, curvetl, situated, in large
specimens, 3 to 5 mm. !)ack of the groove leading to moutli and about
the same distanec apart. They are metallic yellow, brownish at tip,
8.5 to 10.5 mm. long. The flattened, curved exserted portion is more
tapered and sharper than in chilcnsvi. A strong interbasal muscle is

** Npwl.y, IMI, pp. 304. 3I.^; flRS. 1-10. Dr. Ncwby has subJprto<l the sllmo RlnnMs to a lliorouKh histo
loKlcnl <itu'ly. He has kindly contributed tho orlRtnal drnwinKS of ORurct 17 and IN, which arc from the
alKive paper.

268 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

present, and numerous somewhat variable muscles radiate from the
coelomic end. Inside the seta sac a short substitute seta is often
present close beside the functional one.

Anal setae curved terminally, sharp, 10 or 11 in number, and the
dorsal are longer (8.5 mm.) than the ventral (7.3 mm.). The dorsal
are the only ones used to any extent in cleaning out the burrow.
When digging, the posterior end of the body is bent sharply forward,
underneath, so that the ventral setae touch the ventral surface of the
worm, while the strongly exserted dorsal bristles scrape the mud
backward as the body is again straightened. This habit helps to
explain the absence of a midventral seta, there being a broader gap
at that point. The anus is eccentric to the circle of setae, being
nearer to the ventral side (center of anus 6 or 6.5 mm. from dorsal
setae and 4 or 4.5 mm. from the ventral). All bristles show conspic-
uous cross-banding.

There are three pairs of nephridia varying greatly in size according
to degree of distension with eggs or sperm. In one specimen ex-
amined the posterior tubes were 150 mm. long and 10 mm. in diam-
eter, reaching two-thirds the total length of animal. The anterior
pair is situated close to the setae. Rarely, one nephridium of this
pair is missing. The nephrostome is on the anterior side at the base
and the grooved ciliated lips are very long and spirally coiled. Mac-
Ginitie (1935a) has shown that the superficial groove, V-shape in sec-
tion in the male and more C-shape in the female, communicates by
a slit along its bottom with what is virtually an almost closed duct
or tube underneath. In both upper and deeper parts of groove the
cilia beat toward the nephridium while on the outside of the lips,
bordering the superficial groove, they carry materials in the opposite
direction and incidentally help in the circulation of the coelomic
fluid and contained blood cells. "As the eggs, blood cells, and other
coelomic materials pass along the outside of the thread the eggs are
caught in the external portion of the groove, are fed into the inner
channel and then proceed to the opening leading into the storage
reservoir [nephridium]. They are carried toward the reservoir at the
approximate rate of 7 spirals per minute. Thus the eggs are sepa-
rated from all other coelomic materials. As the eggs pass through
the slit between the external portion of the groove to the inner chan-
nel, they are under considerable pressure. They enter as a wedge,
then become disc-shaped in the slit, and finally round out in the in-
ner channel. Only mature eggs with indentations are collected in
this way. As the eggs pass along the inner groove toward the storage
organ they become oriented with the convex surface of one egg
pushed into the indentation of the egg ahead, thus forming a com-
pact chain. Blood cells (which range from 0.014 to 0.02 mm. in di-
ameter) and immature egg cells pass along the collecting threads (i. e.,

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 2G9

tho spiral lips) without lodc:iii<]: in the extornal groove. Sperm is col-
lected ill the outer groove of male collecting threads, fed into the
iimer groove, and carried to the storage organ." "

There is no permanent gonad. "Sex cells in all stages of develop-
ment, from very immature ones to those wliich appear to be fully
mature ore found in the coelomic fluid at all times of the year. In
the case of the male, the apparently mature, free-floating sex cells are
known to be functional." (MacGinitie, 1935c, p. 485.) "I have
examined a male collected in winter as well as several specimens of
both sexes collected in the summer and in these I could establish
neither qualitative nor quantitative dilFerenccs in the sex cells at tho
two seasons. Furthermore, in neither season did I find mitotic fig-
ures in any area of the peritoneum nor could I establish any evidence
of division by any of the cells which were free in the coelom. Thus
there is no evidence in regard to the origin of the sex cells in Urechis."
(Xewby, 1940, p. 7.)

MacGinitie (1935a, p. 342) estimates that there are nearly 3 billion
sperms present in the nephridia of an average-sized male and over 6
million eggs in an equal-sized female. The eggs are 0.115 to 0.12 mm.
in diameter according to MacGinitie (southern California specimens).
C. V. Taylor measured 303 from the Monterey Bay region and found
them to range between 0.123 and 0.144 mm. in diameter.^* The egg is
ver}' clear, with a large nucleus, containing a nucleolus 0.012 to 0.016
mm. in diameter.

The anal vesicles, contracted, have a cauliflower surface beset with
minute ciliated funnels. They empty, vcntrally, into the posterior
part of the cloaca.

In addition to abundant sex cells, the coelomic fluid is filled with
nucleated red or brown blood corpuscles subcircular in shape and up-
ward of 0.035 mm. in diameter, together with very numerous amoe-
i>oid cells, yellowi.sh when aggregated. "The color of the blood
varies from the i)urest oxyhemoglobin red to the darkest brown-black
or a blacidike Chinese ink, even after complete saturation with o.xygen.
The red color is due to liemoglobin homogeneously distributed within
tho blood cells. Wiieiiever the color is brown, besides this hemoglol)in
there is another granular pigment of brown color within the cells
wliich will be proved to be hematin. Red blood was encountered in
some few of the smallest indiviihials and in some of the very largest
sex-mature females. The majority of the individuals, of medium size,
contained brown or brown-bhick blood." (Baumberger and
Miehaelis, 1931, p. 417.)

»» Mnrfilnltlp, llO.'*, p. 340. A carrful pnpcr bawd upon observation of living material and tho only one
dcsrrlMnit tho l)»>hii%ior of the nophrldlnl nppcndoKos.
» Pbyslologlcal Zoology, vol. 4, p. 430, 1031.

270 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Alimentary canal.^^ — As indicated in the family and generic
diagnoses, the most characteristic f eatm-es of the aUmentary tract are
the extremely long presiphonal foregut, with its very extended gizzard
segment, and the ''hind-gut" modified to act as a respiratory organ.
These features are closely similar in the three species, all of which I
have dissected. The apparent differences in the published figures
are due to limitations of material or faulty observation. In plate 34,
figure 1, the greater part of the siphonal and part of the postsiphonal
gut have been removed, A very contracted specimen was used owing
.to limitations of plate. Here it is obvious that the foregut is con-
siderably longer than the body. In a well-expanded preserved speci-
men, 300 mm. long, the pharynx is 30 mm. long; esophagus 40 mm.;
crop 85 mm. ; gizzard 85 mm. ; stomach 50 mm. ; total 310 mm. In an
expanded specimen the distance between the position of the stomach
and anus may equal nearly one-third body length and the muscular
mesentery may be, as in the specimen under discussion, 35 mm. long.
The posterior attachment of this mesentery is indicated also in plate
36, figure 4, M\

The pharyngeal lining is thrown into very coarse longitudinal
folds, which really begin on the ventral side of proboscis and run
directly to the esophagus, diminishing in size. In the esophagus
the much smaller ridges are cross cut by deep narrow channels, which
divide them into rings of oblong verrucae, giving the exterior a ringed
appearance. The esophagus is definitely begun at the last of the
dorsolateral mesenteries of pharynx (pi. 34, fig. 2, 15). In this
figure an attempt is made to show the muscular mesenteries of the
pharynx by a view looking forward into the head region from just
back of the first pair of neplnidia. The pharynx has been pulled
upward by the contraction of these dorsal and dorsolateral mesenteries
1-7 (left side). The ventral mesenteries {IJj) are the least variable.
Muscle 13 (paired) is not connected with the alimentary canal but is
attached below to body wall in front of the seta, and above, dorso-
laterally. All these muscular mesenteries are characteristic of the
genus and on direct comparison appear to be more robust in chilensis
except 13, which is better developed in caupo.

The crop (crop 1 of Seitz) is subtended by a strongly muscular
mesentery not attached to body wall. The posterior part of esoph-
agus (pi. 34, fig. 1, X) loses its ringed appearance, the mucosa hav-
ing deep, fine, longitudinal folds. In the crop the mucosa is again
regularly verrucose, but of finer texture than in esophagus. In the
gizzard the mucosa is thrown into strong ring folds, formed by the
annulate muscles of gizzard wall, conspicuous superficially. The

" Embleton, 1900, pi. 8, gives some figures of the histology of alimentary canal of U. unicinctus; Seitz,
1907, pi. 31, for U. chilensis.

ECHIUROID WORMS OF NOKTH PACIFIC — FISHER 271

crop and gizzard of chilcnsis by direct comparison are definilcly
longer than in caupo by about 25 percent.

The stomach (crop 2 of Scitz) corresponds to that of Thalasscma,
and the mucosa lias about 12 distinct longiseries of compressed ver-
rucac. Externally the stomach has the longitudinal zonation charac-
teristic of Thalassema and allies. It ends abruptly at the beginning
of the much wider midgut with its ciliated groove. About 3 mm. from
end of stomach the siphon begins, while the ciliated groove continues
adjacent to it, along the inside of intestine (pi. 35, fig. 5). A strong
muscular mesentery subtends the stomach and is attached posteriorly
a little to left of nerve cord {^P). In chilcnsis the siphon begins 9 mm.
from end of stomach as compared to 2.5-3 mm. in caupo.

The course of the intestine in a fully expanded specimen is shown
in plate 35, figure 1. The anterior and posterior portions of the body
are omitted. Plate 36, figure 4, shows the cloacal region of the same
specimen. It will be seen that the siphonal part of the intestine is
very long and includes two anterior and two posterior bends. Three
segments of the gut are attached by muscular mesenteries on the right
side of body and three (including the big "hind-gut") are attached
on the left side. The foregut is not attached to body wall except by
the strong phamygeal mesenteries and the mesentery of the stomach
(-IP). In this figure the breadth is accentuated because the body
wall is pinned out flat. The mesenteries of siphonal gut farthest to
right are attached about halfway between midventral and middorsal
lines.

The postsiphonal "small intestine" is rather short and is anchored
by heavier mesenteries than are found on the siphonal portion anterior
to the last loop. The "hind-gut," used as a respiratory organ, varies
in dilation and consequent thickness of wall in difrerent specimens.
The wall is usually thin and translucent. It is firmly anchored along
its entire length on the left side of the nerve cord. On plate 36,
figures 1-3, I have shown the attachment of the anterior end of the
"hind-gut" in the three species. Unless the single specimen of
cliilensis available for dissection is abnormal, there is considerable
difference between it and caupo.

The cloaca is probably normally elongate as shown in plate 36,
figure 4. It is here shown opened for the entire lengtli. 'J'he mucosa
of the posterior third is deeply furrowed longitudinMlly, and in this
part, on the ventral side, are the openings of the two anal vesicles.
The fecal pellets which sometimes crowd the i)ortion of small intestine
shown in plate 34, figure 1, are cylindrical with rounded ends (pi. 35,
fig. (V).

T\\v fresh colors of the viscera in an anesthcti/cd specimen are:
Foregut, pale flesh or skin color; anterior third of midgut i)ale gray-
green mottled Nsith brown; middle third, mottled yellow antl dull

272 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

gray-green; posterior third pale gray-green; siphon, pale jade green;
respiratory gut, translucent raw sienna.

T?/^g.— U.S.N.M. No. 19616.

Type locality. — Elkhorn Slough, an estuary of Monterey Bay,
Calif. ; shallow water, muddy sand.

Distribution. — California: Humboldt Bay, Tomales Bay, Mon-
terey Bay (see note below), Morro Bay, Newport Bay. With one
exception all specimens have been found living under essentially
estuarial conditions; that is, in quiet bays or sloughs in sandy mud.
As a rule the openings of the burrows are under water at low tide,
but are sometimes uncovered. However, in June 1923 I found one
good-sized specimen in a bucket holding flounders and other fish
caught in essentially open sea conditions near the Hopkins Marine
Station, Pacific Grove, Cahf. This record points to the probable
occurrence of the species at moderate depths almost anywhere off the
coast of Cahfornia where mud of the proper consistency for permanent
tunnels is present.^*

History. — The first specimens of which I have any knowledge were
collected in 1903 by C. S. Thompson, at Morro Bay, Calif., and
brought to Stanford University. Some of these, in a good state of
preservation, are still in the museum there. In 1920 I found one
specimen in Elkhorn Slough, Monterey Bay, where a few years later
Dr. Myrtle Johnson collected the examples from wliich the figures in
"Seashore Animals of the Pacific Coast" were drawn. In 1923 a
specimen was brought in by flounder fishermen from the sea bottom
off the Hopkins Marine Station. It was not until 1926 and 1927,
however, that the animal was studied. In connection with an
ecological exploration of Elkhorn Slough, Prof. G. E. MacGinitie,
then a graduate student working at the Hopldns Marine Station,
found them in quantity. By means of narrow aquaria filled with mud
("limoria") and glass-tube facsimiles of the actual burrows, he was
able to observe living animals under essentially normal conditions,
for Urechis seems to be insensible to light. Every important fact
in the ecology of Urechis has been discovered by Professor Mac-
Ginitie.^^

Habitat. — The first field studies were made at Elkhorn Slough, a
shallow estuary, tributary to Monterey Bay, where the water, al-
though slightly warmer than that of the ocean (which here varies
from 49° to 57° F.), has practically the same salinity, there being
usually a free interchange with each tide. In this inlet dwell a con-
siderable variety of bivalves, some of which are much sought for food.
There are two very interesting decapods, Callianassa californiensis
Dana and Upogebia pugettensis (Dana) , which, like Urechis, construct

" Dr. Earle H. Myers tells me he has found Urechis in the stomach of dogfish caught northwest of San
Francisco Bay entrance (Golden Gate).
» Fisher and MacGinitie, 1928b; MacGinitie, 1935b, pp. 682-686, 688, 715, 717; 1938, p. 208.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER

273

tumiels ill the mud tiiul conduct a more or less permanent manage.
The mud teems with amiehds such as Lumbrinireis, and there are
hterally acres that have a greenish tinge from the tentacles of Phoro-
nopsi^ rirulis Hilton. Zostera grows in jiermanent patches and sup-
ports a characteristic association of animals. At favorable times wide
expanses support a growth of green Enteromorpha which, either fresh
or decayed, is an important food element, shice the bulk of animal
life consists of detritus feeders.

At low water broad areas are left bare, but Urechis usually excavates
its home where the entrances are not exposed at lowest tide. A few
places were found where they are exposed at lowest tide.

Figure 19. — A, Urechis caupo Fisher and MacGinitie, in resting posture. B, Position
assumed while dig^'ing with anal setae. C, Plan of Urechis tunnel, the worm in situ
pumping water through the slime-tube. Water enters at C, where there are two Cleve-
landia ios (one outside); at a, Hespernoe lies in wait to feed on tube when it shall be swal-
lowed; b, the tiny clam Cryptomya californica; c, Scleroplax; d, Clevelandia creating
disturbance; e, eruption of mud cloud on ventilating current; castings around exit.

The tunnel (fig. 19, C), never carried very deep, has two entrances
and is in the form of a widely expanded U, of which the uprights are
nearly perpendicular and the bottom horizontal. The apertures are
small, being about one-third the diameter of the tunnel itself. Around
one of the openijigs there is a considerable cjuantity of castings. The
greatest distance between entrances measured 38 inches, the shortest
16 ii\ches. Twenty-seven was the average for many measurements.
The distance apart of the two entrances depends upon the size of the
animal but not proportionately so, for small specimens have more
extensive burrows for their size than larger ones. The largest speci-
men obtained was 19.5 ijiches long when relaxed in anesthesia, and
the smallest was 1 inch.

The burrows have a permanent aspect and none of those contijiutiily
observed was found changed except that occasionally one had a Jiew
entrance. The animals grow very slowly, and so the enlargements need

274 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

be made only at infrequent intervals and then only by widening the
U and extending one of the entrances.

Locomotion. — A Urechis can move along a smooth surface in much
the same manner as an earthworm. It elongates the anterior part
of the body and then forces forward the viscera and the water con-
tained in the respiratory gut by contracting the posterior region and
relaxing the anterior. When most of the body weight is in the anterior
end the posterior portion is draAvn up. These movements are re-
peated as the animal proceeds. Its method of locomotion within the
burrow is quite similar except that the animal has the added advan-
tage of being able to wedge the anterior portion against the sides of
the burrow. Its rapidity of locomotion approximates that of an earth-
worm. It can move considerably faster when in the burrow than
when out of it, and it can move backward nearly as fast as forward.

Digging. — When digging a tunnel Urechis forces its proboscis into
the mud and works out a hole until the body can be drawn into it.
This process is continued until the worm completes a U-shaped
tunnel open to the surface at both ends, so that a supply of fresh
water may be pumped through the tube by peristaltic movements of
the body. Then the bore of the tunnel is enlarged by scraping ma-
terial from the sides by means of the anterior setae, working it back-
ward with the anal setae, and finally blowing it out the "back door"
by a blast of accumulated respiration water from the hind-gut. To
loosen sand from the sides of the burrow, the oral setae are protruded,
then drawn backward through the sandy mud. This digging is done
on all sides of the burrow as the animal can rotate its body at will.
The setae are shed occasionally and renewed.

The use of the anal setae, which form a ring of 10 or 11 a short
distance from the anus, is higlily characteristic and was carefully
observed. The sharp retractile bristles curve forward. The mid-
ventral seta is lacking, ard the pair on each side are distinctly shorter
than the four or five dorsal setae. Wlien a certain amount of loosened
debris accumulates from the activities of the anterior bristles, Urechis
crawls over it and forces it backward, in one of two ways: either by
blowing the sand along with anal-water jets, augmented by the vigorous
ventilating stream of the tube, or by turning under the posterior end
of the body and then vigorously straightening it (fig. 19, B). The
loosened material is shoved along the tunnel, whereupon the anal jet
and ventilation current propel the finer detritus still farther. The
animal backs up and repeats the process. When the posterior end is
folded under, the dorsal setae are strongly everted and their forward
curvature favors efficient scraping. The ventral setae (now dorsal
in position) are against the vential body wall and do not function.
A reason for the smaller ventral setae (and the absence of the mid-

ECmUHOID WOHMS OF XOHTH PACIFIC — FISHER 275

ventral sctji) is now apparent, if wo linve faith in the cfTicacy of use
and nonuse in (Joterniining the rehitive size of siniihir orpins.

Castings are sometimes ejected fioni the burrow by tliis flipping of
the posterior end of the body (wliieh ean be aihnirably imitated with
the forefinger), but usually only by water currents. Castings are
allowed to accumulate and then are ejected in (juantity from one
entrance. When digging downhill the animal shoves the soil along
the body and then out by backing up the burrow, forcing the sand out
the last inch or two by water currents. The opening then resembles
a miniature vok-ano with line dark sand sj)outing out and the roily
water trailing oil' from the crater like smoke. A major convulsion
will carry out fragments of shells 2 or 3 mm. in section. Larger
objects are avoided or allowed to fall toward the lower part of the
burrow where the}' are buried. Doubling the velocity of water in-
creases its carrying efficiency directly as the sixth power. 'J'hc nar-
rowed mouth of the tunnel undoubtedly aids in increasing the force
of these "volcanic" manifestations and hence their efficiency in re-
moving sizable debris.

Once Urechis is settled in a permanent home its daily activities
consist of respiratory movements, obtaining food, cleaning the bur-
row, and resting.

Hespiratory 7noreme7tts. — There are two separate movements con-
cerned with the respiration of Urechis: (1) The peristaltic movements
along the bod\' which pump fresh water into the tunnel and move
that within respirator}' chamber of the intestine; (2) the inhalations
and exhalations, through the anus, for which the muscular cloacal
chamber, resembling that of a holothurian, su|)i)lies the ciiief motive,
power.

The inhalations are from 1 to upward of 30 iji succession (witliout
an exhalation). Exhalation is usually a single discharge although
infrefjuently a rest may occur duiing a period of exhalation. The
rote of breathing is not uniform. For instaiu'c, 2 inspirations cover-
ing 25 seconds were followed by an expiration period of 10 seconds,
whil(! in another instance 7 inspiiations occupied 2") seconds, the ex-
piration 10; 24 hisj)irations occuj)ied 70 secojuls, the single expiration
50; 30 inspirations occupied 90 seconds, the expirations only 25.
Inspirations fewer than 12 predominate in a total of 11 cycles timed.
These tinu's were tak(;n on a specimen lying in a pan of water. In
its natural environnient Urechis bi-eath(!S more .'^lowly, but with the
same irregidarily.

The peristaltic movements of the body which serve to propel water
thr(tugh the tul»(' are oven more erratic. The wa\'e, which expands
the body to lill the burrow, begins at the base of the j)roboscis and
passes along the body at varying rates for dilferent waves or even the
same wave in diirereul parts of the body. As one wave arrives at the

276 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

posterior portion of the body a new one begins at the anterior end,
Urechis is normally always in control of the water and senses any-
thing which may attempt to pass through the burrow.

Feeding. — The unique method employed by Urechis to gather
nourishment is generic and furnishes a striking example of the coordi-
nation of adaptive structure and behavior.

A short distance back of the oral setae is a zone of compound
mucous glands, which form a sort of clitellum very faintly differen-
tiated externally by the ringlike arrangement of the low rugosities of
the skin. These glands are specializations of the simpler and more
numerous multicellular flask-form mucous glands of the integument.
The slime-net or girdle glands, as they have been called, secrete a
fairly long transparent mucous tube, or funnel, whose upper, open
end is fastened near the mouth of the tunnel while the lower remains
attached to the clitellum. This strains from the ventilating, or respir-
atory currents, all minute particles as the water flows through and
when sufficiently loaded the tube is swaUowed. The process is
repeated as long as the animal feeds. Urechis readily adopts a glass
substitute for its normal burrow so that feeding reactions can be closely
followed (pi. 37, flg. 1).

Just before starting a tube, the body is constricted at the clitellum.
This region is then expanded until it presses firmly against the sides
of the burrow (usually near the mouth, but sometimes in the hori-
zontal portion) with which it remains in contact for about 2 seconds.
During the spinning process, which occupies only a few minutes, the
constricted portion of the body anterior to the clitellum (whence the
slime is issuing) undergoes a curious spiral peristalsis (pi. 37, fig. 5)
easily detected by watching the nerve cord, which shows through the
pink body wall, while back of the clitellum the normal respiratory, or
pumping, peristalsis is taking place.

The tubes vary in length from about 2 to 8 inches without apparent
reason. As the tube lengthens Urechis backs down the tunnel, and on
completion the spiral peristalsis anterior to the clitellum ceases, being
replaced by a faint normal peristalsis, the main wave starting just back
of the attachment of slime tube to the body. These normal ventilating
reactions are kept up until the animal, apparently sensing the blocking
of the water current by the clogging of the mucus with detritus, slips
the tube forward "over its head." In doing this it deftly catches the
hind edge of the tube by expanding the proboscis and bending it back-
ward, collarwise, against the inflated nuchal region, until the muscular
pharynx is able to pick up and suck in a portion of the margin (pi. 37,
fig. 7). When diatom culture or detritus is introduced with a pipette
the slime tube is soon swallowed; but if unmolested, Urechis may
continue pumping for an hour before the tube is clogged.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 277

Usually only a few minutes are required for swallowing the tube,
but the time depends upon length of tube and the amount of detritus
intercepted. ^Yhen the tube has been swallowed up to the point of
attachment the animal makes a movement to release it from the
sides of the burrow similar to the reaction while digging with oral setae.

The food funnel is porous to liquid but will intercept the smallest
particles. Phenol red passes through everywhere, but no carmine
particles ever do. Under the microscope no openings can be detected,
but particles approaching a micron in diameter are lodged in the
nmcus.

When first secreted the tube is perfectly transparent, but as it
collects detritus it becomes gray and its outlmes are easily seen.
Peristalsis becomes more energetic as the tube-wall fills.

When spuming the tube or lying at its lower end pumping water
through it, Urechis is very sensitive to disturbances. If water is in-
jected into the mouth of the tunnel, the animal immediately ceases
movement and remains perfectly still for a minute, then slowly
resumes peristalsis. If the disturbance is too great, it will drop out of
the tube and retreat toward the center of the burrow, returning later
to eat the slime tube. While it is lying at the end of a completed slime
tube any sUght disturbance such as the introduction of a little mud or
fresh clean meat will cause Urechis at once to pass the tube forward and
begin swallowing. No large particles are ingested. They are rejected
as the tube is being swallowed.

Urechis feeds to some extent, although not very efficiently, when
lying without its burrow in an aquarium. In such a position it will
swallow sediment from the bottom of the aquarium gathering it with
the proboscis.

Resting. — After a period of feeding Urechis goes to the horizontal
portion of the burrow, contracts its body so that it fits the tunnel
snugly, and lies in a state of suspended activity during which even
respiration ceases. These rests may last for an hour or more, but the
long rests are always preceded by one or more short rests, which last 4
to 8 minutes, and between which respiratoiy water is expclh'd and
more taken in (fig. 19, A).

Commensals (pi. 37, figs. 1-3). — Urechis has three pcrmaiK'nt
commensals: A polynoid annelid, IIes])er()ii<>c adirnior (Skogsberg),
and 2 pinnotherid crabs, ScleropUiT. (franv/uta Katlibun and Piniiixa
Jranciscana Rathbun. Sometimes all three are found in (he same
burrow, but usually only a Ilesperotioe and either a ScleropUix or a
Pinnixa. In addition, the litth; Cryptomya californica (Conrad)
projects its siphons into the burrow to make use of the water in the
burrow for its source of food and oxygen. The goby Clevelnmlia ios
(Jordan and Clilbcrt) uses the burrow as a retreat rather than a resi-
dence, as the little fish freely forages outside, returning when alarmed

278 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9«

or when the entrance is left exposed by low tide. On such occasions
one to five gobies may be taken from the upper part of the tunnel.
A goby left at the laboratory for several weeks in a glass burrow ap-
peared contented. It would pass from one end to the other, wriggling
past the Lrechis as if accustomed to doing so. At Newport Bay a
pair of either Betaeus longidactylus Lockmgton or Crangon [Alphaeus]
californiensis (Holmes) have been found permanently established in
the burrows of U?^echis. The former is also recorded from Elkhorn
Slough from Urechis burrows (MacGinitie, 1935b, p. 706).

Hesperonoe adventor, which ranges in length when alive from 15 to
50 mm,, is commensal with Lrechis throughout its range from Hum-
boldt Bay to Newport Bay and normally is not found outside the
burrows. The food of Hesperonoe consists of particles rejected by
Lrechis when swallowing its slime tube. These particles consist of
either living or dead animals which wash down the burrow with the
current and become entrapped in the slime-net. Sometmies when
Lrechis is swallowing its slime tube the polynoid will crawl forward and
eat part of the tube and contents. It is very aggressive toward in-
truders within the burrow other than the commensal crabs. Only one
Hesperonoe occurs within each burrow, and if another enters the two
will fight until one is killed or driven from the burrow. Other annelid
worms wliich may find their way into the burrow are speedily dis-
patched (by means of the short eversible toothed proboscis) and
devoured.

Hesperonoe rests with its dorsal surface in contact with the body of
Lrechis, moving along the bm-row with the latter by making little
short runs as the peristaltic movement of the body of Lrechis passes
by. It always faces in the same du-ection as Lrechis, and when the
latter turns in its burrow the annelid quickly does likewise. Hespero-
noe is also commensal in Echiurus tunnels.

Scleroplax ranges from 3 to 13 mm. across the carapace and is com-
mensal also in the tubes of Callianassa calijorniensis and Upogebia
pugettensis. It rests facing the side of the burrow, the chelipeds
turned up in front and the last pair of legs raised behind. In this
posture it can travel sidewise along the tube much faster than its host.
Its food consists of particles which wash into the burrows or are un-
covered by the hosts. Pinnixa franciscana screens detritus by
means of its second maxillipeds, and it will also feed on particles of
worms, clams, etc. Scleroplax has never been observed screening
plankton. As many as six Scleroplax have been taken from one
Lrechis burrow. A male and a female are often found together or two
females. In one instance an ovigerous female was found with an
ovigerous Pinnixa, and in another burrow a male Scleroplax and a
male Pinnixa.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 279

Enemies. — Urechis piohahly attains a ripe old aj^o. Five spociinons
of (lifToront sizo, kept in mud in the laboratory for over a year, appeat
not to have p-own. However, as their food is prineipally detritus, and
as natural conditions tire necessary to keep this stirred up in order that
any quantity may be drawn into the slime net, laboratory <;rowtli tests
are not convincini:. Yet what mitrlit be temied the settled habits of
the creature ami the scarcity of very small specimens point stronjj:ly
toward lonjicvity. The only animal known to prey upon them is the
stinj: ray (Myliohattis califorjiicus Gill), which can dijj: out an occa-
sional Urecfii.'^. In tiie ocean, however, small worms are possibly eaten
by flatfishes, which repdarly feed upon Listriolohus pelodes. As
already noteil, Dr. Earle H. Myers found Urechis in the stomach of
small sharks.

The period of mortality probably comes during the larval stajre.
The small goby {Clevelandia ios) is extremelj' numerous, darting here
and there, for any moving particles. These fish range from half an
inch to IK inches in length and often devour objects so small as to be
invisible to the observer. On one occasion 400 of these little gobies
were netted from a hole, 3 by 6 feet, left by clam diggers. In addition
the tiny Urechis must run the gauntlet of a host of small prcdaceous
crustaceaiLs, annelids, nemerteans, and mollusks which forage on the
surface and in the upper layers of mud. Once established in a biuTow
Urechis is relativel}' safe.

Parasite. — I have found rather numerous cestode larvae 0.25-0.32
mm. long in the proximal end of the siphon where they perhaps cause
the hernialike swellings of the siphon wall (pi. 35, figs. 4, 4a, 5).
Probably the adult is to be found in the sting ray.

Spawning. — Stored sex products are found in the ncphridia through-
out the 3'ear. MacGinitie (1938, p. 208) states that normal spawning
takes place during a short season, usually in spring or at the begimiing
of summer as the temperature of the water rises. One male which he
kept in the laboratory for two or three years spawned on May 24 and
25. Just prior to spawning the worm came nearly to the opening of the
glass tube which served as a habitation. Three welts were thrown around
the body so that the circular creases were just anterior to each of the
three pairs of gonopores, and the gonopores themselves were somewhat
protruded and turned toward the anterior end of the body, and,
therefore, t^)war<l the opening of the tube. The gonopores became
quite consj)icuoijs; this was folh)wed by several retching movem(>nts,
as if the aninud were attempting to n'giirgitate, and then sperm issued
in a stream from each gonopore, VVIu-n the sperm ceased to bo
expelled, the animal underwent violent jieristalsis, the waves running
from tin- posterior to the anterior end, causing the sperm to i)our out
of the glass tube. The retching, followed by the violent antiperistalsis,
was performed three distinct times. On both (hiys after s[)awning the

B70329 — 40 5

280 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se

worm went back to the bottom of the tube, pmnped vigorously for
§ome time, and then resumed feeding. During spawning the body of
the worm was much more elongated than normally. The spawning
on May 24 occurred at 4:30 p. m., that on the following days at 9:20
a. m. Although tae nephridia were emptied the first day of spawning,
the movements and procedure on the second day were the same as for
the first spawning, but very little sperm was discharged.

The embryology of Urechis caupo, outside the scope of this paper,
has been thoroughly described and figured by Dr. W. W. Newby
(1940). In this paper, which merits the highest praise, the relation
of the Echiuroidea to other phyla is fully discussed.

ADDENDUM

In 1942 Dr. Sixten Bock published an important memoir "On
the Structure and Affinities of * Thalassema' lankesteri Herdman and
the Classification of the Group Echiuroidea." Owing to delays
occasioned by the war, it has been possible to incorporate only the
most important systematic data in the foregoing report, such as the
new genera Ikedosoma and Maxmulleria. It is to be hoped that
Dr. Bock will continue his fundamental work and will be able to
revise the genus Ochetostoma, badly in need of an overhauling.

As Dr. Bock's scheme of classification differs from mine, it is given
herewith in skeleton form:

Class EcHiuHOiDEA [of phylum Annelida]
I. Order Echiuroinea, nov.

1. Family Echiuridae Baird, 1868. Genera: Echiurus and Urechis.

2. Family Thalassematidae, nov.

a. Subfamily Ikedinae, nov. Genus: Ikeda.

b. Subfamily Thalassematinae, nov. Genera: Thalassema, Ocheto-

stoma (incl. Li stri Globus) , Ikedosoma, Arhynchite.

3. Family Bonelliidae Baird, 1868. Genera: Maxmiilleria, Acantho-

hamingia, Archibonellia, Hamingia, Parabonellia, Protobonellia,
Pseudobonellia, Bonellia.
JI. Order Saccosomatinea, nov.

1. Family Saccosomatidae Theel, 1906. Genus: Saccosoma.
III. Order Poeobiinea, nov.

1. Family Poeobiidae Heath, 1930. Genus: Poeobius.

"The two latter orders comprise each a single species and they
must be regarded as very aberrant Annelids of somewhat doubtful
relationship to the true Echiuroids" (p. 17).

LITERATURE CITED

Annandale, Nelson.

1922. The marine element in the fauna of the Ganges. Bijdr. Dierk., pt. 22,
Feest-nummer TO*'*" Cleboortedag van Dr. Ma.\ Weber, pp. 143-154.
(Anelassorhynchus, new genus; ThalasseTtta gangelica, nomen nudum.)
Annandale, Nelson, and Kemp, Stanley.

1915. Fauna of the Chilka Lake. The Echiuroidea of the lake and of the
Gangetic Delta. Mem. Indian Mus., vol. 6, No. 1, pp. 55-63,
3 figs. (Thalassema dendrorhynchus, T. branchiorhynchus.)
Baird, William.

1868. Monograph of the species of worms belonging to the subclass Ge-
phyrea; with a notice of such species as are contained in the collec-
tion of the British Museum. Proc. Zool. Soc. London, 1868, pp.
76-114, 3 pis.
Bavmberger, James Percy, and Michaelis, Leonor.

1931. The blood pigments of Urechis caupo. Biol. Bull., vol. 61, pp. 417-421.
Blainville, Henri Marie Ducrotat de.

1827. Dictionnaire des sciences naturelles, Vers, vol. 49.
Bock, Sixten.

1942. On the structure and affinities of "Thalassema" lankesteri Ilerdman
and the cla^j^ification of the group Echinoidea. Goteborgs Vet.-
Samh. Handl., ser. B, vol. 2, No. 6, 94 pp., 11 figs., 6 pis.
Brandt, Alexander.

1835. Prodromus descriptionis animalium ab H. Mertensio in orbis terrarum
circumnavigatione obscrvatorum, fasc. 1, p. 62.
Chamberlin, Ralph Vary.

1920. The Gephyrea. Report of the Canadian .Arctic Expedition 1913-18,
pt. D. (Contains excellent bibliography covering Echiuroidea,
Sipunculoidea, and Priapuloidea, pp. 8-21.)
CoN.v, Herbert William.

1886. Life history of Thalassema mcllila. Studies Biol. Lab. Johns Hopkins
Univ., vol. 3, No. 7, pp. 351-401, 4 pis.
Danielssen, D. C, and Koren, Johan.

1S81. Den Norske Nordhavs Expedition (1876-1878). Gephyrea, pp. 20-42,
48, 3 pl.s.
Dendy, Abtiilb.

1898. Notes on a remarkable collection of marine animals found on the
New Brighton beach near Christchurch, New Zealand. Trans.
Proc. New Zealand Inst., vol. 30, pp. 320-38G. (Kx Bock, 1942.)

])KASCHE, R. VON.

1881. Cber cine neue Echiurus Art au.s Jaj)an nebst Bermerkungen tiber
Thalasnema erythrograrnmon S. Lcuckart von der Insel Bourbon.
Verh. zool.-bot. Cos. Wien, 1881, pp. 3-5, pi. 20, fig. 1.
EIllBLF,To^f, A. L.

1900. On the structure and affinities of Echiurus unicinclus. Truii.s. Linn.
Soc. London, ser. 2 (Zool.), vol. 8, pp. 77-97, 1 fig., 4 pis.
Fischer, Wilhelu.

1919. Gephyreen der StldwestkQste Australiens. Zool. An/,., vol. 50, pp. 283-
285, 2 figs.

281

282 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Fischer, Wilhelm — Continued

1926a. Sipunculiden und Echiuriden der Hamburger Siidsee-Expedition
1908-1909. Mitt. zool. Staatsinst. und zool. Mus. Hamburg, vol.
42, pp. 109-117, pi. 3, figs. 5, 7-13.
1926b. Sipunculoidea und Echiuroidea. Die Fauna Siidwest-Australiens,
vol. 5, pp. 207-209, 2 figs., 1 pi.
Fisher, Walter Kenrick, and MacGinitie, George Eber.

1928a. A new echiuroid worm from California. Ann. Mag. Nat. Hist.,

ser. 10, vol. 1, pp. 199-204, 1 pi.
1928b. The natural history of an echiuroid worm. Ibid., pp. 204-213, 3 figs.,
Ipl.
Forbes, Edward.

1841. A history of British starfishes, and other animals of the class Echino-
dermata, pp. 259-267.
Gay, Claudio.

1854. Historia fisica y politica de Chile . . . , Zoologia, vol. 8, Supplement,
p. 475.

GlSL^N, TORSTEN.

1940. Investigations on the ecology of Echiurus. Lunds Univ. Arsskr., new
ser., vol. 36, No. 10, 36 pp., 26 figs. 6 pis.
Greef, Rutger.

1879. Die Echiuren (Gephj^rea armata). Nova Act. Leop. Carol., vol. 41,
172 pp., 9 pis.

GufiRIN-M^NEVILLE, F^LIX EdOUARD.

1831. Iconographie du regne animal de G. Cuvier, vol. 2 (Zoophytes), pt. 6.
Hall, Victor Ernest.

1931. Muscular activity and oxygen consumption of Urechis caupo. Biol.
Bull., vol. 61, pp. 400-416.
Heath, Harold.

1930. A connecting link between the Annelida and the Echiuroidea (Gephyrea
armata). Journ. Morph., vol. 49, pp. 223-246, 4 figs., 3 pis.
Herdman, William Abbott.

1897. Note on a new British echiuroid gephyrean, with remarks on the genera
Thalassema and Hamingia. Quart. Journ. Micr. Sci., new ser.,
vol. 40, pt. 3, No. 159, pp. 367-384, 2 pis.
Horst, R.

1881. Die Gephyrea gesammelt wahrend der zwei ersten Fahrten des Willem
Barents. 1, Echiuria. Niederl. Arch, fiir Zool., Suppl. Band 1,
12 pp., 1 pi.
Htman, Libbie Henrietta.

1940. The invertebrates: Protozoa through Ctenophora, 726 pp., 221 figs.
Ikeda, Iwaji.

1904. The Gephyrea of Japan. Journ. Coll. Sci. Imp. Univ. Tokyo, vol. 20,

art. 4, pp. 1-87, 4 pis.
1907. On three new and remarkable species of echiuroids. Journ. Coll. Sci.

Imp. Univ. Tokyo, vol. 21, art. 8, pp. 1-64, 4 pis.
1908a. Note on a new deep-sea echiuroid, Protobonellia mitsukurii, nov. g.
et nov. sp. Annotationes Zoologicae Japonenses, vol. 6, pt. 4,
pp. 259-265, 4 pis.
1908b. On a new echiuroid {Hamingia ijiniai) from the Sagami Bay. An-
notationes Zoologicae Japonenses, vol. 8, pt. 1, pp. 61-68, 1 pi.
1910. Notes on a deep-sea echiuroid, Acanthohamingia shiplei (n. g. et n. sp.),
with remarks on the species Hamingia ijimai Ikeda. Quart. Journ.
Micr. Sci., new ser., vol. 56, pp. 135-147, 1 pi.

ECHIUHOID WOKMS OF NORTH PACIFIC — FISHER 283

Ik EDA, IwAji — Continued

1924. Further notes on the Gephyrea of Japan with descriptions of some new
species from the Marshall, Caroline, and Palau Islands. Jap. Journ.
Zool., vol. 1, No. 2, pp. 23-44, 1 pi.
Jameson', H. Lyster.

1899. Contributions to the anatoni_v and histology of Thalassema neptuni
Gaertner. Zool. Jahrh., Abt. Anat., vol. 12, pp. 535-566, 1 fig., 3 pis.
JoHN'soN', Myrtle Elizabeth, and Snook, Harry James.

1927. Seashore animals of the Pacific coast, 659 pp., 700 figs., 12 col. pis.

New York.
Johnston'. T. Harvey, and Tiegs, O. W.

1919. Pseudobonellia, a new echiuroid genus from the Great Barrier Reef.
Proc. Linn. Soc. New South Wales, vol. 44, pp. 213-230, 3 pis.
Lamarck, Jean Haptiste Pierre de Monet.

1801. Systome des aniniaux sans vert^bres, 432 pp.
1816. Hi.stoire naturclle des animaux sans vert^bres, vol. 3, p. 65.
Lam pert, Ktrt.

1883. Uber einige neue Thalassemen. Zeitschr. wiss. Zool., vol. 39, pp.
334-342.
Lanchester, W. F.

190oa. On the sipunculids and echiurids collected during the Skeat Expedi-
tion to the Malay Peninsula. Proc. Zool. Soc. London, 1905,
pp. 35-41, 1 pi.
1905b. The marine fauna of Zanzibar and British East Africa (Cyril Cross-
land). Gephyrea. Ibid., pp. 28-35, 1 pi.
Lankester, Edwin Ray.

1881. On Thalassema neptuni Gaertner. Zool. Anz., vol. 4, p. 350.
Leigh-Sharpe, W. Harold.

1928. Thalassema neptuni, Gaertner — a British echiuroid. Ann. Mag. Nat.

Hist., ser. 10, vol. 2, pp. 499-504, 2 pis.
Lelckart, Fuiedrich Sigismund, and RDppell, Wilhelm Peter Eduard
Simon.

1828. Neue wirbellosen Tiere des Roten Meeres. Atlas zu der Reise im
nordlichen Afrika, .\bt. Zool., i)p. 7, 8.
.MacGimtie, George Eber.

1935a. Normal functioning and experimental behavior of the egg and sperm
collectors of the echiuroid, Urechis caupo. Journ. Exp. Zool.,
vol. 70, pp. 341-355, 1 fig., 1 pi.
1035b. Ecological a.spectH of a California marine estuary. Amcr. Midi. Nat..

vol. 16, pp. 629-765, 21 figs., maj).
1935c. The fertilization of eggs and the rearing of larvae of Urechis caupo
within the blo(jd cavity of the adult animal. Journ. Exp. Zool.,
vol. 71, pp. 483-487.
1938. Notes on the natural history of some marine animals. ,\mer. .Midi.
Nat., vol. 19, pp. 207-219, 2 pis.
.Monro, C. C. A.

1931. Polychaeta, Oligochaeta, Echiuroidca and Sipunculoidea. Great
Barrier Reef Expedition, 1928 29. vol. 4, No. 1, j)p. 1 37. {I'seudo-
honellia biulcrina JohijHton and Tiegs recorded from Low Islands.)
Mt)LLER, Max.

1852. ObscTvationes anatomicac dc vermibus quibusdam maritimis, pp.
14 22, 1 1)1. Ikrlin.

284 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Newbt, William Wallace.

1932. The early embryology of the echiuroid, Urechis. Biol. Bull., vol. 63,
pp. 387-399, 4 pis.

1940. The embryology of the echiuroid worm Urechis caupo. Mem. Amer.

Philos. Soc, vol. 16, 219 pp., 85 figs.

1941. The development and structure of the slime-net glands of Urechis.

Journ. Morph., vol. 69, pp. 303-316, 10 figs.
Onoda, Katsuzo.

1934. On Pseudobonellia, a new genus of the bonellian echiuroids. Annota-

tiones Zoologicae Japonenses, vol. 14, pp. 413-420, 3 figs., 1 pi.

1935. Parabonellia (nom. nov.) misakiensis (Ikeda), correction of my paper

on Pseudobonellia. Annotationes Zoologicae Japonenses, vol. 15,
p. 141.
PocHE, Franz.

1920. tJber eine in Vergessenheit geratenen Art der Echiuroidea. Arch.
Naturg., Jahrg. 86, Abt. A, pp. 102-104.
Prashad, B.

1919a. Zoological results of a tour in the Far East. Echiuroids from brack-
ish water with the description of a new species from the Andamans.
Mem. Asiatic Soc. Bengal, vol. 6, pp. 323-338, 1 pi. (Thalassema
kempi.)
1919b. Notes on echiuroids from Chandipore, Orissa. Rec. Indian Mus.,

vol. 16 pp. 399-402. {Thalassema microrhynchus.)
1920. On a new species of Thalassema from the Gulf of Manaar with notes
on Thurston's species T. formulosum. Rec. Indian Mus., vol. 19,
pt. 2, pp. 35-37. (Thalassema hornelli.)
1935. On a collection of echiuroids of the genus Thalassema Lamarck in the
Indian Museum, Calcutta. Rec. Indian Mus., vol. 37, pt. 1, pp.
39-43, 1 pi. {Thalassema arkati, T. marshalli.)
Prashad, B., and Awati, P. R.

1929, On a new species of the genus Thalassema from Bombay. Rec. In-
dian Mus., vol. 31, pt. 4, pp. 259-261, 1 pi. (Thalassema bombay-
ensis.)

QUATREFAGES, JeaN LoUIS ArMAND DE

1865. Histoire naturelle des annells marins et d'eau douce, vol. 2, pp. 591-

599, pi. 16, fig. 13.
Redfield, Alfred C, and Florkin, Marcel.

1931. The respiratory function of the blood of Urechis caupo. Biol. Bull.,

vol. 61, pp. 185-210, 5 figs.

RiETSCH, MaXIMILIEN.

1886. fitude sur les g6phyriens arm^s ou ^chiuriens. Recueil Zool. Suisse,
vol. 3, No. 3, pp. 313-515, 6 pis.
RiCKETTs, Edward F., and Calvin, Jack.

1939. Between Pacific tides, 320 pp., 112 figs., 46 pis. Stanford University,
Calif.
Sato, Hatao.

1931. Report of the biological survey of Mutsu Bay. 20, Echiuroidea. Sci.
Rep. Tohoku Imp. Univ., ser. 4, vol. 6, pp. 171-184, 4 figs.

1934. Report on the Sipunculoidea, Echiuroidea and Priapuloidea collected

by the Soyo-Maru Expedition of 1922-1930. Sci. Rep. Tohoku
Imp. Univ., ser. 4, vol. 9, pp. 1-32, 31 figs., 1 pi.

1935. Sipunculoidea and Echiuroidea of the West Caroline Islands. Sci.

Rep. Tohoku Imp. Univ., ser. 4, vol. 10, pp. 299-329, 17 figs., 3 pis.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 285

Sato, Hayao — Continued

1937. Echiuroidea, Sipunculoidea and Priapuloidea obtained in northeast
Honshu, Japan. Saito Ho-on Kai Mus. Res. Bull., No. 12, pp.
137- 17G, 3 pis.
1939. Studies on the Echiuroidea, Sipunculoidea and Priapuloidea of Japan.
Sci. Rep. Tohoku Imp. Univ., ser. 4, vol. 14, pp. 339-400, 60 figs.,
5 pis.
Seitz, Philipp.

1907. Der Hau von Echiunis chilcnsis {Vrechis n. g., chilensis). Zool.
Jahrb., Abt. Anat., vol. 24, pp. 323-350, 3 pis.
Selenka, Euil.

1885. Report on the Gephyrea collected during the voyage of H. M. S.
Challenger. Challenger Reports, vol. 13, pp. 1-24, 4 pis.
Shipley, Arthur.

1899. On a collection of echiuroids from the Loyalty Islands. Willey's

Zoological Results, pt. 3, pp. 335-356, 1 pi. (Review of nominal
species of Echiuroidea.)
Skorikov, a. S.

1909. Subfam. Echiurini nov. (Gephyrea armata). Ann. Mus. Zool. Acad.
Imp. Sci. St.-P6tersbourg, vol. 14, pp. 77- 102, 1 pi.
Sluiter, G. p.

1883. t)ber einige Sternwlirmer des indischen Archipels. Zool. Anz., vol. 6,
pp. 222- 228.
Spengel, Johann Wilhelm.

1879a. tlber die Organisation des Echiurus pallasii, I. Zool. Anz., vol. 2,

pp. 542- 547.
1879b. Beitrage zur Kenntniss der Gephyreen, I: Die Eibildung, die Entwick-
lung und das Miinnchen der Bonellia. Mitt. Zool. Stat. Neapel,
vol. 1, pp. 358-419, 5pla.
1880. Idem, II: Die Organisation des Echiurus pallasii. Zeitschr. wiss.

Zool., vol. 34, pp. 458-538, 4 pis.
1912a. Idem, III: Zum Bau des Kopflappens der armaten Gephyreen.

Zeitschr. wiss. Zool., vol. 101, pp. 342-385, 21 pis.
1912b. Idem, IV: Revision der Gattung Echiurus. Zool. Jahrb., Abt. Syst.,

vol. 33, pp. 173-212, 1 pi.
1912c. t)ber den Ilautmuskelchlauch gewisser Thalassema Arten und Seine
Bcdeutung flir die Systematik dieser Tiere. Verh. Deutsch. zool.
GeH., vol. 22, PI). 309-317.
Steinbeck, John, and Ricketts, Edward F.

1941. The s*;a of Cortoz, 598 i)p., 40 pis. New York.
Tn^Er., Hjalmar.

1906. Northern and Arctic invertebrates in the collection of the Swedish
Stale Mu.seum (Riksmu.stnjm), II: Priapulids, echiurids, etc.
Kungl. Svenska Vet.-Akad. Handl., vol. 40, No. 4, pp. 1-26, 2 pis.
Wharto.v, L. D.

1913. A description of sorno Philippine; Thalassemac with a revision of the
genus. Philippine Journ. Sci., sec. D, vol. 8, pp. 243-270, 8 figs.,
2 plH.
WiL-soN, Charles Branch.

1900. Our North American echiurids. Biol. Bull., vol. 1, pj). I63-17H, 1 pi.

EXPLANATION OF PLATES

All figures were made by the writer directly from dissections or
specimens.

Plate 20

Echiurus echiurus alaskanus, new subspecies

1, Dorsal view of anatomy of anterior portion of body, X 5. Body wall is indi-

cated only in pharyngeal region. Junction of diaphragm to body wall is
indicated by dashes for dorsal half and dots for ventral half. Coiled loops
of esophagus are shown emerging from posterior opening of diaphragm,
below the interbasal muscle. Lines radiating from pharynx are the muscu-
lar frenula. The peripharyngeal coelom is shaded.

2, Anal bristle from posterior ring, 7.5 mm. long.

3, Anal bristle from anterior ring, 8 mm. long.

4, Diagram of the diaphragm (ventral half dotted) showing the posterodorsal

opening. The esophagus is omitted except where it pierces the right wall
and becomes the gizzard. The edges that are attached to body wall and
ventral mesenteries are dashes.

5, Diagram of a section through diaphragm showing how its lower border merges

with the ventral mesenteries.

B^B*, dorsal, ring, neurointestinal, and ventral blood vessels, respectively,"
C, stomach; CG, ciliated groove; D, diaphragm; G, gizzard; MD, dorsa
mesenteries of esophagus; MV, ventral mesenteries of esophagus (fig. 5);
A^, nephridium; NC, nerve cord; 0, esophagus, indicated by arrows, ventral
mesentery omitted, dorsal inesenteries shown as lighter lines; P, pharynx;
Si, anterior end of siphon; X, perivisceral coelom (Sg. 4); A^^, peripharyngeal
coelom.

Plate 21

Listriolohus pelodes, new species

1, Ventral view of a large specimen from Monterey Bay, Calif., X l}4.

2, Same specimen, X 5, ventral view of anterior end showing nerve loop in pro-

boscis, a section of which has been removed.

4, Small phase before muscle bands are evident, natural size.

4c, Side view of anterior end of a living specimen, X 3.

46, A female, X 3, showing nephridia and fecal pellets. At this size the muscle
bands are not apparent unless the specimen is stronglj^ contracted. Out-
line from living animal.

Ocheiostoma octomyotum, new species

3, Ventral view of type specimen from Newport Bay, Calif., natural size. A

specimen from Cabrillo Beach, near San Pedro, 95 mm. long, has a pro-
boscis 93 mm.

Plate 22

Listriolohus pelodes, new species

Type specimen, X 7, dissected to show organs of anterior portion of the body;

the alimentary canal is drawn to the right of its natural position.
B', dorsal blood vessel; B^, ring vessel; B^, neurointestinal connective; B*, ventral
vessel; C, stomach; CG, ciliated groove of intestine; G, gizzard; I, intestine;
286

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 287

Ml, iiitorbftsal muscle of sctao; X, iiophridiiiin; \C, nerve cord; 0, esoph-
ag:iis; P, pharynx; P\ posterior end of pharynx; 8, seta;«Si/, anterior end of
siphon; VM, ventral mesentery; X, parasite.

Plate 23

Ochetostoma octomyotum, new species

1, Dissection, X 2; the spiral funnels of the nephridia and longitudinal muscle

hands have been omitted except the midventral, which is lighter shaded.

2, Interval between right ventrolateral and lateral muscles at middle of body

showing fascicles of the oblique layer. On the lower left corner the oblique
layer has been removed, X 20.

3, Cloaca and adjacent part of intestine opened to show relation with intestinal

coecum and anal vesicles, X 5.
Lettering as for plate 24.

Pl.\te 24

Ochetostoma octomyotum, new species

The anterior portion of plate 23, figure 1, enlarged X 5 and with addition of
details.

A, anus; AV, anal vesicles; AV\ their opening into cloaca; B', dorsal blood ves-
sel; D*, ring vessel; IP, neurointestinal connective of which B^ is merely a
part; B*, ventral vessel; C, stomach; CF, ciliated funnel or nephrostome;
CG, ciliated groove of intestine; CI, cloaca; DM, dorsal mesentery of pharynx,
G, gizzard; IC, intestinal coecum; IC\ its opening into cloaca; MC, outer
circular muscle layer; MD, dorsal muscle band; MDL, dorsolateral muscle
band; ML, lateral muscle band; MO, oblique inner layer of muscles; MVL,
ventrolateral muscle band; N, nephridium; NC, nerve cord; O, esophagus;
P, pharynx; S, seta; Si, siphon; Sil, entrance to siphon; Si2, end of siphon;
VM, ventral mesentery of pharynx.

Plate 25

Arhynchite inamoenua, new species

1, \'entral view of paratype, X 1.

2, Seta of type, X 10.

3, Di.s.section of anterior complex of type, X 7, showing foregut in situ.

4, Skin of figure 1, from near midventral lino, enlarged.

."J, TyjK>, X 7; the interbasal muscle has been cut aii<l the liberatod foregut drawn

to the right; ventral mesentery is dotted.
B', B*, B*, dorsal, neurointestinal, and ventral blood ves.sels, respectively; CF,

nephrostome; C, gizzard; M, mouth; MI, interbasal muscle; A^, nephridium;

NC, nerve cord; O, e.soi>hagus; P, ])liarynx; S, 8(;ta.

Plate 2G

BonelliopKig aUiHknnn, new geriu.M and sjjecies

1, Di.s.Hoction of si)ecirnen from I'liala-Hka, dorsal view, X 4.

2, Ti]) of one of the priinar\- l)rancheH of an anal vesicle, showing two secondary

branches with their ciliated funnels, X 50.

3, One of the ciliated funnels, X 200.

4, Nephrostome, X 10.

288 PROCEEDINGS OF THE NATIONAL MUSEUM vol. se

5, Anterior end of an individual, the short proboscis apparently in process of

regeneration, X 3.

6, Ventral view of a worm life size; the second proboscis indicates about the

maximum length in preserved specimens.

7, Anterior ventral portion of worm showing the contracted verrucose skin, X 10;

when the skin is fully distended the verrucae disappear, the glandular thick-
enings then appearing as squarish patches, closely spaced.
Go, gonad; other lettering as for plate 24.

Plate 27

Bonelliopsis alaskana, new genus and species

1, Dissection of anterior end of a specimen with a right nephridium filled with

eggs, X 5. A "window" has been cut in the dorsal wall of pharynx and also
one in the nephridium to show the invaginated tip, within which is the
nephrostome. The longitudinal ridges on inner wall of nephridium are indi-
cated by dotted lines.

2, Same, anterior end, in contracted state, with right half removed to show

relation of dorsal and ventral mesenteries to alimentary canal and ventral
mesenteries to alimentary canal and nephridium, X 5. Anteriorly only a
few of the frenula of pharynx indicated; back of these the transverse mesen-
tery of figure 1 (M' ) is indicated as a black line. DM, dorsal mesentery, and
DM^ (dot-dash), its attachment to right wall of body (removed); VM,
ventral mesentery, attached to lower side of intestine and mostly to left side
of f oregut and involving posterior part of dorsal artery (B') . The nephridium
lies in a sort of anterior cul-de-sac or egg trap. When the animal is extended
the upper margin of dorsal mesentery is at a distance behind the nephridium,
here shown at minimum size.

Af' , a transverse mesentery below the dorsal blood vessel;

M', special muscular mesentery between gizzard and stomach, not shown in figure
2; PV, position of pyloric valve, mentioned in text; other lettering as for
plate 24.

Plate 28

Eubonellia valida, new genus and species

1, Dissection of anterior end of body from above, X 4. The f oregut has been

drawn to the left to show the large nephridium (N) with its terminal nephro-
stome (CF). Note the unusually long stomach (C), filled with pellets,
between the gizzard (G) and intestine (/). The ventral blood vessel (B*)
and the much contracted neurointestinal vessel (B') have been cross-
hatched. The dorsal blood vessel (B') is unshaded.

2, A cluster of nephric elements of an anal vesicle, X 30.

3, Type, natural size, from below.

4, A pellet from the postsiphonal intestine, X 10.

5, Male, from mouth cavity, X 50; anterior end to right; the sperm receptacle and

duct shown.
Lettering as for plate 24.

Plate 29

Nellobia eusoma, new genus and species

1, Ventral view of anterior end of type showing the short truncate^snout, X 5.

2, Ventral view of type, X 1.

ECHIUROID WORMS OF NORTH PACIFIC — FISHER 289

3, Teruiinal portion of intestine, the cloaca, anus, and anal vesicles, X 5.

4, Branchlot of anal vesicle, X 50.

NP, nephridiopore; Go, posterior part of gonad with blood vessel and nerve cord
underneath; M, mesenteries; other lettering as for plate 24.

Plate 30

Nellobia euaoma, new genus and species

1, Dissection of anterior portion of type, seen from above, X 5. The nephridium,

filled with eggs (0.85 mm. in diameter) on the left, has a window cut in the
wall to show the constricted duct from the egg chamber. The swollen duct
leading to external opening lies under the nerve cord and ventral blood
vessel.

2, Pharynx, X 5. Interior; anterior end looking toward mouth. The dorsal side

has been cut open.
B\ B*, B*, dorsal, neurointestinal, and ventral blood vessels, respectively; CF,
nephrostome; D, peripheral portion of peripharyngeal diaphragm (the central
portion adjacent to pharynx has been removed); DAI, dorsal mesenteries;
G, gizzard; Go, gonad; M, mesenterial sheet holding loop of pharynx-esopha-
gus; NC, nerve cord; O, esophagus; Sil, beginning of siphon; VM, ventral
mesentery of pharynx-esophagus; X-Y, probable extent of gizzard; at Y
the canal was broken, and it is possible that a portion of the succeeding
stomach was lost.

Plate 31

Acanthohamingia paradola, new species

1, Ventral aspect of paratype, X 1.

2, Same specimen; genital groove, extending forward from nci)hridiopore (AO

and showing four males in situ, X 5 ( $ , males attached to skin).
2a, Anterior portion of figure 2, X 15.

3, Type; genital groove in probably the normal closed state, X 5.

4, Genital groove of third specimen that has two nephridiopores {N), X 5.
4a, Nephridiopores of above, enlarged.

5, Male from genital groove of paratype (fig. 2), 1.19 mm. long, X 50.

6, Tyj)e; anal vesicles and thin-walled cloaca from above, X 3. The anus can be

seen through the thin wall of the cloaca, and on the left most of tlie tubes
spring from a rudimentary bladder.
I, intestine Cmi.ssing from type); M, mouth; A'^, nephridiopore; S, spermotheca
(nephridium).

Plate 32

Acanthohamingia paradola, new species

Dissection of anterior part of animal from above, X 4. Note the very long forogut
ending at B^ and the long (as compared with Bonellia) segment of intestine
between B^ and Sil.

B\ B*, B*, dorsal, neurointestinal, and ventral blood vcssel.s, respectively; C,
portion corresponding to stornarli of other bonelliids; CF, nephrostome; G,
probable gizzard; /, intestine; A^, ncphridia; NC, nerve cord; 0, esophagus;
P, pharynx, Pe, pellet, X 10; Si, siphon; Sit, anterior end of siphon.

290 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Plate 33

Urechis caupo Fisher and MacGinitie

1, Dissection of contracted specimen from above, showing the intestine in hap-

hazard convolutions. The principal mesenterial bands, which anchor the
intestines to the body wall, are shown but not lettered. The coelomic aper-
tures of the nephridia are recognizable by the conspicuous coiled lips. The
arrow indicates point where pharynx becomes esophagus. The figures in
sequence on the midgut are intended to aid in following the convolutions;
1 is at the beginning and IS near the end. Beyond 13 the slight diverticulum
of the respiratory gut is indicated.

2, Pharynx contracted and slit open along ventral side to show the straight

longitudinal folds of lining, continuous with those of the proboscis. Pos-
teriorly is shown characteristic lining of esophagus; on each side are the
dorsal mesenteries. In front of these the ventral mesenteries are spread
laterally since pharynx has been opened ventrally (see pi. 34, fig. 2, 14)-

3, Ventral surface of proboscis and anterior end of body.

4, Posterior end of body showing eccentric anus and circle of setae ; + marks the

midventral line.

5, Two views of the anterior setae. The line indicates 1 mm.

6, An anal seta, same scale as figure 5, with, below, a tip enlarged.

AS, anterior setae (accessory seta shown at side; muscles not drawn); AV, anal
vesicles; Ci, C2, crops 1 and 2 of Seitz, subtended by muscular bands Mi and
M2; C2 is the stomach; CL, cloaca, the posterior portion lined with heavy
longitudinal ridges (arrows mark apertures of anal vesicles) ; G, gizzard, a
portion of the foregut lying between Ci and C2, characterized by thick
muscular walls and circular muscular ridges and constrictions; HG, respira-
tory gut, specialized posterior segment of midgut; Mi, M2, muscular bands
of crop and stomach (C2) ; A'', nephridium; NC, nerve cord; 0, esophagus,
anterior limit marked by an arrow; P, pharynx; PS, posterior or anal setae;
S, siphon or accessory intestine; SI, beginning of siphon near beginning of
midgut; S2, end of siphon; VM, ventral mesenteries of pharynx; 1-13, these
figures are in sequence along the midgut and are intended to aid in following
the course; 13 is near the junction of midgut and its terminal specialized
portion, the respiratory gut.

Plate 34

Urechis caupo Fisher and MacGinitie

1, Dissection of contracted individual showing the generically characteristic

parts of alimentary canal, most of the "small intestine" having been removed.
The very long foregut consists of pharynx (anterior to P), esophagus (0),
crop (C) with its strong muscular mesentery, gizzard (G), and stomach
(C^) anchored posteriorly by a strong mesentery here shown in maximum
contraction. Si is beginning of siphon (pi. 35, fig. 5). Attachment of
respiratory gut is always on left of nerve cord (NC) . Along its dorsal surface
_ is shown the muscle strand continued from the small intestines and serving

posteriorly for attachment of a few dorsolateral mesenteries.

2, Head region of coelom looking forward from just behind first pair of nephridia

{16) showing arrangement of muscular mesenteries of pharynx: 1-7, dorsal
and dorsolateral; 8-12, lateral and ventrolateral; 13, the dorsoventral
muscles mentioned in text; I4, ventral mesenteries of pharynx; 17, nerve

ECHIUROID WOKMS OF NOUTH PACIFIC — FISHER 291

cord; crossiiiu; the ventral mesenteries between the two figure i.{'s is the
interbasal muscle of setae; radiating muscles of setae shown in solid black
on right.

3, Anterior ainpoct of a nephridium of second pair showing nephrostome and

elongated lips spirally coiled, X 5.

4, A nephridium from a specimen 40 mm. long, not yet sexually mature, X 20.

Plate 35
Urechis caupo Fisher and MacGinitie

1, Arrangement of intestine and mesenteries in fully expanded sjx'cimen, X ^.

Anterior and posterior portions of body have been omitted and the intestine
has been spread to right and left to show attachments. Normally these
lateralmost parts overlie the darker and more mesially located portions. A
section has been removed from respiratory gut to show attachment of mes-
enteries. C, crop; C^ stomach; G, gizzard; IIG, respiratory gut; M^,
mesentery of stomach; NC, nerve cord; Si, siphon; Sil and Si2, anterior and
posterior end of siphon.

2, Portion of midgut at A' of figure 1, showing its highly sacculate structure;

mucosa with fine anastomosing plications, transverse in direction; Si,
siphon, X 2.

3, Postsiphonal midgut at XX of figure 1, X 2, showing the mucosa and longitudi-

nal muscle band marking position of ciliated groove.

4, 4a, Cestode larvae from anterior end of the siphon, X 60. These larvae vary

in length from 0.25 to 0.32 mm. and are free in the lumen of siphon and in
the hernialike swellings, which may be caused by them (.see fig. 5, above Sil).

5, Sagittal section, X 5, of the distal end of stomach and beginning of midgut and

siphon, showing macroscopic character of mucosa; C^, stomach; CG, one side
only the ciliated groove; the groove is constituted by two of these finely
plicated folds or ridges of the mucosa. The plications are coarser and the
groove is broader in the short segment CG'; Si, siphon, showing foliose
mucosa. The cestode larvae were found in this portion and in the hernia-
like swellings shown just above SjI, the narrow passage connecting midgut
and siphon.

6, Fecal pellets, X 3. Specinjen from Monterey Bay, Calif.

Plate 36

1, I'rrchin chilcnaia (Miillcr): Anterior end of respiratory gut showing mode of

attachment by muscular mesenteries, X 3. Compare wit It figures 2 and 3.

2, r'rec/ii« c«i/po Fisher and MacGinitie, X 1.5.

3, IJrechis unicinctu8 (von Drasche), X 5.

4, L'rechin caupo: Cloacal region of a relaxed specimen (pi. 35, fig. 1) with the

posterior part of respiratory gut ; dorsal wall of cloaca removed. ..1 V, anal
vcHicle; M^, nmscular masentery anchoring the stomach; M*, muscular
fronula of cloaca; A^, ncphridia; NC, ncrvo cord; PS, posterior setae.

Plate 37

Urechis caupo anfl Conmiensals

1, Portion of tunnel showing one position of worm while pumping water through
its slime-net and characteristic stations of commen.sals, X *i. A, Cleve-
landia ion (Jordan and Gilbert) at mouth of tube; B, IlespcronoS advrntor
(Skogsberg) ; C, Scleroplax granulata Rathbun; D, Cnjplomya californica
(Conrad). At U|)i)er [)f)irit where tube is interrupted one inch has been
omitted, at lower point four inches.

292 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 8«

2, Scleroplax granulata Rathbun, male, X 3.

3, Hesperonoe adventor (Skogsberg), type X 1.

4, Specimen of U. caupo with slime-tube in place, with the thickening at point of

attachment to body indicated. The worm is shown in characteristic pump-
ing posture; X 3.

5, Characteristic posture while tube is being secreted; tube just begun; entrance

indicated by dots; X 1/4.

6, Expression of worm while swallowing slime-tube.

7, Grasping slime-tube at moment of starting to swallow. The proboscis is

holding the posterior edge of tube while a portion is being sucked in on
ventral side. This step occupies about three seconds.

U. S. BOVERHMENT rRINTIN* OFFlCli l»4«

S NATIONAL MUSEUM

PROCEEDINGS VOL "^6 PLATE 20

LCMIURUS ECHIURUS ALASKANUS. NEW ^UOsHH.Itt.
POn t.Xfn.*NATION SEE PAOE ?««

U. S. NATIONAL MUSEUM

PROCEEDINGS, VOL. 96 PLATE 21

LISTRIOLOBUS PELODES. NEW SPECIES. AND OCHETOSTOMA OCTOMYOTUM, NEV

Species

FOR EXPLANATION SEE PAGE 286

U. S. NATIONAL. MUSEUM

PROCEEDINGS. VOL 96 PLATE ?2

LISTRIOLOBUS PELODES. NEW SPECIES
roB i;kpi_anation see paocs a»» m

U. S. NATIONAL MUSEUM

PROCEEDINGS, VOL. 96 PLATE 23

OCHETOSTOMA OCTOMYOTUM, NEW SPECIES

FOR EXPLANATION SEE PAGE 287

U S NATIONAL MUSEUM

PROCEEDINGS. VOL 96 PLATE 24

MV)7-*^\J MVL

^ ML ^M

OCHETOSTOMA OCTOMYQTUM Nf W ijPtCIES
row tXPLANATION SEE PAGE .'»?

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 25

ARHYNCHITE INAMOENUS, NEW SPECIES
FOR EXPLANATION SEE PAGE 287

U S NATIONAL MUSEUM

PRCX:EEDINGS. vol. 96 PLATE 26

BorJLLLIOPSIS ALASKANA. NEW GENUS AND SPECIES
FOn FXPUANATION SEE PAr.l

U. S. NATIONAL MUSEUM

PROCEEDINGS, VOL. 96 PLATE 27

BONELLIOPSIS ALASKANA. NEW GENUS AND SPECIES
FOR EXPLANATION SEE PAGE 288

U S NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 28

EUBONELLIA VALIDA. NEW GENUS AND SPECIES
FOR EXPLANATION BEE PAGE /»»

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 29

- -i

i ^

N P

/^ \

lb,

*^V 'i)

^-''^^~t.^,jJL-/UVly-

Nellobia eusoma. New Genus and species

FOR EXPLANATION SEE PAGES 298-289

U S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 30

f-JhLioBiA tijsoMA. New Genus and Species

FOR EXPUANATION Str PACiE I<W

U. S. NATIONAL MUSEUM

PROCEEDINGS, VOL. 96 PLATE 31

■/

/ i

\ t

"* t^

.i.

J- /

."?^

/¥^ -'

r .

ACANTHOHAMINGIA PARADOLA, NEW SPECIES

FOR EXPLANATION SEE PAGE 289

U S NATIONAL MUSEUM

PROCEEDINGS. VOL 96 PLATE 32

ACAN f HtJHAMINICIA PARADOLA NEW SPECIES

ro« FxriANATiOfj sie I'A'.c ;;•»

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 33

^P ^'^^

..M);. J''

\ ^

ji. '^

F^<:4-

J^r/y/ 1

■<f---*jAi

Urechis caupo Fisher and MacGinitie

FOR EXPLANATION SEE PAGE 290

U S NATIONAL MUSEUM

PROCEEDINGS. VOL S6 PLATE 34

URECHis CAUPO Fisher and MacGinitie

rOR eXPl^NATION r.EE PACES .■»- »1

U^ S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 35

Urechis caupo Fisher and MacGinitie

FOR EXPLANATION SEE PAGE 291

U. S NATIONAL MUSEUM

PROCEEDINGS VOL 96 PLATE 36

unlchis caupo. U chilensis. and U. UNICINCTUS

FOR EXPLANATION SCC PAGB 291

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL. 96 PLATE 37

URECHIS CAUPO AND COMMENSALS
FOR EXPLANAXrON SEE PAGES 291-292

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

by the

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM

Vol. % W.>hington: 1946 No. 3199

THE OSTEOLOGY OF THE FOSSIL TURTLE TESTUDO
PKAEEXTANS LAMBE, WITH NOTES ON OTHER
SPECIES OF TESTUDO FROM THE OLIGOCENE OF
WYOMING

By Charles W. Gilmore *

Among the fossil vertebrate materials collected from the Oligocene
of the Indian Creek Basin area in Wyoming by the Smithsonian
Paleontological Expeditions of 1932 and 1942 were six well-preserved
specimens of the land tortoise Testudo. Interest in these specimens
centei-s primarily in the fact that they contribute the first information
to be had of the skull and internal skeleton of Testudo j^raeextans
Lambe, as well as illustrating tlie individual variation found in the
shell structure of that species. Two of these specimens pertain to
Testudo laticunea Cope, this being the first time this species has been
recognized in this area.

Family TESTUDINIDAE

Genus TESTUDO Linnaeus

TESTUDO PKAEEXTANS Lambe

Plates 38-41; Platk 44, Fkjckk 2

Testudo praeextana Immbk, Ottawa Nat., vol. 27, j)!). '>7 (W, |)ls. 4, !">, 1913. — Hay,
Carnegie Inst. Washington Pul.i. 390, p. 101, 1930.

Diagnosis. — The cliaractcrs that at tliis time appear to (iistiii^'uisii
Testudo praeextans arc a.s follows: Cara{)ace depressed, with Jhitteiied
toj) in the vertebral region; prominent epiplastnd lip, project iiig
beyond the borders of the curai)ace and with j)arallel sides; sulci
usually shallowly impressed and usually ending, on free borders, in

'.Mr. fJilmorc died on 8vptomh«r 27, 104fi.— Ki>.

873370— IC 1 -'03

294 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

a projecting point or miicro; neurals variable but usually less dif-
ferentiated than in most species of the genus; first neural longest
of the series; entoplastron pointed in front, rounded behind; gular
scutes constantly encroaching on the entoplastron ; suprapygal usually
in contact with the eleventh peripheral only, rarely reaching the
pygal; median longitudinal ridge in front of the choanae.

Figure 20. — Skull of Testudo praeextans Lambe (U.S.N.M. No. 15874), superior view:
exoc, exoccipital; /, frontal; /, jugal; mx, maxillary; occ, occipital condyle; op, opisthotic;
p, parietal; pf, prefrontal; pmx, premaxillary; prf, postfrontal; so, siipraoccipital; sq,
squamosal. Natural size.

Materials. — ^The four specimens pertaining to Testudo praeextans
are all from Niobrara County, Wyo., and from the Brule division of the
Oligocene. The best-preserved specimen, U.S.N.M. No, 15874, col-
lected in 1932, consists of the complete carapace, plastron, skull,
lower jaws, pectoral and pelvic girdles, both humeri, both ulnae, one
radius, both femora, both tibiae, one fibula, and parts of all four feet.
It was found on the Anderson Ranch on the south side of Young
Woman Creek; specimen U.S.N.M. No. 15878, collected in 1932, was
found on the east side of Little Indian Creek and consists of the com-
plete shell, parts of both humeri, both ulnae, both radii, one articulated
forefoot, and portions of the other three forefeet; specimen U.S.N.M.

OSTEOLOGY OF TESTUDO PRAEEXTANS — GILMORE

295

No. ir»72S was foiiiul in 1942 about a inik" north of Wliitnian Post-
oflice and consists of carapace, plastron, hunuMus, both scapulae, both
coracoids, both tii)iao, both fibulao, tarsals, and sonic foot bones;
specimen I'.S.X.M. Xo. 10)732 was coUccti'tl in 1942 on the Anderson
Kanch on the nortli side of Young Woman Creek and consists of
carapace, })lastron. pelvis, two humeri, coracoid, two scapulae, tibia,
two ulnae, two radii, parts of both fibulae, and foot bones.

DESCRIPTION

Skull. — Of the 48 or more species of Testudo described from North
America, the skulls of only sLx are known at the present time. These
are Testudo thom.soni Hay, T. peragrans Hay, T. oshorniana Hay, T.
impensa Hay, T. orthopygia (Cope), and T. gilberti Hay; and only one
of these, T. thomsoni, is from the Oligocene. To this list we may now
add T. jiraeertans Lambe, represented by an unusually well preserved
skull and lower jaws belonging to specimen U.S.N.M. No. 15874 and
illustrated in three views in figures 20-22.

Figure 21. — Lateral view of skull and lower jaws of Testudo praeextans Lanibc (U.S.N.M.
No. 15874): an, angular; ar, articular; c, coracoid; d, dentary; /, frontal; /, jugai; mx^
maxilla; occ, occipital condyle; p, parietal; pf, prefrontal; p7nx, premaxillary; prj, post-
frontal; qu, quadrate; sa, surangular; so, supraoccipital; sq, squamosal. Natural size.

The skull and lower jaws of this specinicn arc practically complete
and undistorted, with numy of the cranial sutures clearly distinguish-
able (fig. 21). Th(! skull is of moderate size. Its length, from the tip
of the premaxillary to the occipital condyle, is 81 mm.; its width across
the sf|UHinosals is 57 mm.; the height from tiie cutting edge of the
mu.xiila to the upper surface of the frontal is 2\\ mm. 'V\\v. sides of the
skull forward to the backs of the orbits arc ncniiy parallel; beyond
here tliev regtdarly converge to the broadly rounded snout. Viewed
from the side tiie upper outline of the skull is nearly straight to a i)oint
above the anterior borders of the orbits where the prefrontals bend
strongly downward.

296

PROCEEDINGS OF THE NATIONAL MUSEUM

The prefrontals meet along the median line, a distance of 16 mm.,
and this is also the length of the suture between the frontals. The
lateral angles of the skull extend backward to a point slightly posterior
to the occipital condyle. The orbit has a greatest anteroposterior
diameter of 23 mm.; the nasal opening is 17 mm. wide; the interorbital
space is 22 mm. wide.

Viewed from above, the skull of Testudo praeextans has its closest
resemblances in the cranium of the Miocene T. osborniana, differing

Figure 22. — Palatal view of the skull of Testudo praeextans Lambe (U.S.N.M. No. 15874):
bo, basioccipital; bsp, basisphenoid; ect, ectopterygoid; mx, maxillary; occ, occipital
condyle; pi, palatine; pmx, premaxillary; pt, pterygoid; qu, quadrate; so, supraoccipital;
sq, squamosal; v, vomer. Natural size.

in its smaller size, the more abruptly truncated snout, the more
bluntly pointed squamosal region, the longer supraocciptal process,
and the straighter cutting edge of the maxillary. The orbit also has
a greater anteroposterior diameter, being one-fourth the over-all
length of the skull. The anteroposterior extent of the otic region
measured across the paraoccipital and the prootic is 22 mm.

The incomplete skull of Testudo thomsoni, the only other Oligocene
species in which the cranium is now known, differs from the skull

OSTEOLOGY OF TESTIDO PHAKKXTANS — GILMORE 297

before me in hav^ing shorter prefontals, smaller frontals, and longi-
tudinal channels on the masticatory surfaces unequal in width.
Wlii'ihcr those diU'eronces constitute stable characters that can be
relied upon for their specific separation cannot be determined until
more skulls ai-e available.

The roof of the mouth is moderately vaulted and traversed along
the midline by a sharp ridge (fig. 22), the anterior end of which abrupt-
ly widens where the vomer articulates with tiie premaxillaries and

Figure 23. — Lower jaw of Testudo praeextans Lainbe (U.S.N.M. No. 15874), superior view:
ar, articular; c, coronoid; d, dentary; prar, prearticular; sa, surangular. Natural size.

maxillary bones. Posteriorly this vomerine ridge underlaps the
pterj-goid part of the ridge, which merges into the palatal surfaces
of the pterygoid slightly in advance of their median union with the
basisphenoid.

The masticatory surface of the ma.xillary is divided by a low metlian
ridge that is received in a groove of the mandible. This ridge is
bordered on each side by longitudinal grooves of about equal width.
In this respect this specimen differs markedly from Testudo thomsoni,
w liich has a very narrow inner groove and a widened out(^r groove.
The imier ridge is unusually low, and it meets its fellow of the opposite
side on the posterior border of the premaxillary. At the midline
along th(! symphysis of the premaxillae there is a prominent longi-
tudinal ridge, as in Oopherus and in some living species of Testudo.
This ridge is absent in the skulls of other American fossil Testudo
now known. It was the sup[)osed absence of this ridge in front of the
choanae that led Hay to use it as one of the important characters for
distinguishing Testudo from Stylemys}

' Hay. O. ['., Fossil turtUs of .North Amcrlcn. rarncRlc trust. Washington Publ. 75. p. 307, \W».

298 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

The lower jaw, as in Testudo generally, is massive and heavy. Its
length from the symphysis to the angle is 59 mm., height at the
coronoid process 59 mm., height at the symphysis 10.5 mm. The
masticatory surface is traversed longitudinally by a deep groove.
Inner cutting edge of the mandible is nearly as high as the outer.
Inner cutting edge of the rami is separated by a notch at the sym-
physis; cutting edge not denticulated. The extent of the several
elements forming the mandible is clearly shown in figures 21 and 23.

Figure 24. — Carapace of Testudo praeextans Lambe, type (C. N. M. No. 8401): 1, 2, 3, 4,
5, 6, 7, 8, neurals 1-8, respectively. One-sixth natural size.

Carapace. — In the general contour of the carapace and plastron,
but more especially in the depressed character of the shell as a whole,
all these specimens are in close agreement with the type of the species.
The carapaces of all are flattened on top in the vertebral region and
when viewed from the side present a nearly straight profile except
where they curve downward at the ends. This depression of the cara-
pace, with the exception of Testudo laticunea Cope, appears to dis-
tinguish this species from all other North American members of the
genus. Among the large land tortoises only Testudo grandidieri
VaUlant, of northern Madagascar, has a somewhat simUar depressed
and flattened shell.

The four specimens under discussion show a considerable difference
in size of the shell, as indicated in table 1.

OSTEOLOGY OF TESTl'DO PR.\KEXTANS — GILMORE

299

T.\BLE 1. — Comparative vieaaurements of carapaces of Testudo praeextans

Specimen

Greatest
length

Oreafest
width

Greatest
height

Sex

Width to
length

C.N.M. No. 8401 (type).--

U.S.N.M. No. 15878-

r.S.N.M. No. 16728

r.S.N.M. No. 16732

r.S.N.M. No. 15874

Mm.
479
485
525

' 497
540

Mm.
410
392
438
450
452

Mm.
120
140
184
182
160

Female

Male

Percent
85
80
83
90
83

• Shortened by crushing.

Tlic larger size of the National Museum specimens as compaied
with the type may be accounted for partly by difference in age and
l)artly by sex. The hollowcd-out plastra of the National Museum
s])c'cinu'ns clearly indicate them to be males, whereas the flattened
})lastron of the typo of T. praeextans shows it to be a female. Among
living land tortoises it is a well-established fact that the males of a
species attain a larger size than the females. Furthermore, that the
larger specimens (U.S.N. M. Nos. 15874 and 16732) are aged individ-
uals is shown by the strong incurving of the pygal region (pi. 40, fig.
1 ; pi. 41, fig. 2), for it has been observed by Lord Rothschild,^ in his
study of the Galdpagos tortoises, that "very old individuals of both
sexes show an inclination often very strong, for the supracaudal to
curve round towards and even under the posterior end of the
plastron."

Viewed from above, all the specimens (pis. 38, 39), including the
type, are very similar in having their anterior borders broadly rounded
from side to side; the breadth of the shell decreases more rapidly to-
ward the front than toward the back, and so the posterior half has a
squarer outline than the anterior half. This is brought about chiefly
by the enlargement of the ])eripherals. The posterior border may i)e
described as angularly lounded, though this contour dilFers c<ui-
siderably with the age of (Ik; individual. The oldest s])ecimens
(Nos. 15874 and 10732) are more squarely tniiicate across the pygal
region tlian the otlieis, owing to the do^viiward and f(tI•^\aI•d deflection
(»f this ini(l])ortion of the shell.

The anterior peripherals in all specimens agree in being produced al-
most horizontally forward; the hiteral ones continue the geiu'ral con-
vexity of the shell downward, whereas those more posteriorly Mare
outward and somewhat njjward above the o[)enings for the posterior
extremities.

The nuchal, as shown by the measurements in table 2, is fairly con-
stant in its extent and proportions. The same observation may be
nuide of the pygal, suprapygal, and second su])rapygal.

1 liothschlld. I-. W., Nov. Zof.l.. vol. 22. i>. 42N, 1015.

300

PROCEEDINGS OF THE NATIONAL MUSEUM

Table 2. — Comparative measurements {in jnillimeters) of nuchals, pygals, and
suprapygals of Testudo praeextans

Nuchal

Pygal

Second
suprapygal

First
suprapygal

Specimen

Maxi-
mum
length

Maxi-
mum
width

Maxi-
mum
length

Maxi-
mum
width

Maxi-
mum
length

Maxi-
mum
width

Maxi-
mum
length

Maxi-
mum
width

C.N.M. No.8401 (type).
U.S.N.M. No. 15878_.
U S N M No. 16728

102

107

102

103
103
113
113
113

70
66

62
76

82
84
83

84
81

102

U.S.N. M. No. 16732._
U.S.N. M. No. 15874_.

50
49

""'74'

87
100

118

Table 3. — Comparison of shape of neurals of Testudo praeextans

Neural
No.

C.N.M. No.8401
(type)

U.S.N.M.
No. 15878

U.S.N.M.
No. 16732

U.S.N.M.
No. 15874

U.S.N.M.
No. 16728

1
2
3

4
5
6
7
8
9

Ovate

Hexagonal

Quadrangular
Hexagonal- -.

Ovate

Octagonal

Quadrangular

Octagonal

Hexagonal

do..

Ovate

Octagonal

Quadrangular
Hexagonal- --

Hexagonal- --

Quadrangular
Hexagonal- --

Hexagonal

Do.
Quadrangular
Hexagonal

Do.

The free border of the pygal in specimen No. 15874 differs from the
others in being strongly toothed. The bifurcated first suprapygal
in the type and in specimen No. 15874 is in contact with the pygal
and eleventh peripheral, whereas in the other three specimens it
articulates only with the eleventh peripheral.

In the shape of the neural bones three features are found common
to all available specimens: (1) The first neural is the longest of the
series; (2) the thii'd neural is always quadrangular; (3) the fifth to
eighth neurals, inclusive, are hexagonal. One individual, No. 15878,
has two octagonal neurals, the second and fourth; one. No. 16732,
has only the second neural octagonal; and No. 16728 has none octago-
nal. In most species of Testudo there are two octagonal neurals, the
second and fourth, but a series of specimens of other species might
show them to be equally variable as in T. praeextans. The variation
in the form of the neurals in T. praeextans is shown in table 3.

In discussing the neurals of T. laticunea, Hay^ says, "The nem-als
have not attained so high a degree of differentiation of form as they
have in most of the species of the genus." If reference is made to the
absence of octagonal neurals the statement is true of the types of

3 Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 403, 1908.

OSTEOLOGY OF TESTUDO PRAEEXTANS— GILMORE 301

both T. laticunea and T. -praeextans. In my judgment the dirt'orence
found in the neural bones of the specimens under consich-ration repre-
sents individual variations well within the species.

Specimen U.S.N.M. No. 15878 (pi. 38, fig. 2) has an extra neural,
or nine in all. It is quite evident from an examination of the speci-
men that the extra element has ileveloped between number 7 and the
last, which is without corresponding costals. Lambe * has described
a greater deviation of carapace structure in a specimen of Stylemys
nebrascensis, which has not only an extra neural but also a ninth pair
of costal bones and an additional vertebral scute. Tliis same authority
points out that the type of Stylemys culhertsoni likewise has an addi-
tional neural.

The variation in shape of the neurals has brought about a corre-
sponding variation in the proportions of these bones, as shown in table
4. In Tf.stvdo praeertans the first neural is consistently the longest of
any of the series.

The costal plates in all these specimens are highly modified, and as
usual in Testudo the second, fourth, and sixth are widened distally and
narrowed proximally, while the third and fifth are narrowed distally
and widened at the proximal ends.

The sulci in most of the specimens are narrow and shallowly im-
pressed, and where they reach the free edges of the peripherals there is
usually a small obtuse projection or mucro.

In the type of Testudo praeextans the sulcus forming the posterior
boundary of the third vertebral curves strongly forward at the center
where it crosses the fifth neural, as in U.S.N.M. Nos. 15878, 16728, and
1G7.32. In specimen No. 15874, however, the sulcus crosses the sixth
neural as in the type of T. laticunae Cope.

The proportions of the vertebrals are variable, as shown in tal)le 5.
In the type of 7'. laticunea and No. 15878 the fifth vertebral is the
longest of the series, whereas in No. 15874 the first is the longest.
In the type of T. laticvnea and No. 15874 the first vertebral is longer
than wide, but in the type of T. praeexiuns and No. 15878 this same
vertebral is wider than long. No. 15874 is the only specimen having
the third vertebral longer than wide.

The most distinctive feature of the plastron is the extended epiphis-
tnd lip that always projects well Ix'vond the line of the front margin of
the carapace. It was the "marked prominence and size of the
epiplastral lip" that Lanibe' stressed as the most important character
for distinguishing Testvdo praeextans, a character to which the specific
name refers. In the light of these additional spc'cimens it is clearly
shown that the extent and shape; of this lip constitute one of the more
stable characters of this species.

< I-ambp. L. M., Ottawa Nat., vol. 27, p. W, 1013.

• LanilH!, L. M., Ottawa Nat., vol. 27, pp. 57-fil, pl«<, 1. o. UM.1.

302

PROCEEDINGS OF THE NATIONAL MUSEUM

The relative proportions of the lobes and other measurements of the
plastron are clearly shown in table 6.

Table 4. — Comparative measurements (in millimeters) of neurals of Testudo

praeextans

Specimen

Length

Width

C. N. M. No. 8401 (type)

U.S.N.M. No. 15878

U.S.N.M. No. 16732

U.S.N.M. No. 15874

U.S.N.M. No. 16728

68
62
67
76
76

40
42
46
48
43

41
37
43
44
42

37
39
41
39

36
36
55
42

34
34
36
34

41
38
35

15
46
51

44
33
34
46

47
42
49
51
55

34
40
41
39
43

49
42
54

45
51

44
50

58
46

44
49
57
55

35
38
53

Table 5. — Comparative measurements (in millimeters) of vertebrals of Testudo

praeextans

Specimen

Length

Width

C. N. M. No. 8401 (type)

U.S.N.M. No. 15878

U.S.N.M. No. 16728 _ _

100
100
103
120
122

77
82
91
97
109

78
81
86
90
112

97
110
113
114
100

82
79
91
91
86

103

101

16?

103

105

170

U.S.N.M. No. 16732

U.S.N.M. No. 15874

109

106
117

97
89

211

188

Table 6. — Comparative measurements (in millimeters) of plastra of Testudo

praeextans

Measurement

C.N. M.
No. 8401

U.S.N.M.
No. 15878

U.S.N.M.
No. 16732

U.S.N.M.
No. 15874

U.S.N.M.
No. 16728

Greatest length of plastron

Length of anterior lobe

Width of anterior lobe .

493
163
229
147
239
72
97

495
154
205
135
225
60
81

47
34

540
178
258
145
245
71
95

72
39

580
187
243
155
270
85
102

88
48

191

Length of posterior lobe

Width of posterior lobe

Length of epiplastrai lip

Width of lip at base , _

76
97

Extension of epiplastrai lip be-
yond border of carapace. _

Depth of posterior notch

The free borders of the plastra in all five specimens are acute,
except those parts of the borders adjacent to the notches, which
are thickened and rounded. The posterior lobes are terminated
behind in two broadly rounded apices separated on the median line
by wide V-shaped notches. The borders of the apices are usually
slightly toothed. Likewise, the anterior ends of the epiplastrai lips
are notched at the center and the borders are toothed with six blunt
teeth, except in the type of Testudo praeextans, as shown in figure 25.

OSTEOLOGY OF TESTIDO PHAEEXTANS — GILMORE 303

The entoplastra in all the specmiens except No. 15878 are in
agreement in having pointed anterior ends and a broad somewhat
rounded posterior border, with the gular scutes overlapping the
front of this bone.

There is some variation in the extent of the plastral scutes in these
specimens, as shown in table 7.

Pelvis. — The pelvis of specimen No. 15874 is in an unusually perfect
state of preservation, the two halves being coalesced along the median

Figure 25. — Plastron of Testudo praeexlans Lambc, type (C. N. M. No. 8401): ab, abdominal
scute; an, anal scute; ent, entoplasiron; /^m, femoral scute; g, gular scute; hum, humeral
scute; hyo, hyoplastral bone; hypo, hypoplasiral bone; pec, pectoral scute; xiph, xiphi-
piastral bone. One-sixth natural size.

line, as reprcscntt-d in figures 20 and 27. The i)elvis of No. 107i{2
is also well preserved, but the two halves were separated along the
median suture. Except for minoi- details these two pelves ai-e in
close agreement and thus are probably tyi)ical of Testudo pratcxtaiis.

Of these two turtle specimens of nearly equal size and presumably
of equivalent age om(> has the pelvic sutures coalesc(>d, while in
the other these same sutures are open, thus indicating that the
relative age of an individual cannot always be judged by the non-
coalescence of the sulural junetures.

The anterior extremities of the pui)es are complete in specimen
No. 15874, which shows that border to be broadly rounded, thus

304

PROCEEDINGS OF THE NATIONAL MUSEUM

suggesting that Hay may have erred m restoring this missmg end
in the pelvis of the type of T. laticunea as being bilobed.® The un-

FiGURE 26. — Pelvis of Testudo praeextans Lambe (U.S.N.M. No. 15874), viewed from above:
il, ilium; is, ischium; pu, pubis. One-half natural size.

Figure 27. — Pelvis of Testudo praeextans Lambe (U.S.N.M. No. 15874), viewed from the
left side: il, ilium; is, ischium; pu, pubis. One-half natural size.

usual length of the anterior pubic process, forward of the lateral
pubic tuberosities, appears to be a distinctive feature of this species.
The apices of the ischial tuberosities are bluntly pointed and 80 mm.
apart.

« Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 404, flg. 512, 1908.

OSTEOLOGY OF TESTUDO PRAEEXTANS — GILMORE

305

Careful comparison made of the other skeletal parts preserved
with these various specimens, with each other and with homologous
bones of other species of Testudo, failed to disclose, except that of
size, differences that would be of assistance in specific differentiation.
On that account there seems do point in illustrating or describing
these parts here, although most of the bones present are in excelhMit
preservation. In order, however, that there may be a record of rela-
tive proportions between carapace and limb bones, a table of measure-
ments of the more important elements of the specimens here dis-
cussed is given (table 8).

Table 7.

-Comparative measurements {in millimeters) of plastral scutes of Testudo
praeextans

Scute flength on the midline)

C.N.M.

No. 8401

(type)

U.S.N.M.
No. 15878

U.S.N.M.
No. 16732

U.S.N.M.
No. 15874

U.S.N.M.
No. 16728

Gular -

80
103

43
147

162"
44

143
72
51

H umeral

Pectoral

Abdominal

135"
63
50

Femoral

Anal

Table 8. — Comparative measurements (in millimeters) of girdle and limb bones of

Testudo praeextans

Measurement

U.S.N.M.
No. 15878

U.S.N.M.
No. 16732

U.S.N.M.
No. 16728

U.S.N.M.
No. 15874

Greatest lenp;th of coracoid

Greatest width at inner end

Greatest length of scapula from tip to tip.

Greatest length of humeru.s

Greate.'^t length of ulna

Greatest length of radius

Greatest length of femur

Greatest length of tibia.

Greatest lengt h of fibula

88
77
153
155
99
98

151

153

77
80

1 150

157
99
97

113
82
82

I Estimated.

Feet. — Ail four of the National Museum specimens considered in the
I)receding pages have various parts of the carpi, tarsi, and feet present,
l>Mt only in No. 15878 arc the bones preserved at all in articulation.
Tlie right forefoot, with cari)us and radius, although lacking a few
elements, has the others articulated in normal relationship as siiown in
plate 44, figure 2. This foot clearly shows the presence of five well-
developed digits, each having a daw. The snudl blocklike bones of
the carpus are but little disturbed and can be quite certainly identified.
The ulnare and intermedium furnish the main articulation of the ulna,
which is missing. Occujjying the position of the radiale at th(> distal
end of the radius is an angular blocklike element that, following Baur

306 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

and Williston/ we may identify as centrale 1. Its outer side is in
juxtaposition to a smaller pentagonal bone regarded as centrale 2.
These two centralia although distinct elements in this individual are
often found fused into a single bone, and this appears to have happened
in specimen No. 15874. Williston, in the book cited, observes/' Among
terrestrial tortoises the radiale has disappeared until nothing is left of
it but a nodule of cartilage united with the first centrale which has
usurped its place." Hay, however, in his "Fossil Turtles of North
America," continues to regard the first centrale as the radiale.

The fifth carpale (pi. 44, fig. 2, C5) is present and remains in
articulation with metacarpal 5. Carpalia 3 and 4 are missing. The
third digit lacks its metacarpal, and a phalange is missing from the
fifth digit; otherwise the foot is complete. Comparison of these
wrist and foot bones with those of the living Galapagos Testudo of
comparable size shows a close correspondence in form, as well as in the
arrangement of the individual elements.

The hind foot of No. 15878 contributes nothing new to our knowl-
edge of the pes of Testudo.

SUMMARY

In the original description of Testudo praeextans Lambe called
attention to the close similarity in the form of the epiplastral beak to
that of T. thomsoni but concluded that the "much greater propor-
tionate size of the epiplastral lip and differences throughout of the
elements forming the lobe" were sufficient to indicate their specific
distinctness.

In view of the very fragmentary character of the type of T. thomsoni,
and especially of the considerable variation in the form of the anterior
lobe as shown by the present specimens, it would now appear that
Lambe was not justified in establishing the new species T. praeextans.
On plastral parts alone I should unhesitatingly regard T. praeextans to
be a synonym of T. thomsoni, which has priority by several years.
However, when the skulls are compared, differences in proportion of
the elements forming the skull roofs, and the different widths of the
channels on the triturating surfaces, strongly suggest that the dis-
covery of more complete materials of T. thomsoni may disclose other
and more important distinctive characters. For the present, there-
fore, it seems desirable to continue the use of both names.

In the preliminary study of the present materials it was first
thought that these specimens could not be specifically distinguished
from Testudo laticunea Cope and that such differences as existed might
be attributed to the female sex of the type of that species. This idea
was abandoned, however, with the discovery that the Wyoming

' WUliston. S. W., The osteology of the reptiles, p. 179, 1925.

OSTEOLOGY OF TESTUDO PRAEEXTANS — GILMORE 307

spociinons all hail i)it)aclly roundtil aiitoiior carapace borders and
prominent epiplastral lips with parallel sides, as contrasted with the
more truncate carapace hoidi^r and wedge-shaped epiplastral lip in
T. lat'ivuiua.

This study of a number of Testudo specimens from a restricted area
antl apparently of a single species shows enough variation in shell
structure to cast much doubt on the validity of many described species,
especially those foundetl on fragmentary materials. Furthermore, it
raises the perplexing question as to what featvn'es are to be relied on
for specific difrcrentation. It has been demonstrated (pis. 38-41)
that the neural, costal, and other elements forming the carapace, as
well as the form and proportions of the vertebral and plastral scutes,
are seldom in themselves suflicientl}' constant in shape and proportion
to be relied on for specific designation.

^^^lether characters of the skull in fossil Testudo will be found more
stable can only be determined when a series of crania is available for
comparison, and at this time no such series exists. It is in the general
form of the carapace and plastron and especially in the development
of epiplastral beak that most reliance has been placed in diagnosing
the present species.

TESTUDO LATICUNEA Cope

Plates 42, 43

Testudo lalicunea Cope, Paleont. Bull. No. 15, p. 6, 1873. — Hay, Carnegie Inst.
Wa.shington Publ. 75, p. 402, pi. 67, figs. 1, 2, 1908.

Two specimens (U.S.N. M. Nos. 15854 and 16731) in the Oligocene
collections from eastern Wyoming are identified as pertaining to
Testudo laticunea Cope, the first recorded occurrence of the species
in this area. These specimens are remarkably alike (cf. pis. 42 and
43) both in size and structural detail, a condition calling for comment,
as anyone will agree who has had occasion to study a series of Testudo
specimens.

The type of T. laticunea is an essentially complete shell, with some
of the internal skeleton, and was found in the Oligocene badlands
along the head of Horse Tail Creek in northeastern Colorado. Two
other specimens, one from the Tiianotherium beds (Chadron) of South
Dakota and one from the Oreodon beds (Brule) of Siou.x County,
Nebr., were identified by Hay * as jx'rtaining to this species. These
together with the two in the U. S. National Museum comi)rise all
the known materials of the .species.

Specimen U.S.N.M. No. 15854 consists of a nearly complete cara-
pace and plastron (pi. 43), left humerus, proximal end of right ; left
ulna, radius, and part of forefoot; incom])letc right scai)ula, left

' Hiiy, o. v., CamoKie Inst. Wa.shlDKlon I'ubl. T.'i, \t\). IOt-40:., 1908.

308

PROCEEDINGS OF THE NATIONAL MUSEUM

coracoid; left femur, tibia, fibula, and portion of hind foot. It was
collected by C. W. Gilmore on the Thomas Ranch, Niobrara County,
Wyo., in 1932. Specimen U.S.N.M. No. 16731 consists of a complete
carapace and plastron (pi. 42). It was collected 1 mile northeast of
Whitman Postoffice, Niobrara County, Wyo., by George B. Pearce
in 1942. Both of these specimens are from the Brule division of the
Oligocene.

In size and general contour of the shell these specimens are in close
agreement with the type, as shown by the measurements given in
table 9.

Table 9.

-Comparative measurements {in millimeters) of carapace and plastron of
Testudo laticunea

Measurement

Type spec-
imen

U.S.N.M.
No. 15854

U.S.N.M.
No. 16731

Greatest length of carapace _■

Greatest width of carapace

Greatest length of plastron

Gi-eatest length of anterior lobe

Greatest width of anterior lobe

Greatest length of posterior lobe

Greatest width of posterior lobe

Greatest width of bridge

Greatest width of anterior lip at gUlar notch.

408
356
440
132
200
120
200

436
332
440
132
202
110
193
200
90

435
340
443
137
203
110
196
197

Specimens U.S.N.M. Nos. 15854 and 16731 are practically free from
distortion and thus give a true picture of the normal shell. It will be
noted in the table of measurements that the Wyoming carapaces are
considerably narrower than that of the type. This difference may be
partly due to the crushing to which the type has been subjected. In
cross section at midlength the shells of Nos. 15854 and 16731 are evenly
rounded from side to side, whereas the type is said to be flattened on
top. All three, however, can be classed as having a depressed style of
shell, and all are of female sex, as shown by the flatness of their plastra.

Other minor differences observed between these two specimens and
the type fall well within the variations expected in individuals of a
single species.

The skeletal parts preserved with the shell of U.S.N.M. No. 15854
have been carefully compared with the homologous bones of T.
praeextans and other species of Testudo, but except for their smaller
size no other characters for distinguishing between them were dis-
covered.

Hay ^ has pointed out in his study of the type of T. laticunea that the
"neurals have not attained so liigh a degree of differentiation as they
have in most species of the genus." Although the complete neural
series cannot be traced out in either specimen, the correctness of the

» Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 403, 1908.

0STE01.0(.Y OF TESTUDO PUAEEXTANS — GILMORE

309

above coju'liision is verified ])y these new materials. S])eeinien No.
16731 np])enrs to sliow the seeoiitl neural to he oetagoual as it is in the
South Dakota specimen studied hy Hay.''

The slight variation in the vertehrals of the three s])eeiniens dis-
cussed here is shown by the measurements given '\u table 10.

Table 10. — Comparative measurements {in millimeters) of veriebrals o/ Testudo

laticunea

Specimen

Length

Width

Type specimen

86
78
91

80
89

81
83
78

75
83
78

90
88
90

102
94
94

82
74

85
82
78

82
'79'

1 :',:,

U.S.X.M. No. 15854

U.S.N. M. No. 16731

133
142

Table 11. — Comparative measurements (in millimeters) of epiplastral beak of Tes-
tudo quadra ta

Measurement

Type
(A.M.N.H.)

U.S.N. M.
No. 16737

Width of beak at base . .

120
70
29

135

Length of beak from gular groove

Thickness of beak at base .

The combination of characters that appear to distinguish Testudo
laticunea Cope is as follows:

Diagnosis: Carapace depressed with truncated anterior border, but
slightly rounded from side to side across the vertebral region; prom-
inent epiplastral lip, projecting beyond the borders of the carapace
with converging sides; neurals less diU'erentiated than in most species
of the genus, and usually w^ithout those of octagoiuil form.

TESTUDO QUADRATA Cop*
PlATK 44, FiGURK 1

Testudo quadratus Cope, The V'ertebrata of the Tertiary formations of the West,
p. 704, pi. 01, fiK. 5, 1884.

Testudo quadrata Hay, HiblioKraj)hy ami catalogue of the fo.s.^il V'ertebrata of
North America, p. 451, 1902; CarneRJe Inst. Washington I'ubl. 75, p. 410,
figs. 532, 533, 1908; Seconrl bibliography and catalogue of the fossil Verte-
bra ta of North America, vol. 2, p. 104, 1930.

A third species of 7'€.s<7/(/o occurring in the Indian Creek Hiisin area
is represented in the National Museum OligoccMu- ((tllcction by an
e])iplastral beak (U.S.N.M. No. 10737), shown in ])\iiiv. 44, figure 1.
This specimen was collertcd from the Brule, aixiut 1 mihr north of
W hitman Postoflice, Niobrara County, Wyo., in 1942.

» Ibid., p. 4fH.

310 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

The quadrate form of this beak with diverging lateral edges is more
like that of T. quadrata Cope than of any other species, but it differs
in having the gulohunieral sidci running backward and inward to
meet on the median line instead of directly across this bone at right
angles to the midline, as in the type of the species. This feature,
stressed by both Cope and Hay, is so at variance with all other known
species of Testudo, both living and extinct, that it leaves one wondering
if it is not an abnormal condition peculiar only to that individual.

Specimen No. 16737 is larger than the type, as shown by the
measurements given in table 11.

With the possible exception of length, the other differences observed
between the two specimens may be regarded as individual variations.
The beak is shorter than that of the type although exceeding it in all
other dimensions. Regardless of the question of doubt that this
difference may engender as to its proper specific assignment, this
fragmentary specimen certainly indicates the presence in the Brule
of the Hat Creek Basin of a third species of Testudo.

NOTE ON THE GEOLOGICAL DISTRIBUTION OF TESTUDO
IN NORTH AMERICA

In the "Fossil Turtles of North America," p. 397, 1908, Hay lists six
species of Oligocene Testudo as occurring in the Oreodon beds (Brule)
and two as coming from the Titanotherium beds (Chadron), as follows:

Brule Chadron

Testudo amphithorax Cope T. brontops Marsh

T. ciiltrnta Cope T. exornata Lambe

T. laticunea Cope
T. ligonia Cope
T. quadrata Cope
T. ihomsoni Hay

In a subsequent publication, "Second Bibliography and Catalogue
of the Fossil Vertebrates of North America," Hay assigned all the
Brule species to the Chadron. One is at a loss to understand whether
this change was made on the basis of new information or whether it was
a slip of the pen.

In view of the recognition of two, if not three, of the species in im-
doubted Brule deposits, it would appear to show the original geologic
assignment was correct. That some of these species may have per-
sisted from the Chadron into the Brule is suggested by a specimen in
the Yale Aluseum from the Titanotherium beds, which Hay '^ identified
as belonging to Testudo laticunea. That such was the case needs ad-
ditional verification. Tliis assertion is made on the fact that the
commonest of all land tortoises of the Brule, Stylemys nebrascensis, has
never been recognized as occurring in the Chadron.

11 Hay, O. P., Carnegie Inst. Washington Publ. 75, p. 404, 1908.

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issued

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM
VoL % Wuhington: 1946 No. 3200

EIGHT NEW SPECIES OF CHALCID-FLIES OF THE GE^^US
PSEUDAPHYCUS CLAUSEN, WITH A KEY TO THE
SPECIES

By A. B. Gahan

The species of chalcid-flies of tlie genus Psevdaphycus Clausen
(family Encyrtidae) are apparently all parasites of pseudococcine
scale insects, and some of the species are known to be of considerable
economic importance through the control they exercise over their
hosts. P. utilis Timberlake is credited with having all but exter-
minated Pseudococciis nipae (Maskell) in Hawaii, and the new species
malinius (described on a subsequent page of this paper), which was
introduced into the eastern United States from Japan to combat
Pncudoroccu.<i cowHtocki (Kuwana), apparently gives promise of
bringing that serious orchard pest under control.

Tlie genus is widely distributed. Species are now known to occur in
Austria, Spain, Canary Islands, Cuba, Puerto Kico, the United States
from New Jersey to California, Hawaii, Korea (Cho.sen), and Japan.

PypAidaphyffUA belongs to the group of genera related to Aphycus
Mayr. It may be distinguished from Aphycus as well as most of the
other related genera by the fact that it has only live, instead of six,
segments in the antennal funicle. It is most closely related io Both-
ri^orcwra Timberlake and A<-e,ropha(ni.f< Smith, both of which have
the funicle 5-s('grnented. It may be separated from linthriocrarra
by the dull sculpture of its frontovertex and tlioraric dorsum and
by the fact that it is never shining black in color. As pointed out by
Mercet (liull. Knt. Kes., vol. 2H, p. ;H7, 1!»;57), the dillerences between
Pftetidaphycus and AcerophnguJi are very slight. About the only real
differences seem to l)e that in Arerophng'>t-f( the front(»vertox is less than

311

705901— 4e

312 PROCEEDINGS OF THE NATIONAL MUSEUM tol. 9e

twice as long as wide, the ocellar triangle is more or less obtuse, and
the antennae are unicolorous and always pale, while in Pseiidaphycus
the f rontovertex is nearly always two or more times as long as broad,
the ocelli are arranged in an equilateral or acute triangle, and the
antennae are always contrastingly colored with the scape, pedicel,
and at least some of the funicular segments black or fuscous and the
club white.

The species of Pseuda'phycus offer very few good characters for
their separation. Slight differences in sculpture are hard to define
and differences in color consist largely of differences in shades of yel-
low, since all the species are yellowish sometimes varied with admix-
tures of blackish or fuscous. The following key to species is offered
as an aid to identification but should be supplemented by comparison
with types or authentically determined specimens whenever possible :

KEY TO THE DESCRIBED SPECIES OF PSEUD APHYCUS

FEMALES

1. Pedicel of antenna as long as or longer than first three segments of funicle

combined, funicular segments from first to last successively increasing in

width and all more or less transverse 2

Pedicel of antenna not longer than first two funicular segments combined;
first funicular segment small, transverse, narrower than pedicel ; segments
2 to 5 of funicle subquadrate, subequal, and each a little broader than
pedicel 1. meritorius, new species

2. Ovipositor exserted one-fifth to nearly one-half length of abdomen 3

OvijDOSitor extending barely beyond apex of abdomen, at most distinctly less

than one-sixth length of abdomen 2. prosopidis Timberlake

3. Wings hyaline; scape not expanded beneath 4

Wings at least faintly fuscous ; scape often but not always somewhat thick-
ened or slightly expanded beneath 5

4. General color pale lemon-yellow ; hairs on mesoscutum white; ocellar triangle

rather small and nearly equilateral; ovipositor exserted about one-third

length of abdomen 3. graminicola Timberlake

€reneral color bright orange-yellow; hairs on mesoscutum dark brown or
blackish ; ocellar triangle obviously acute and unusually large ; ovipositor
exserted about one-fifth length of abdomen 4. utilis Timberlake

5. Porewing fuscous with a more or less distinct paler hyaline or subhyaline

transverse streak beyond apex of stigmal vein; scape obviously a little

'''' thickened or broadened 6

S'',Forewing uniformly fuscous or subfuscous, without a paler transverse streak;

,.v scape usually slightly thickened but sometimes cylindrical 9

<5. Anterior ocellus at center of f rontovertex ; speculum of forewing not inter-
rupted; mesoscutum, axillae, and scutellum bright orange-yellow, with a
conspicuous narrow transverse band of black at anterior margin of meso-
scutum 5. websteri Timberlake

Anterior ocellus distinctly above center of f rontovertex ; speculum of forewing
interrupted a little behind middle ; mesoscutum, axillae, and scutellum more
or less dusky orange to blackish yellow, anterior margin of mesoscutum with
suture sometimes blackish but never forming a conspicuous and well-
defined transverse band 7

CHALCID-FLIES OF GENUS PSEUDAPHYCUS — GAHAN 313

7. Mesoscutum, axillae, scutellum, and abdomen blackish yellow; lateral borders

of mesoscutum white; posterior ocelli as far from anterior ocellus as from
margin of occiput; Inner orbits parallel for whole length of froritovertex.

6. maculipennis Mercet

Mesoscutum, axillae, and scutellum dusky orange-yellow; lateral borders of

mesoscutum not white; posterior ocelli a little more distant from anteriof

ocellus than from margin of occiput; inner orbits diverging very slightly

below 8

8. Frontovertex about two and one-half times as long as broad ; head and thorax

dorsally rather pale orange-yellow, axillary sutures of mctanotum usually

not blackish 7. angelicus (Howard.)

Frontovertex fully three times as long as broad; head and thorax dorsally
a deeper shade of orange-yellow mixed with blackish, and axillary sutures
brownish black 8. abstrusus, new species

9. Middle and hind tibiae each with two fuscous or blackish bands 10

Middle and hind tibiae without fuscous bauds 12

10. Frontovertex as broad as long or somewhat broader than long; ocelli in an

equilateral triangle; dorsum of thorax dirty yellow; antenna fuscous with

first funicular segment and club white 9. austriacus Mercet

Frontovertex twice to two and one-half times as long as broad ; ocellar triangle
at least slightly acute; dorsum of thorax orange-yellow, antennal scape
yellowish below, brown above; pedicel and funicle brownish or fuscous,
fifth funicular segment and club white 11

11. Ocellar triangle distinctly acute, posterior ocelli about half as far from each

other as from anterior ocellus ; speculum of forewing interrupted below

middle 10. orientalis Ferrlere

Ocellar triangle nearly equilateral ; posterior ocelli only slightly closer to
each other than to anterior ocellus ; speculum of forewing not Interrupted.

11. malinus, new species
12 Last two segments of funicle (sometimes entire funicle) and club white,
first three segments of funicle usually fuscous; frontovertex unusually
narrow, fully three times as long as broad ; ocelli in a very acute triangle,
posterior pair separated by about diameter of an ocellus ; ovipositor one-
third to half as long as abdomen 12. angustifrons, new species

Entire funicle black or at least fuscous; frontovertex at least a little broader;
ocellar triangle less acute and posterior pair of ocelli separated by more
tlian diameter of an ocellus; ovipositor usually shorter 13

13. Mf?«o8cutum, scutellum, and axillae weakly sculptured, distinctly shining;

dorsum of thorax dirty yellowish or grayish in color ; antennal club blackish

basal ly 14

Mesowutura, scutellum, and axillae (Wiely and densely Hcul[)tured, dull ; dorsum
of thorax orange-yellow; antennal club entirely white 15

14. Scape of antenna obviously a little thickened or expanded beneath ; ovipositor

exserted one-third h'ngth of abdomen ; 8p<'culuni of fonnviug uninterrupted.

13. mundus, new sp«'cle»*

Scape of antenna not obviously expandt-d ; f)vlposltor about on<!-!lfth length

of abdomen ; speculum at least partially Interrupted a little behind middle.

14. liinatulus, n«'w six-cles

15. Anterior margin of face narrowly dark brown or blncklHh, forming a dark

transverse line between bases of mandibles; occiput exct'pt around margins
black or blackish, anterior face of pronotum mostly black; suture between
pronotum and meso.scutum blackish; tegula pale at base, blackish aplcnily;
abdomen dorsally mostly blackish; posterior tibiae often with two fusrons
bands 11. malinus, new siM'cles

314 PROCEEDINGS OF THE NATIONAL MUSEUM vol. »«

Anterior margin of face without a dark line between bases of mandibles;
head and thorax dorsally bright orange-yellow without any blackish mark-
ings; tegula white; abdomen with four rather definite dark transverse
bands, one of which connects vibrissal plates, one basad of this and two

apicad of it; hind tibiae not banded 16

16. Frontovertex bright orange-yellow and very finely granulosely sculptured
with only very faint indication of alveolation even under high magnifi-
cation 15. meracus, new species

Frontovertex paler, dilute orange-yellow, entire surface covered with fine
but very distinct facetlike punctures 16. alveolatif rons, new species

1. PSEUDAPHYCUS MERITORIUS, new species

The subquadrate second to fifth segments of the f unicle distinguish
this species from all other species treated here.

Female. — ^Length 0.92 mm. Frontovertex, mesoscutum, axillae,
and scutellum slightly dusky orange-yellow ; head (except frontover-
tex), tegulae, pronotum, all pleura, and sterna and all legs yellowish
white; propodeum fuscous; abdomen dorsally yellowish with four
transverse fuscous bands, the anterior two bands coalesced at the
lateral margins of abdomen and enclosing an ellipsoidal area between
the cerci, the posterior two bands straight ; abdomen beneath fuscous
to blackish medially with the margins pale, ovipositor pale at base,
darker apically ; eyes black ; ocelli dark red ; antennal scape pallido-
fuscous; pedicel and entire f unicle black or blackish; club white;
wings uniformly faintly fuscous, or subhyaline ; marginal and stigraal
veins dark fuscous, submarginal vein pale.

Frontovertex a little less than twice as long as broad, granulosely
punctate; inner orbits parallel; ocelli in an equilateral triangle;
anterior ocellus situated only slightly above middle of frontovertex;
lateral ocellus less than its own diameter from margin of eye and
about its own diameter from occipital margin ; eyes clothed with short
hairs. Antenna clavate; scape not expanded beneath, five to six
times as long as thick; pedicel nearly twice as long as thick and less
than one-third as long as scape; first funicular segment small, nar-
rower than pedicel ; segments 2 to 5 of f unicle subequal and subquad-
rate, and each a little thicker than pedicel; club a little thicker than
funicle and aipproximately as long as the four preceding funicular
segments combined.

Mesoscutum, axillae, and scutellum finely granulosely punctate,
mat; propodeum in front of spiracle weakly reticulated, elsewhere
smooth ; pleura weakly reticulated, slightly shining. Forewing with
nearly uniform, rather dense discal ciliation ; speculum closed at pos-
terior margin of wing but otherwise uninterrupted; marginal vein
punctif orm ; postmarginal vein shorter than stigmal.

Abdomen short-ovate in outline, about as long as thorax or slightly
longer, slightly narrower than thorax ; weakly reticulately sculptured

CHALCID-FLIES OF GENUS PSEUDAPHYCUS GAHAN 315

and more or less distinctly shining dorsally and with similar sculpture
vent rally. Ovipositor exserted one-fourth to one-third the length
of abdomen.

}fale. — Length 0.8 mm. Similar to female except as follows:
Funicular segments subequal in length, but successively increasing in
width from first to last, the fifth about as broad as pedicel and a little
broader than long; club nearly as long as funicle; abdomen shorter
than thorax, nearly circular in outline, blackish dorsally and ventrally
with yellowish lateral margins above and beneath except at cerci,
where the black on dorsum extends to the margins.

Type locality. — Winchester, Va.

Type.—V. S. X. M. No. 57324.

lie marks. — Described from numerous specimens received from
D. W. Clancy of the Charlottesville, Va., laboratory of the Bureau of
Entomology and Plant Quarantine and said to have been reared from
Fseudococcus virgatus (Cockerell) collected at Winchester, Va., June
20, 1944, and recorded under laboratory note Nos. 67G, 677, 678, and
681. Other specimens identified as this species were reared from the
same host collected at Winchester, May 31, 1944, under laboratory No.
638; August 4, 1943, laboratory No. 529; August 3, 1944, laboratory
No. 850; and at Hamilton, Va., August 12, 1943, laboratory No. 530,
and May 30, 1944, laboratory Nos. 634 and 635.

2. PSEUDAPHYCUS PROSOPIDIS Timberlake
P-< udaphyrus prosopidin TiMiJFmL.\KK, Proc. U. S. Nat. Mus., vol. 50, p. 571, 1916.

Described from Mesilla, N. Mex., as a parasite of Fseudococcus
prosopidh ( Cockerell ) .

I. PSEUDAPHYCUS GRAMINICOLA Timberlake

Pfifudaphyrus graminUola Ti.mbeklake, Proc. U. S. Nut. Mus., vol. 50, p. 570, 1916.

Kecorded by Timberlake as parasitizing pseudococcine scales on
tStipa sp. at Las Vegas, N. Mex., and on Elymus condctu'<atus at Kim-
balls, Utah. The host insect involved in the record from Stipa sp.
has recently been ideiifified by Aforrison as Trionyiuu.s ncomcxicamus
var. utahercHis (Cockerell) . The host on fJhjnius remains unidentified.

4. PSEUDAPHYCUS UTILIS TlmbcrUke

pMMila^hycuH ufiliH TiMfJE:Ri..\KK, proc. Hiiwriiiaii Kiit. Soc, vol. ."">, p. .'{23, Akh. 1-4,
lfrj3.— SwcETi', Pr.tc. Hiiwaiiiui Kiit. Soc. vol. .1, p. »U, 11)2:{. — Ti.MitFniLAKK,
Proc. llawaiiiiri Eiit. Soc., vol. .""), p. 431. WS.i. — Williams. IlMinlborik of iii.vtrts
Jiiid other iiiv(Ttcl>ratt's of Hawaiian su^j.-ir cane flchlH. ]). L'ri.'i, lll.'n.- Swkzky,
Pror. nth Pafiflc Sol. Con^rr. (Canada, 19:{.3), vol. .'>, [». rj.WJ, H>34.— PrMBianoN,
Hawaiian Planters Hoc, vol. 45, p. lOO, 1J>41. — Anon., Puerto Hico Kxp. Stat.
Hrj.t. for imo. p. 68. lt)42.

This species is a parasite of PseudococoiM nipae (Maskell) and was
originally described from the state of Veracniz, Mexico. It was intro-

316 PROCEEDINGS OF THE NATIONAL MUSEUM vol. w

duced into Hawaii in 1922 where, according to Pemberton, it proved so
efficient as practically to exterminate its host. More recently it was
introduced into Puerto Rico where, according to the above-cited report
of the Puerto Eican Experiment Station, it gives every indication of
bringing the host insect under complete control.

5. PSEUDAPHYCUS WEBSTERI Tiraberlake
Pseudaphycus wetsteri Timberlake, Proc. U. S. Nat. Mus., vol. 5, p. 570, 1916.

Known only from the type, a single female, said to have been reared
from the stems of Elyrrms virginicus, at Villa Ridge, 111. The real
host was probably a pseudococcine scale on the Elymus, as Timber-
lake pointed out.

6. PSEUDAPHYCUS MACULIPENNIS Mercet

Pseudaphycus macuUpennis Mebcbt, Bol. Real Soc. Espan. Hist. Nat, voL 23,
p. 140, fig. 2, 1923; Rev. Espan. Ent., vol. 1, p. 12, fig. 1, 1925.

Originally described from material collected on the island of Tene-
riffe, Canary Islands. Mercet later collected it at Barcelona, Spain.
No host has as yet been recorded for the species.

7. PSEUDAPHYCUS ANGELICUS (Howard)

Aphycus angelicus Howard, Proc. U. S. Nat. Mus., vol. 21, pp. 241, 245, 1898.

Pseudaphycus angelicus (Howard) Timberlake, Proc. U. S. Nat. Mus., vol. 50,
p. 573, 1916.— Clausen, Univ. California Techn. Bull., Entom., vol. 3, pp. 258,
280, 1916.— Flanders, Journ. Econ. Ent., vol. 28, p. 552, 1935 ; vol. 33, p. 758,
1940.

P. angelicns was originally described from Los Angeles, Calif., as
a parasite of Pseudococcus sp. on passionflower. It has since been
recorded by Timberlake as parasitizing Pseiuiococcus {longispinus
Tsirgiom) = adonidum (Linnaeus), ryani (Coquillett), and {citro-
philus Clausen) = gahani Green; by Clausen from Pseudococcits
maritiinus (Ehrhorn) and gahani Green; and by Flanders from
Pseudococcus adonidu/m (Linnaeus) and Phenacoccus gossypii Town-
send and Cockerell. All these records are of occurrences of the species
in California, and thus far it is not known to occur outside of that
State.

Timberlake has given a satisfactory description of this species in
the paper cited above, except that it should be not^d that the trans-
verse fuscous band on the cheeks which he mentions is not always
present.

8. PSEUDAPHYCUS ABSTRUSUS, new species

The specimens to which this name is here assigned have proved
exceedingly puzzling (hence the specific name).

Female. — Length 0.94 mm. Apparently agreeing in every respect
with females of angelicus except that the frontovertex is very slightly
narrower than in that species, and the general color is somewhat

CHALCID-FLIES OF GENUS PSEUDAPHYCUS GAHAN 317

tiarker, dirty orange-yellow, with the anterior margin of mesoscutum
as well as the axillary sutures blackish and the metanotum, propodeum,
and entire dorsum of abdomen deeper black. The color of antennae,
uudeipnrts of the head and thorax, tegulae, wings, and legs is as in
angelicus.

Male. — Length 0.8 mm. The single male specimen available for
study has the antennal pedicel, first funicular segment, and the entire
club white, the remainder of antenna black. In angelicus the pedicel
is brownish, the first two segments of the funicle white, and the club
mostly white but with its base narrowly blackish. Otherwise I can see
no differences.

In both angelicus and ahutrvMis the f rontovertex is densely and very
finely alveolately sculptured, the alveoli very minute and discernible as
such only under high magnification. This sculpture is very similar to
tliat found in alvcolatifronx, new species, described beyond, but in that
species the aveoli are coarser and more distinct, the frontovertex is
broader and shorter, and the ocelli are arranged in an equilateral
instead of a distinctly acute triangle.

Type locality. — Roseland, Va.

ry;?e.— U.S.N.M. No. 57325.

Remarks. — The holotype female and one female paratype were col-
lected by D. W. Clancy, August 1. 1944, at Roseland, Va., on the bark of
an apple tree infested with mealybugs and bear laboratory No. 745.
The male allotype and one female paratype are labeled as having been
reared from Pscudococcus comstocki (Kuwana) collected Octx^ber 18,
1944, by Clancy at Colesville, Va., and bear laboratory Nos. 855 and
850, re.spectively.

». PSEUDAPHYCUS AUSTRIACUS Mercet

Ptcudnphycus austrincus Mfbct;t, Uev. Kspan. Eiit., vol. 1, p. 13, flR. 2, 1925.
The type locality is Weyer, Austria. No host is known.

10. PSEUDAPHYCUS ORIENTALIS Ferriere

I'sijudaphycus orimtalis Ffkrikke, Bull. Ent. Ros., vol. 28, p. 315, flg. 1, 1987.— liU
I'yAjj-jr, Trans. Roy. Ent. Soc. London, vol. 0.3, p. 82, 1043.

This species was originally de.scribed as a parasite of Pseudococcus
lilacinus (Cockerell) collected at Los Banos, Luzon, Philippine
Islands. Le Pellcy gives an account of its habits and biology.

11. PSEUDAPHYCUS MALINUS, new ipecifi

Aphvcvs Kp. Hakishi.kr and Clancy, Jotirn. Eoon. Ent., vol. 37, p. WH, 1944.

The description of orientalis Ferriere fits this species in many par-
ticulars but differs in respect to the ocellar triangle and the speculum
of the wing. In ornentali-s the ocellar triangle is said to be acute, "the
hind ocelli half as far from each other as from tlie front ocellus," and
the wing is said to have "the hairle.ss streak interrupted below the

318 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9»

middle." In malinics the ocellar triangle is somewhat acute, but the
posterior ocelli are only slightly closer to each other than to the front
ocellus and the hairless streak is not interrupted. One inconspicuous
but seemingly constant character distinguishing malinus from all
other species examined is that the clypeus has a very narrow brownish
or blackish anterior border which constitutes a nearly straight and
continuous dark line between the bases of the mandibles.

Female. — Length 0.9 mm. Frontovertex, mesoscutum, axillae, and
scutellum dirty orange-yellow ; temples, cheeks, and face below f rons
whitish except for a very narrow brownish or blackish line along the
oral margin between bases of mandibles ; occiput above the neck more
or less blackish ; antennal scape, pedicel, and f unicle dark brownish
or fuscous, the last funicular segment usually mostly whitish; club
white ; apices of mandibles dark brown ; eyes black ; ocelli dark red-
dish ; pronotum whitish above, its anterior face in large part blackish ;
anterior margin of mesoscutum along the suture narrowly blackish;
scutellum frequently with indications of an obscure brownish trans-
verse band before apex; tegulae pale basally, fuscous apically; all
legs and underside of thorax whitish, the middle and hind tibiae some-
times each with two more or less distinct fuscous bands ; propodeum
and dorsum of abdomen dark brown, the latter with a narrow trans-
verse band of blackish at about basal one-third ; under side of abdomen
blackish with broad whitish margins. Forewing subhyaline, with a
faint fuscous cloud behind the stigmal vein but without a hyaline
band.

Frontovertex about two and one-half times as long as broad,
granulosely sculptured; inner orbits parallel; ocelli in a nearly
equilateral triangle, the posterior ocelli slightly closer to each other
than to the anterior ocellus; ocellocular line equal to about half the
diameter of an ocellus; anterior ocellus situated distinctly above
middle of frontovertex; eyes clothed with short hairs. Antennae
strongly clavate; scape subcylindrical, approximately four times as
long as thick; pedicel about one-third as long as scape and subequal
to first four funicular segments combined; funicular segments all
transverse and successively increasing in width from first to last, the
fifth segment fully twice as wide as the first; club solid, distinctly
wider than last funicular segment and approximately as long as
pedicel and entire funicle combined.

Mesoscutum, axillae, and scutellum finely sculptured, mat, and
clothed with short pale hairs; propodeum very faintly reticulated,
shining; mesopleuron with very fine and weak reticulation. Forewing
with short dense discal ciliation distad of speculum, the cilia basad
of speculum longer and not quite so dense ; speculum closed at posterior
margin of wing by about three rows of hairs, otherwise not inter-

CHALCID-FLIES OF GENUS PSEUDAPHYCUS GAHAN 319

riipted; marjiinal vein piinctiforin; sti*rnial vtMii loiif^^T tliau marginal
anil postniarginal conibinod.

Abdomen abont as long and as broad as thorax, short-ovate to nearly
circular in out lino, weakly reticulated and more or less shining dor-
sally and vi'ntrally; ovipositor exserted approximately one-sixth to
one-fifth the length of abdomen.

Male. — Length 0.75 nnn. Like female in all respects except that
frontovertex is only about twice as long as broad, anterior ocellus
only slightly above middle of frontovertex, and ahdonun a little
shorter and narrower than thorax.

Type locality. — AVinchester, Va.

Type.—V.S.^.U. No. 57320.

Remarks. — The holotype female, allotyi)e, and a largvr number of
paratypes of both sexes were reared by D. AV. Clancy from Pseudo-
COCCU.S comstockl (Kuwana) collected August 11, 194->, at Winchester,
Va., and recorded under Charlottesville, Va., laboratory No. 557.

This species was introduced from Asia into Virginia and several
other eastern States in 1941-42 by the Bureau of Entomology and
Plant Quarantine to combat the Comstock mealybug. Numerous
specimens of the species reared from P. com-'^tockt collected by R. W.
Burrell and C. P. Clausen in Japan and Chosen have been examined
and compared with the types, but since none of this Asiatic material
is in fir.st-class condition the description has been drawn from speci-
mens representing recoveries of the species in Virginia. Since its
original introduction specimens have been distributed for colonization
in other localities, and the National Collection now contains material
representing recoveries at the following points: Batesville, Berryville,
and Hamilton, Va.; Proctorville, Ohio; South River, Morganville,
:in<l Moorestown, N. J.; Branfurd, Guilford, and Meriden, Conn.;
;iiid Bridgeville, Del.

12. PSEUDAPHYCUS ANGUSTIFRONS. new speoiia

Thf- unusually narrow frontovertex will distinguish tJiis species from
all others known to me.

Female. — Length 0.8 mm. Fi-ontovcrtex light orange; occiput more
or less blaekish above the neck: eyes black; ocelli dark red: remainder
of head whitish. Antennal scape and pedicel black or bhickish, the
scape usually pale at base and .sometimes with the dor.sal margin j)ale;
first three segments of funicle more or less fuscous, fourth and (iflh
M'gments white; club white. Pronotum whitish dorsally and laterally
but with anterior face mostly blackish; mesoscutinn, axillae, and
scutellun) brownish orange, with the sutures darker; propodeum dark
brown; pleura, .sterna, and all legs whitish; abdomen doisally dark
blown with whitish margins, ventrally entirely while. Wings uni-
formly faintly fuscous; marginal vein daik fuscous, other veins \m\W\\
TooOei 4»; —2

320 PROCEEDINGS OF THE NATIONAL MUSEUM vol. oe

Frontovertex unusually narrow, fully three times as long as broad,
gi'anulosely punctate ; inner orbits parallel or very nearly so ; ocellar
triangle strongly acute, posterior ocelli separated by a distance equal
to r.bout one to one and one-half times the diameter of an ocellus,
nearly twice as far from anterior ocellus as from each other, almost
touching the eye margins and a little more than the diameter of an
ocellus from the occipital margin; anterior ocellus situated a little
above the middle of frontovertex; eyes large, with very short and
inconspicuous pile. Antennae strongly clavate ; scape distinctly a 1 ittle
exj^anded beneath, widest beyond middle, very slightly less than four
times as long as broad; pedicel about one-third the length of scape
and about equal to first four segments of f unicle combined ; funicular
segments all wider than long, successively increasing in width from
first to last, the first segment narrower than pedicel, the fifth segment
the largest and about twice as broad as long; club solid, distinctly
broader than last funicular segment, and approximately as long as
pedicel and funicle combined.

Mesoscutum, scutellum, and axillae finely sculptured but not wholly
mat, clothed with short whitish hairs ; propodeum smooth and shining;
mesopleuron weakly sculptured. Forewing with speculum closed at
posterior margin of wing and also interrupted a little behind tlie
middle ; discal ciliation distad of speculum short and moderately dense,
proximad of speculum a little coarser and not quite so dense ; marginal
vein punctiform, postmarginal a little longer than marginal, stigmal
distinctly longer than marginal and postmarginal combined.

Abdomen as broad as thorax and usually a little longer than thorax,
ovate, weakly sculptured dorsally, smooth and shining ventraily;
ovipositor sheaths exserted about half the length of abdomen.

Male. — Length 0.7 mm. Closely resembling female but with fronto-
vertex only slightly more than twice as long as broad ; anterior ocellus
located very nearly at middle of frontovertex; abdomen no longer
than thorax and subcircular in outline.

In some specimens of both sexes the prescutum is narrowly bordered
anteriorly with black and in some males the ocellar triangle is blackish.

Type locality. — Cuba.

Ty/^e.— U.S.N.M. No. 57327.

Remarks. — Described from 26 females and 4 males, all intercepted
at quarantine, at three different ports of entry, on shipments of pine-
apples originating in Cuba and in each case infested with mealybugs
(probably Pseudococcus hrevipes Cockerell). In most instances the
parasites were actually reared from the mealybugs. The holotype
female and two female paratypes were reared at Detroit, Mich., April
18, 1932, by W. W. Wood, under Detroit No. 1094. The allotype and
three female paratypes were reared April 28, 1936, at New York, and

CHALCID-FLIKS OF GLNUS PSEUDAPHYCUS GAHAN 321

bear New York No. 58589. Other paratypes were intercepted May
13, 1D3G, uiKler New York No. 59153; Doceniber 28, 19;}L>, under New
Orleans No. 5953; June 3, 1934, under New Orleans No. 8715; April
>20, 193G, under New Orleans No. 17;V20; May 22, 1934. under New
Orleans No. 8-lSS; and April 28, 193r), under New York No. 58577.

18. PSEUDAPHYCUS MUNDUS, new species

I'seudococcobiun terryi (FuUaway) Bynum (not tenyi Fullaway), Jdui'h. Ec«>u.

Eiit., vol. 30, pp. 75G-7G1, 1937.
I'lciidaphynts sp. nov. Wolcott, Insectae Borlnquenses, pp. 128, .''i2!), 103U.

This species closely resembles angeUcus (Howard) but may be dis-
tinguished from that species by the fact that the forewing is more
weakly and more uniformly infuscated, without a transverse hyaline
bund, and the speculum is not interrupted. The female differs further
from angdicU'S by having the antennal club black or blackish at base
instead of entirely white. The male differs from the male of angelicus
by having the entire funicle of the antenna nearly uniformly pale
grayish instead of mostly black with the first two segments only pale.

Female. — Length 1.1 mm. Frontovcrtex, mesoscutum, axillae, and
scutellura dilute orange with a slight dusky tinge ; head except fronto-
vcrtex, tegulae, pronotum, entire underside of thorax and all legs
white or whitish; propodeum and dorsum of abdomen blackish,
abdomen beneath whitish; ovipositor pale at base, black at apex;
eyes brownish black, ocelli dark reddish; occiput immediately above
tlie neck more or less infuscated and suture between pronotum and
mesoscutum often fuscous; antennal scape black with a white dorsal
margin; pedicel, entire funicle, and usually the greater part of basal
segment of club black; last two segments and apex of first segment of
club pure while; wings uniformly faintly fuscous, almost hyaline.
Ilairs on mesonotum white.

Frontovcrtex two and one-half to three times as long as bioad,
granulosely punctate; inner orbits practically parallel; ocelli in a
slightly acute triangle; anterior ocellus situated distinctly above the
middle of frontovcrtex, lateral ocellus about its own diameti-r from eye
margin; eyes clothed with short pile. Antenna strongly clavale;
scape distinctly a little expanded beneath, about four times as long as
bnjad ; pedicel a little more than one-third as long as sctipe and nearly
equal in lenglii to four following segments of funicle; funicular seg-
ments all strongly tiansverse, sube(]ual in length but successively in-
cieasing in width, the fifth segment approximately twice as wide as the
first and three times as broad as long; club large, indistinctly 3-seg-
inented, bn^ader tlian last funicular segment, and as long as or a little
longer than entire funicle.

Mesoscutum, axillae, and scutellum weakly .sculptured, rather dis-
tinctly shining and sparsely haii'y; propodeum smooth: mesopleuron

322 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

uniformly weakly reticulated. Forewing Avith discal cilia basad of
speculum a little sparser and slightly longer than distad, the speculum
closed at posterior margin of wing by several rows of hairs but other-
wise uninterrupted; marginal vein a little longer than broad, post-
marginal distinct, about as long as stigmal.

Abdomen as long as head and thorax combined, as broad as thorax
at base and tapering to a point at apex, distinctly reticulately sculp-
tured over the whole dorsal surface, nearly smooth ventrally. Oviposi-
tor exserted one-third the length of abdomen.

Male. — Length 0.85 mm. Like female except as follows: Whole
antenna much paler in color, the scape whitish with a longitudinal
fuscous stripe, pedicel and funicle grayish white, and club white;
abdomen not longer than thorax, broadly rounded at apex, nearly
circular, blackish dorsally but with lateral margins broadly margined
with whitish.

7'ype locality. — Houma, La.

Type.—\J. S. N. M. No. 57328.

Remarks. — Described from numerous specimens received from D. W.
Clancy of the Charlottesville, Va., laboratory of the Bureau of Ento-
mology and Plant Quarantine, where they were reared from Pseudo-
coccus honinsis (Kuwana) collected at Houma, La., in November
1943. Besides this type series the following identifications of this
species have been made by me : One specimen labeled as having been
reared February 10, 1916, from Pseudococcus calceolariae (Maskell)
[ — misidentification of P. honinsis (Kuwana)] collected at Audubon
Park, New Orleans, La., by E. R. Barber ; three specimens reared in
1943 from Pseudococcus honinsis (Kuwana), collected at Houma and
Raceland, La., by E. K. Bynum; two specimens indicated as having
been reared from P. honinsis at Cairo, Ga., in 1932, by E, K. Bynum ;
two specimens taken at quarantine at Brownsville, Tex., on cut flowers
from Mexico; nine specimens reared from the pink mealybug of
sugarcane {THonymus sacchari Cockerell), December 28, 1932, at Rio
Piedras, Puerto Rico, by F. Sein, under P. R. Ace. No. 178-32. Also
before me are specimens reared in breeding tests at the Charlottesville,
Va., laboratory from the following hosts : Phenacoccus gossypn Town-
send and Cockerell, Pseudococcus comstocki (Kuwana), and P. adoni-
dvm (Linnaeus).

This species has become confused in the literature with Aphycus
terryi FuUaway {^Pseudococcohius terryi FuUaway). In the arti-
cle by E. K. Bynum cited above, it is indicated that living material
of A. terryi was received in 1932 by T. E. Holloway, of the Bureau of
Entomology and Plant Quarantine laboratory in New Orleans, from
the Hawaiian Sugar Planters' Experiment Station in Hawaii. The
original stock was apparently increased by propagation in the labora-

CHALCID-FLIES OF GENUS PSEUDAPHYCUS — GAHAN 323

tory. and during the same year releases are said to have been made
on three sugtircane plantations in Louisiana; at Cairo, Ga.; at Belle
Glade. Fla.; and a small shipment sent to G. N. Wolcott for release
in Puerto Rico. Other releases of the parasite are said to have been
made in 1933, 1934, and 1936 in several additional localities in the
three states mentioned. According to Bynum the parasite was re-
covered in the field of introduction at Houma in 1932 and each year
thereafter up to the time of publication. Recoveries were allegedly
made at several other points of introduction including Cairo, Ga., and
Belle Glade, Fla. At some points of introduction no recoveries were
made. According to Wolcott attempts in Puerto Rico to recover
terryi resulted only in the rearing from Trionymus saccJmri of speci-
mens which were identified b}- Muesebeck as a new species of the genus
Pseiidaphycus.

In 194:3 I received from D. W. Clancy, of the Charlottesville, Va.,
laboratory, a series of specimens of a parasite reared from Fseudococ-
OU3 honirusis that had been sent to him by J. W. Ingram, of the Bureau
of Entomology and Plant Quarantine laboratory at Houma, La.
These specimens had been reared from mealybugs collected in 1943
from fields in the vicinity of Houma and were supposed to be repre-
sentatives of the introduced Aphycus terryi. They proved not to be
A. terryi^ however, but the above-described new species, Pseudaphycus
inundus.

Following this discovery a request was made to the Houma labora-
tory for samples of the original shipment of parasites received from
Hawaii for introduction into Louisiana. Such a sample was received
and proved to be true A. terryi. Another lot labeled, "Cairo, Ga.,
1932, ex Psev/lococcufs honinsis E. K. Bynum SC #333," and appar-
ently constituting a part of the original release at that point, was also
received. This sample proved to be not terryi but Pseudaphycus
mundus. No representatives of the other releases were obtained, but
.specimens taken at Houma and Ilaceland, La., in 1943, supposedly
representing recoveries of terryi^ again turned out to be P. mundwi.
The specimens mentioned by Wolcott as having been taken in Puerto
Rico and identified by Muesebeck as Pseudapkycus n. sp. were located
in the National Museum collection and were also found to be the same
as those from Louisiana and Georgia.

Tlie above data show that the parasite received from Hawaii was
certainly Aphycus terryi but that this species has not been recovered at
any of those points of release frcjni which material has been submitted
for identification. All the alleged terryi reared from field a)llect ions
at Houma that I have seen have been the new species Pseudaphycus
munduji. Api)arently alsf) it was this sj)ocies, not terryi., which was
released at Cairo, Ga. Furthermore, it ap[)ears probable that this was

324 PROCEEDINGS OF THE NATIONAL MUSEUM vod. oe

the species introduced into Puerto Rico and that this introduction was
successful even though the species introduced was not the one intended.
Pseudaphycus mundus may be indigenous in Louisiana. As noted
above it was reared at Audubon Park, New Orleans, as early as 1916, at
least 16 years prior to the attempted introduction of Aphycus terryi.
Circumstantial evidence would seem to indicate that in the attempt to
increase the stock of the Hawaiian parasite in the laboratory, field-
collected material of the host which had already been attacked by
P. mundiis was introduced into the cages and the two species thus
became confused.

14. PSEUDAPHYCUS LDIATULUS, new Bpceies

The dorsum of the thorax in this species is more weakly sculptured,
more distinctly shining, and of a darker yellowish-gray color than in
any of the other species. Most closely resembles graminicola but
distinguishable by the above characters and its slightly narrower
frontovertex.

Female. — ^Length 0.75 mm. Frontovertex light orange-yellow ; occi-
put slightly fuscous; eyes black; ocelli reddish; remainder of head
whitish. Antennal scape and pedicel pale yellowish gray; funicle
black or blackish ; club mostly white, more or less stained with blackish
basally. Mesoscutura, scutellum, and axillae yellowish gray; prono-
tum whitish, with its anterior face stained with blackish on each side of
neck; propodeuni dark brown; pleura, sterna, and all legs whitish;
tegula whitish, its apex stained with fuscous ; abdomen dorsally mostly
dark brown or blackish, ventrally varying from mostly blackish to
mostly whitish with only the middle blackish. Wings nearly uni-
formly faintly fuscous with faint indication of a paler transverse band
beyond apex of venation. Ovipositor mostly dark brown, paler at
base.

Frontovertex about two and one-half times as long as broad, mat,
the sculpture made up of extremely minute alveolate punctures ; inner
orbits parallel for most of their length but diverging rapidly just
above the scrobe ; ©cellar triangle nearly equilateral, or slightly acute ;
posterior ocellus about half its own diameter from eye margin and
about its own diameter from occipital margin; anterior ocellus dis-
tinctly above middle of frontovertex; eyes with very short, incon-
spicuous pile. Antennae clavate ; scape not expanded beneath ; pedicel
approximately one-third as long as scape and about as long as follow-
ing four segments combined ; funicular segments all broader than long,
successively increasing in width from first to last, the fifth segment
about twice as broad as long; club solid, distinctly broader than last
funicular segment and only slightly shorter than funicle and pedicel
combined.

CHALCID-FLIES OF GENUS PSEUDAPHYCUS — GAHAN 325

Mesoscutuni, axillae, and scutelhim nearly flat, weakly sculptured,
distinctly shining, and clotlied with short pale pubescence; propo-
(hnirn smooth and shining: mesopleuron weiikly sculptured. Speculum
of forewing purtially interrupted near middle and closed at posterior
margin of wing b}' one or two rows of cilia; discal ciliation moderately
dense, very slightly coarser basad of speculum than distad of it; mar-
ginal vein punctiform, postmarginal short, stigmal about twice as
long as marginal and postmarginal combined.

Abdomen ovate, very slightly longer than thorax and slightly nar-
rower than thorax; tergites with distinct, shallow, reticulate sculpture,
somewhat shining; sternites practically smooth and shining; oviposi-
tor sheaths exserted approximately one-fifth the length of abdomen.

Male. — Length 0.G5 nnn. Similar to female but with frontovertex
only about twice as long as broad, mesoscutum along its anterior mar-
gin narrowly black, abdomen distinctly shorter and narrower than
thorax and circular in outline.

Type locality. — Stevensville, Kent Island, Md.

ry/>e.— U.S.N.M. No. 57;3i:i).

Remarks. — Described from 21 females (1 holotype) and 5 males
(1 allotype) reared by H. S. McConnell, from Phenacoccus sp. found
feeding on a species of Andropogon at Stevensville on Kent Island in
Chesapeake Bay, August 3, 1942.

IS. PSEUDAPHYCUS HERACUS, new species

This species is very similar to meritoritts, new species, described
elsewhere in this paper, and the two species are found infesting appar-
ently the same species of mealybug in some localities in Virginia.
It differs from jrwritorius, however, by having the segments of the
funicle shorter and more transverse, by having the club somewhat
longer than the entire funicle, and by having the anterior ocellus
situated distinctly above the middle of frontovertex.

Female. — Length 0.8G mm. Frontovertex, mesoscutum, axillae,
and scutellum orange with no dark markings; head except frontover-
tex, tegulae. ])r()notum, pleura, sterni, and all legs whitish; metanotura
and propodeum orange, a shade lighter than scutellum ; abdomen dor-
sally mostly yellowish with a narrow blackish cross stripe connecting
the cerci and basad of this stripe with one and apicad of it with two
similar but less distinct fuscous stripes; abdomen beneath slightly
paler; ovipositor pale at base, darker apically; eyes grayish black,
(xpjli dark red ; antennal scape nearly uniformly pale yellowish, pedi-
cel and funicle fuscous, the apical funicular segments. usually paler
than the basal ones; club white. Wings uniformly subhyalinc; mar-
ginal and stigmal veins fuscous. Hairs on mesonotum whitish.

Frontovertex somewhat more than twice as long sis broad, finely
granulose but under high magnification with faint indications of

326 PROCEEDINGS OF THE NATIONAL MUSEUM vol. »«

alveolate sculpturing ; inner orbits parallel for most of their length ;
ocelli forming a slightly acute triangle ; anterior ocellus about one and
a half times the width of f rontovertex above the edge of scrobal cavity ;
lateral ocellus less than its own diameter from eye margin and more
than its diameter from occipital margin ; eyes with, distinct short pile.
Antenna clavate; scape not expanded beneath, slightly broader at
middle than at either end, four times as long as broad ; pedicel about
one-third as long as scape, very nearly as long as first four funicular
segments combined; funicular segments all broader than long, suc-
cessively increasing in thickness from first to last, the basal segments
distinctly narrower than pedicel, last segment about as broad as
pedicel; club large, broader than last funicular segment and a little
longer than entire funicle.

Mesoscutum, axillae, and scutellum uniformly granulosely punctate,
mat; propodeum smooth; mesopleuron distinctly finely reticulate-
punctate, subopaque. Forewing with rather dense discal ciliation,
the cilia basad of speculum a little coarser than those beyond ; speculum
closed posteriorly by one or two rows of hairs, otherwise uninter-
rupted; marginal vein punctiform; stigmal vein a little longer than
postmarginal.

Abdomen short ovate, slightly narrower and a little shorter than
thorax, very weakly reticulated and shining dorsally, and practically
smooth ventrally. Ovipositor exserted about one-fourth the length
of abdomen.

Male. — Length 0.75 mm. Similar in practically every respect to the
female except that the abdomen is nearly circular in outline, obviously
shorter and narrower than the thorax and blackish medially with the
margins broadly yellowish.

T'i/pe locality. — Hamilton, Va.

Type.—V.S.'^.M. No. 57330.

Remarks. — Holotype female, allotype, and 13 paratypes reared July
5, 1943, from Pseudococcus virgatus (Cockerell) , collected at Hamilton,
Va., and received from D. W. Clancy of the Charlottesville, Va., lab-
oratory under No. 628 ; also 35 paratypes reared May 16, 1944, from
the same host collected at Batesville, Va., under Clancy's No. 611.
Many additional specimens reared by Clancy from P. virgatus col-
lected at Hamilton, Batesville, Crozet, Winchester, Greenwood, and
Covesville, Va., are in the collection identified as this species but not
considered a part of the type series.

16, PSEUDAPHYCUS ALVEOLATIFRONS. new species

Differs from all other forms studied in the coarser and much more
distinct alveolate sculpture and paler orange-yellow color of the
frontovertex.

CHALCID-FLIES OF GENUS PSEUDAPHYCl'S (iAHAN 327

Female. — I-/en«rth O.S'2 mm. Frontovertox very dilute orange yellow,
its entire surface uniformly densely covered with fine but perfectly
distinct facetlike punctures. In all other respects apparently agree-
ing with the foregoing description of wrrar?As so that the description
of that species except for the frontovertcx will serve for this species
also.

Mall. — The head is gone from the single male specimen received
so that the characters of the head are unknown. With respect to the
other characters this male seems to differ in no significant respect from
the male of meracus.

Type locality. — North Bergen, N. J.

Type.—V.S.^.^l. No. 57331.

/ifmarl.-s. — Described from a unique female and one headless male
reared in November 1938 by George Rau from Pseudococcus com-
^tocki (Kuwana).

The frontovertcx in most of the forms studied is granulosely sculp-
tured and mat, with little or no indication of distinct separate alveoli
or punctures even under the higher magnifications of a binocular
microscope. In anf/eh'cus, ahstru^ii-s, meraou.s., and limafulits the sculp-
ture of frontovertex is more or less distinctly alveolate but the alveoli
are finer and less definite. P. alveolatifrons differs from angeliciis and
ahstnuiiwi further by having the ocelli in an equilateral, instead of a
distinctly acute, triangle and l)y lacking the transverse hyaline streak
on the wing. From Umatulus it may be distinguished by the distinctly
duller sculpturing of the mesonotum as well as by its brighter orange-
yellow color.

It is possible that additional material may show that the differences
between meracus and aJ^vcolatifrons are not of specific value, but in
view of the fact that the types of the two species are from different
hosts and from different localities and this sculptural difference does
Lxist, it seems wisest to give a separate name to this form from
/-'. comntocki even though it may eventually turn out to be a synonym.

I tOVlaNMIIIT PRINTJRt OFriCI II4«

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

iutui ISiJlvA, 0*0 hy the

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM
Vol. % Wa.hington: 1946 No. 3201

NEW CERAMBYCID BEETLES BELONGING TO THE TRIBE
DISTENIIXI FROM CENTRAL AND SOUTH AMERICA

By W. S. Fisher

DuRixG the process of rearranging the American species of the tribe
Disteniini (Coleoptera: Cerambycidae) in the United States National
Museum, four new species were found. These are described herein.

Genus DISTENIA Lepelitier and ServiUe
DISTENIA LATERALIS, new species

Head, pronotum, and underside of body reddish brown; elytra and
scutollum yellowish brown, the elytra with a distinct, broad, longitudi-
nal, green vitta on each side near lateral margin ; antennae brownish
black; palpi and legs palo brownish yellow.

Head glabrous, sparsely, irregularly punctate. Antenna nearly
one and one-half times as long as body, sparsely clothed with long
flying hairs on underside: first segment slightly shorter than third,
slender, cylindrical, gradually expanded toward apex, not depressed
on top at base, rather densely, finely punctate, and sparsely clothed
with long and short, semierect hairs.

Pronotum, not including lateral spines, slightly longer than wide,
widest at middle; sides strongly constricted near base iind ajiox, tri-
angidnrly expanded on each side at middlo jind ai-med with a long,
f^onical, acute s[)ine; disk broadly, transvei-sely depressed near autcfior
margin, transversely, sinuately grooved near base, with a sliglitly
ol'vatod space oji each side in front of transvoi-se, basal groove; sur-
face glabrous, coarsely, sparsely, irregularly puiu-tate. with a smooth,
elongated, median space.

Elytra at base slightly wider than pronotum including lateral
spines; sides gradually converging from bases to apices, wliieh are
T13015— «« 320

330 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9fl

unispinose, the lateral spine on each rather short and acute, and the
sutural angle feebly produced; surface very densely, coarsely, deeply
punctate, becoming nearly impunctate near apices, very sparsely
clothed with rather short, inconspicuous, erect, yellowish hairs.

Body beneath indistinctly punctate, very sparsely, irregularly
clothed with short, inconspicuous, recumbent, yellowish hairs, with a
few longer, erect hairs intermixed; femora slender, cylindrical,
slightly clavate, each armed with a long, acute spine at apex.

Length 22 mm., width at base of elytra 4.5 mm.

Type locality. — Rurrenabaque (Beni River), Bolivia.

T^^z/^e.— U.S.N.M. No. 57612.

Remarks. — Described from a single specimen collected at the type
locality during October by William M. Mann in connection with the
Mulford Biological Exploration during 1921-1922.

This species is closely allied to Distenia limhafa Bates, but it differs
from that species in having the head and pronotum reddish brown
with only a vague bronzy reflection, the lateral spines on each side of
the pronotum long and acute, the scutellum yellowish brown, the elytra
more coarsely and densely punctured and with the apices armed with
a long, acute, lateral spine, and the antennae brownish black.

DISTENIA SPINIPENNIS. new species

Head, pronotum, scutellum, elytra, and underside of body greenish
black (elytra slightly more greenish) ; antenna (except first segment
which is black) and palpi reddish brown; legs pale brownish yellow,
the femora slightly darker at apices.

Head nearly glabrous, sparsely, irregularl}', indistinctly punctate.
Antenna one and one-half times as long as body; first segment dis-
tinctly shorter than third, robust, very strongly clavate, strongly,
longitudinally depressed on top on basal half, scabrous and coarsely
punctate basally, and sparsely clothed with long, fine, semierect hairs.

Pronotum, not including lateral spines, slightly longer than wide,
widest at middle ; sides strongly constricted near base and apex, tri-
angularly expanded on each side at middle and armed with a rather
long, acute spine; disk broadly, transversely depressed near anterior
margin, narrowly, transversely, sinuately grooved near base, narrowly,
transversely grooved along base, and with three slightly elevated,
elongate, smooth spaces, one median and one on each side of middle;
surface sparsely, indistinctly punctate in depressions, and clothed with
a few short, indistinct, erect hairs.

Elytra at base as wide as pronotum including lateral spines ; sides
strongly converging from bases to apices, which are unispinose, the
lateral spine on each long and acute, and the sutural angle rectangu-
lar; surface coarsely, densely, deeply punctate, becoming nearly im-

NEW CERAMBYCID BEETLES — FISHER 331

punctate at apices, rather densely, uniformly clothed with short, erect,
whitish hairs.

Body beneath indistinctly, irre«;ularly punctate, sparsely clothed
toward sides with long and short, seniierect, whitish hairs; femora
slender, slightly clavate, unarmed at apices.

Length 19.5 mm., width at base of elytra 4 mm.

Type lotality. — "Guapiles," 250-300 meters, Costa Rica.

Ty/^d.— U.S.N.M. No. 57613.

Remarks. — Described from a single specimen collected at light dur-
ing May 1934, by Ferdinand Nevermann. The specimen is labeled
Guapiles, which is probably an error for Guaplies.

This species is closely allied to Distenia phaeocera Bates, but it
dili'ei-s from the description given for that species in not having the
elytra stria-punctate or the surface alutaceous, and in the slightly
elevated smooth spaces on each side of the pronotum being not divided.

Genus COMETES Lepelitier and Serville

COMETES EMARGINATA. new species

Elongate and strongly shining, black, except anterior coxae, anterior
femora (except tips) , basal halves of middle and posterior femora, and
tibiae in part brownish yellow.

Head finely, irregularly punctate on top, somewhat transversely
rugose anteriorly, with a smooth, longitudinal groove extending from
clypt'us to occiput, sparsely clothed with long, erect and recumbent,
whiti.-h hairs. Antenna one and one-half times as long as body,
sparsely clothed with l<->ng, flying hairs on underside; first segment
slightly longer than third, robust, cylindrical, gradually expanded
toward apex, finely, rugosely punctate, sparsely clothed with long erect
and short recumbent, whitish hairs.

Pronotum as wide as long, widest at middle; sides strongly con-
stricted near base and apex, triangularly expanded on each side at
middle but not distinctly spinose; disk broadly, transversely depressed
near anterior margin, transversely, sinuately grooved near base, nar-
rowly, transversely grooved along ba.se, and with two slightly elevated,
rouiul, smocjth gibbosities on each side <jf middle; surface densely,
finely punctate; in median depression, the punctures nearly concealed
by a dense, recumbent, yellowish pubescence, and with a few long, erect,
white hairs.

Elytra at base as wide as [)ron<jlum at middle; sides gradually con-
verging from bases to apices, which are separately enuirginale. the
lateral spine slightly longer than su(ural si)ine; surface coarsely,
deeply, densely punctate, tiie puncture forming more or le.ss distinct
rows but becoming obsolete toward apices, rather den.sely clothed
toward sides with short, indistinct, it'iinibent, whitish hairs, with a

332 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

few long, erect liairs intermixed, and each elytron ornamented with a
narrow vitta of transversely recumbent, white hairs along sutural
margin.

Body beneath indistinctly punctate, sparsely clothed at sides with
short, recumbent, whitish hairs ; legs sparsely clothed with long and
short, semierect, white hairs.

Length 11 mm., width 2 mm.

Ti/pe locality. — Hamburg Farm (on Keventazon River) , Costa Rica.

Type.—V.S.'NM. No. 57614.

Remarks. — Described from a single specimen collected at the type
locality on grass, May 26, 1934, by Ferdinand Nevermann.

This species can be separated from the other described species of
this genus by having the tips of the elytra emarginate.

COMETES BICOLOR, new species

Elongate, rather strongly shining except elytra which are sub-
opaque; antennae bluish black with a vague violaceous reflection;
head, pronotum, scutellum, underside of body, and legs bluish black
with a distinct greenish or violaceous tinge; elytra brownish yellow
with apical fourth and elevated lateral margins violaceous blue.

Head rather coarsely, irregularly punctate on top, somewhat trans-
versely rugose anteriorly, with a smooth, longitudinal groove extend-
ing from clypeus to occiput, very sparsely clothed with inconspicuous,
semierect hairs. Antenna nearly one and one-half times as long as
body, sparsely clothed on underside with long, flying hairs ; first seg-
ment as long as third, slender, cylindrical, narrow at base, strongly
expanded toward apex, rather densely, coarsely, shallowly punctate,
sparsely, uniformly clothed with short, semierect, whitish hairs.

Pronotum slightly wider than long, widest at middle ; sides strongly
constricted near base and apex, obtusely expanded on each side at mid-
dle but not spinose ; disk broadly, transversely depressed near anterior
margin, shallowly, transversely, sinuately grooved near base, nar-
rowly, transversely grooved along base, with two irregular, smooth
elevations on each side of middle, and an elongate, smooth, median
elevation; surface coarsely, deeply, confluently punctate, sparsely
clothed with moderately long, erect, inconspicuous hairs.

Elytra at base as wide as pronotum at middle; sides parallel from
bases to near apices, which are separately narrowly rounded ; surface
densely, coarsely, deeply, uniformly punctate, the punctures becom-
ing more confluent toward apices, rather densely clothed with short,
inconspicuous, erect hairs, and each elytron with two more or less dis-
tinct longitudinal costae, the inner costa extending from base to mid-
dle of elytron, and outer one from base to apical fourth.

Body beneath sparsely, shallowly, irregularly punctate, sparsely,
irregularly clothed with moderately long, semierect, fine hairs; legs
rather densely clothed with long and short, semierect, whitish hairs.

NEW CERAMBYCID BEETLES — FISHER 333

Length 11.5 mm., width 3 mm.

Type locality. — Colombia, South America (no definite locality).

ry^e.— U.S.X.M. No. 57G15.

litmarlca. — Described from a single specimen collected August 31,
1942, by Francisco J. Otoya (No. 2053).

This species is allied to Comctes pulcherrivins Bates, but it differs
from that species in iuiving the pronotum more densely and coarsely
punctured, and the basal three-fourths of the elytra brownish yellow,
with only the apical fourth and narrow lateral margin of each elytron
violaceous blue. The head is mounted separately on the same pin
with the rest of the specimen.

u I aaviNMHiNr PMiNTiMa orricit lt4t

PROCEEDINGS OF THE UNITED STATES NATIONAl MUSEUM

issued jvAjfV vi» Cjyj ^ fy the

^^^^

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM

V' I. ')6 Washingtor: 1946 No. 3202

MACHAEROIDES EOTHEN MATTHEW, THE SABER-
TOOTH CREODONT OF THE BRIDGER EOCENE

By C. Lewis Gazin

The 1940 Smithsonian Institution expedition to the Bridger
Basin of Wyoming' had unusual good fortune in securing skeletal
remains of the rare creodont Machaeroides eothen. The materials,
consisting of skull, lower jaws, and other skeletal portions of one
individual, were found by Franklin Pearce in low exposures,
probably of Bridger "C," immediately to the north of Twin
Buttes, about 30 miles southwest of Green River, Wyo. Previous
finds of this remarkable sabertooth form, so far as known, are
limited to the lower jaw portions described by Matthew ^ in 1909.

The skull, U.S.N.M. No. 17059, v/as found with the lower jaws
in a position of articulation, and although essentially complete
there has been a small amount of transverse crushing and dis-
tortion so that the riglit side of the skull is higher than the left.
The remainder of the skeleton includes portions of the vertebral

lumn, both humeri and femora, the right radius and ulna, an
incomplete left tibia, and fragments of the pectoral and jielvic
girdlrs. The feet were not present oxcopt for a carpal and two
metacarpal bones.

Matthew readily appreciated the indications in the lower jaw
portions of Machaeroides eothen of a modification nearly parallel-
ing that of the machairodont forms among the Fissipedia. In
the materials he had at hand these modifications were not too
evident, and R. H. Denison - was inclined to consider the M.
eothen jaw as resembling Felin and not truly "sabertooth." The

' Matthew, W. D.. Mem. Amcr. Mus. N*t Hist., vol. 9, pt. 6, pp. 4B2-<63, 1909.
' n..nl<»on. R. II.. Ann. N«w York Acad. 8cl.. vol. 87. p. 181, 1938.

335

336 PROCEEDINGS OF THE NATIONAL MUSEUM VOL.96

National Museum specimen, however, leaves no doubt of the
direction and extent of specialization, which quite parallels that
seen in the machairodonts but not to the extent exhibited in the
Uinta Apataelurus kayi described by Scott. ^ Structurally no
characters are observed in the illustrations of the lower jaws of
A. kayi that would preclude derivation of this form from M,
eothen, with the characteristics attendant upon sabertooth de-
velopment reaching a high degree of specialization within an
interval of time suggesting a comparatively rapid evolutionary
tempo.

Matthew regarded Machaeroides eothen as an oxyaenid type
of creodont, closely related to forms included in the subfamily
Limnocyoninae, to which he allocated it (p. 410) , * having created,
however, the subfamily name "Machairoidinae" on an earlier
page (330). Denison ^ retained the supergeneric separation,
which was entirely justified by the discovery of Apataelurus,
recognizing a natural phyletic subfamily. However, Denison
removed the Limnocyoninae and Machaeroidinae from the Oxyae-
nidae and placed them in the Hyaenodontidae. Justification for
this was claimed on the basis of morphological differences between
the Oxyaeninae and Limnocyoninae and similarities between the
Limnocyoninae and Proviverrinae, particularly between Prolim-
nocyon and Sinopa. Separation from the Oxyaenidae seems sup-
ported, and, moreover, carnassial specialization, which has been
the key to their supposed affinity, may well have developed inde-
pendently in the two groups. I cannot, however, but regard the
carnassial differentiation that so readily distinguishes the Lim-
nocyoninae from members of the Hyaenodontidae as being of
fundamental importance, and hold that the morphological simi-
larities may be as easily attributed to similar adaptation or
parallelism. It is in a similar manner that the Machaeroidinae
so markedly resem.ble the machairodonts, although obviously not
derived one from the other. Derivation of the Limnocyoninae
from an early proviverrine stock is not disproved and may be
reasonable as suggested by the similarity of lower jaws belonging
to forms of Prolimnocyon and Sinopa morclax, but their di-
vergence must originate in a primitive form with molars as yet
undifferentiated as to carnassials, and where M-, for example, has
not taken on characteristics so markedly different in the two
groups.

3 Scott. W. B., Ann. Carnegie Mus., vol. 27, art. 6, pp. 113-120, 1938.

* Matthew, W. D., ibid.

» Denison, R. H., ibid., p. 181.

MACHAEROIDES EOTHEN MATTHEW GAZIN 337

Inasmuch as it seems advisable to exclude the Limnocyoninae
from the Oxj-'aenidae, and since I am unable to reconcile it with
the Hyaenodontidae on the basis of carnassial differentiation, I
propose, at the risk of censure, that the Limnocyoninae and
Machaeroidinae be given family recognition as the Limnocyonidae,

Description of skull (pi. 45) . — The skull of Machaeroidcs eothen
is significantly smaller than that of Limnocyori verns but much
larger than Thinocyon vclox, corresponding closely in size to
Siyiopa rapax among its less closely related contemporaries. The
skull is moderately slender but with a noticeably deep rostrum,
high srgittal crest, and a narrow occiput.

The rostrum, in addition to depth, exhibits a well-inflated
maxillary portion covering the long root section of the canine,
extending parallel and immediately posterior to the suture join-
ing the premaxilla. The nasals extend posteriorly in a nearly
F-shaped wedge between the frontals, terminating fully as far
back as the postorbital processes. In Thinocyon and Limnocyon
the nasals as exposed terminate distinctly forward of this position.
The postorbital processes of the frontals are moderately well
developed and, although possibly not complete, do not appear so
prolonged as in Thinocyon. The lachrymal bone in M. eothen is
lar;;e and extends well forward of the orbit and exhibits a most
unusual feature in that the lachrymal foramen enters the skull
anterior to the orbital rim and anteroventral to the lachrymal
crest or tubercle. A smaller foramen also enters the lachrymal
bone outside and immediately dorsal to the lachrymal tubercle.
This arrangement was not ob.-.erved in other creodonts, but it
occurs in marsupials and sloths. Matthew, " however, noted that
the lachrymal foramen in Limnocyon was very near the orbital
rim.

The cranial portion of the skull is characterized by an exceed-
ingly small brain case, relatively much smaller than in Thinocyon,
and a very high sagittal crest, which joins a high but narrow
occipital crest. The latter is overhanging but, quite unlike Lim-
nocyon, is noticeably constricted transversely immediately above
the occipital condyles. A prominent vascular foramen is noted
at the suture between the parietal and squamosal at a position
about over the trough for the audital tube, and one also in the
posterior portion of the temporal fossa, presumably at about the
juncture of the parieto-squamosal and parieto-occipital sutures.

In ventral aspect the palatal portion of the skull shows no
unusual features. No evidence exists of the grooves and ridges

• Matthew. W. D , ibid., p. 438.

338 PROCEEDINGS OF THE NATIONAL MUSEUM VOL.96

characterizing' the later fissiped sabertooth cats. However, palatal
excavations between the deuterocone or protocone portions of
successive 3-rooted teeth, for reception of the much-elevated pro-
toconids of the lov/er cheek teeth, are as well developed as in
several of the other creodonts. The posterior margin of the palate
is extended posteriorly a very short distance below the narial
passage, somewhat as in Limnocyon. The palatal margin outside
. the narial passage, however, shov/s a conspicuous notch for the
palatine vein, with well-developed tuberosities on both the palatal
and maxillary sides of the groove.

^ The zygomatic arch arises from perhaps a slightly higher posi-
tion on the relatively deep rostrum of Machaeroides eotken than
it does in Limnocyon verus, although in M. eothen the left arch-
is broken down to a position below normal on that side of the skull.
The depth of the arch is moderate, but because of crushing the
extent to vv^hich it is expanded laterally cannot be determined.
Posteriorly the arch terminates at a position relatively low with
respect to the basicranial surface. The zygomatic process of the
squamosal projects conspicuously dovv^nward from the basicran-
ium, placing the glenoid surface for articulation of the lower jaw
at a much lower level than observed in other creodonts, a condition
noted in machalrodonts and in the sabertooth marsupial Thyla-
cosmilus atrfT. This lowering of the fulcrum gives leverage to
the temporal muscle acting on a coronoid process of reduced
height. The reduction of the coronoid presumably permitted the
lov/er jaw to open v»'ide enough for the mandible to clear the
saberteeth, apparently much wider than necessary in other carni-
vores, except Apataelurus and the machalrodonts.

The basicranial portion of the skull is the least distorted by
crushing and is relatively elongate and clearly much narrower
than in Limnocyon verus. The paroccipital process shows very
little development and does not project posteriorly as in Limno-
cyoii verus. However, in addition to the downward-projecting
pedestallike base for the glenoid surface, the mastoid process, as
in the machairodonts, is very well developed, projecting downward
and forvv^ard, and noticeably expanded in an anterointernal-
posteroexternal direction. The mastoid process is moderately
developed in most creodonts including Limnocyon, but nowhere in
the suborder is it relatively so important, particularly in a for-
ward medial extension, as in M. eithen, and in Apataelurus by
inference. Its prominence and rugosity demonstrate the strength
and importance of the sterno-cleido-mastoid muscle, the actions
of which include depressing the head, as in striking with the

MACHAEROIPES EOTHKN MATTHEW — GAZIN -^39

sabers. The mastoid process is essentially a part of the mastoid
portion of the periotic ; however, the extent to whicli the squamosal
enters into its composition cannot be determined, limited possibly
to a portion of the anterior surface of the process.

The foramina of the hasicranium show certain significant dif-
ferences in relative position from those in Limiiocyon vems or in
Thinocyon relox. For the most part these maintain a primitive,
creodont arrangement with differences noted in Machaeroides
eothcn that are in part due to structural modification attendant
upon sabertooth development. The alisphenoid canal, if present,
is decidedly long as in Limnocyon, but with the posterior opening
confluent with or not distinguished from the foramen ovale, so
that it was not certainly identified in the material at hand. In
Limnocyon veirus the posterior opening of the alisphenoid canal
's shown by Matthew ^ as well forward of the foramen ovale.
The foramen ovale in M. eothcn is located medial to the post-
glenoid process, at the root of the pterygoid wing of the alisphe-
noid. The postglenoid foramen enters the skull at the base of
the posterior surface of the postglenoid process but exhibits a
groove for about half the length of this surface before closure is
complete. The foramen lacerum medius is in a customary posi-
tion antero-internal to the exposed portion of the petrosal. From
a position medial to the promontorium of the petrosal a narrow
cleft extends posteriorly along the outwardly convex lateral
margin of the basioccipital to the foramen lacerum posterius,
about halfway to the occipital condyles. The internal carotid
evidently entered the brain case at some point along the anterior
part of the cleft and adjacent to the petrosal. The condylar or
hypoglossal foramen is well forward of the condyles and sepa-
rated by a thin partition from the foramen lacerum posterius,
quite unlike Limnocyon or Thinocyon but resembling certain
specimens of the Oligocene Daphoeims in this respect. The
stylomastoid foramen shows as a groove on the medial margin
of the mastoid process. Dorsally this is completely enclosed,
presumably by bone belonging to the mastoid portion of the
periotic, at the root of the mastoid process, posterolateral and
very close to the promontorium of the petrous portion.

Machaeroides eothcn is without a tympanic bulla, and the
tympanic ring was not preserved. The site of the audita]
tube is a deep and compressed J7-shaped trough between the
postglenoid and mastoid jjrocesses, extending laterally and some-
what posteriorly from below the anteroexternal portion of the

' M«tth-w. W. D.. ibid.. t\z. 66.

340 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. %

petrosal. In Limnocyon and Thinocyon the trough is more widely
open and relatively shorter. The petrosal is partially exposed
on both sides of the skull of M. eothen. It exhibits an acutely
projecting promontorium located immediately inward from the
medial margin of the mastoid process. The petrosal is more
broadly rounded ventrally in Thinocyon. The fenestra rotunda
in M. eothen is large and faces posteriorly and slightly outward
and downward below the flattened ventral surface of the pos-
terior portion of the petrosal. The anterior surface of the
petrosal is broad and flattened, facing anterolaterally and ven-
trally and joining the medial surface in a bluntly rounded angle
that extends anteromedially and dorsally from the promontorium.
The fenestra ovale faces slightly forward of lateral and is situ-
ated very deep in the mesotympanic fossa, almost directly above
but well separated from the promontorium. It is much higher
in position than the fenestra rotunda and is very much above the
lip of the trough for the audital tube. Further description of
the petrosal and its relation to the cranial cavity is not under-
taken, inasmuch as such additional information cannot be ob-
tained without damage to the skull.

Upper dentition. — The dental formula for the upper teeth of
Maclmeroides eothen is 3-1-4-2, as noted by Matthew in Lim-
nocyon and as observed in Thinocyon. Moreover, the first upper
molar is the carnassial as in these forms and as in the Oxyae-
nidae. The teeth resemble those in the Limnocyoninae with
certain exceptional characteristics which for the most part are
modifications seemingly accompanying sabertooth development.

The incisors are slender, recurved, conical teeth adapted for
piercing and with transversely fiattened roots. Unlike Lim-
nocyon they increase in size from first to third, P being much
more robust and with a root portion about twice as long as in I'.
P in Limnocyon is much smaller than P and may not be present
in some individuals, as indicated by Matthew. * In Thinocyon
the three are subequal and slightly spatulate.

The enlarged canine is preserved only on the left side and is
broken away a short distance below the alveolus. It is removed
from P by a short diastema. The tooth as exposed has no cingu-
lum, is nearly oval in cross section, and has a long gently curved
root section, as inferred from the inflated portion of the maxilla.
In Matthew's ® illustration of Limnocyon verus the upper canine
appears to have an exposed cingulum and the short crown is

• Matthew, W. D., ibid., p. 434.
•Matthew, W. D., ihid., fig. 58.

MACHAEROIDES EOTHEN MATTHEW — GAZIN 341

noticeably recurved below this point. The length of the canine
in M. eothen cannot be determined from the present material;
however, if this tooth extended as far as the flange of the lower
jaw it would have had a length of about 3 cm. beyond the alveolus.
The canine shows no evidence of serrations along the anterior or
posterior margins.

The premolars of M. eothen are 1-, 2-, 3-, and 3-rooted, respec-
tively. F\ preserved only on the right side when found but
subsequently lost, is a small, simple, conical tooth without an
accessory cuspule and separated by a diastema from both the
canine and P-. In both Limnocyon and Thinocyon P^ is 2-rooted
and exhibits a small posterior cuspule. P- of M. eothen, in addi-
tion to being 2-rooted, retains a vestige of a posterior cuspule.
The tooth is relatively much smaller and transversely more
compressed than in Limnocyon. P% on the other hand, is much
better developed than in Thinocyon and apparently than in
Limnocyon. This tooth, preserved only on the left side, has
a minute parastyle and a rather well developed posterior cusp
or crest, approximating P* in this respect. The lingual root
is slender but extends markedly inward from about midway
of the tooth length and supports a very small deuterocone.
P* is relatively robust with a prominent parastyle before and a
trenchant cusp posterior to the large backward sloping primary
cusp. The deuterocone portion is broad and well defined, extend-
ing lingually from the midportion of the tooth. The deuterocone
portion does not project forward as it does in Thinocyon or (to a
less degree) in Limnocyon. The deuterocone is situated on the
lingual margin of the talon and is connected with the primary
cusp by a low crest across the forward part of the talon. A small,
shallow basin is enclosed between this forward crest and the
low cingular crest around the posterior margin of the talon.

The difference in development of the various premolars is
rather striking in comparison with related forms. P' and P* are
relatively small and less progressive, probably reduced from an
earlier but more advanced state, paralleling the machairodonts in
this respect, as an accompanying factor in sabertooth specializa-
tion. P\ however, has retained a more advanced stage of devel-
opment and undoubtedly continued to be a significant and func-
tional tooth in the later and more advanced Apntaeluntfi, as
indicated by the well-developed and trenchant P^ with which it
occludes in the lower jaw of the Uinta form.

^r, the upper sectional tooth in Machneroides eothen, has taken
on a very trenchant appearance as compared either with the

342 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. w

preceding P* of MacJiaeroides or with M^ in Limnocyon and
Thinocyon. The talon portion is much reduced and far forward
in position, the deuterocone being but a very small cusp at the
anterolingual angle of the tooth. The posterior crest is elongate
and the greater part of the lingual face of the tooth forms a flat
shearing blade which, though oblique, is directed more nearly
longitudinal than in either Limnocyon or Thinocyon. The para-
cone and metacone of M^ are closely connate, more so than in
Limnocyon, approximately as in Tritemnodon. The parastyle is
but weakly developed, being represented only by an enlargement
of the cinguium anteroexternal to the paracone.

M^ has a nearly transverse shear and is characterized in dis-
tinction from that in Limnocyon and Thinocyon in having lost all
trace of the metacone. Also, the talon is reduced to a very sub-
dued projection from the paracone, the shear being effected essen-
tially by the paracone and parastyle, whereas in Limnocyon and
Thinocyon the talon participates very largely in the shearing
structure, cutting against the metaconid and occluding with the
talonid of Mo. The reduction of the talon of M^ in M. eothen
accompanies the loss of the metaconid and extreme reduction of
the talonid of the lower carnassial.

Mandible (pi. 46, a, b) . — The lower jav/ of Machaeroides eothen
has been briefly described and figured by both Matthew and Deni-
son, and the relationship to other creodonts discussed at length.
However, the specimens available to them for study were incom-
plete, lacking particularly the extremity of the flange, coronoid
process, and the crowns of the canine and carnassial teeth. Res-
torations of these parts in drawings were conservative in indicat-
ing less modification from the limnocyonine pattern than the more
complete material demonstrates.

The lower jaw is relatively very deep and transversely slender
compared to jaws belonging to forms of the Limnocyoninae. Its
flange projects downward to a greater extent than anticipated but
not to the extent shown in Apataelurus kayi, and much less than
in Hoplophoneus. In M. eothen it projects but a little below the
dorsoventrally elongate symphysis and curves gently outward so
that the width across the extremities of the flanges is greater than
at any point above. The deep symphyseal surface has a nearly
straight, steeply inclined anterior margin, which makes an abrupt
angle with lower margin. The coronoid process of the lower jaw
appears truncated and much reduced from the large and fully
developed coronoid observed in Limnocyon and Tkhiocyon. The
reduction, nevertheless, has not gone so far as that in Apataelurus.

MACHAEROIDES EOTHEN MATTHEW — OAZIN 343

Moreover, the condyle is lower with respect to the tooth row.
The angle, less widely separated from the condyle, does not curve
downward from the nearly straight lower margin of the jaw
but preserves the alignment practically to its extremity, more so
than in Apataeh/jus. The masseteric fossa is well defined with
a sharp masseteric crest anterodorsally ; however, the fossa, as
in Apataelurus and also Pafriofelis, extends farther forward be-
neath the carnassial than in Limnocyon and Thinocyon. The
mental foramina are beneath Pj and the posterior root of P-j and
placed relatively low on the side of the jaw.

Lower dentition. — The lower teeth are all present, although
the right canine and Po as well as the left carnassial and median
incisors are slightly damaged. The lower incisors of Machae-
roides eothen were most certainly reduced to two. The lateral
of these is the larger of the two, and both, as in the case of the
upper incisors, are piercing type structures with transversely
flattened roots. In lateral view the canine appears moderately
robust at the alveolar border but is transversely flattened and
tapers rapidly to a point only a little above the closely adjacent
incisors. Moreover, it shows a pronounced scar or bevel verti-
cally along its posterolateral surface, worn through occlusion with
the superior canines.

The cheek teeth appear decidedly slender, and the two anterior
premolars, both of which, however, retain two roots, are of rather
small size. P3 is about intermediate in size and development be-
tween P2 and P^. It has a crested talonid, almost as in P4; a
structure but very feebly expressed in Po. P3 is without a para-
conid. whereas in P, this cusp is low but distinct. Both P3 and P4
are more progressive than in Limnocyon and Thinocyon; however,
in Apataelurus P.-^ has become relatively much reduced in size.
P^ in both Machaeroides and Apataelurus has become a relatively
large and functionally significant tooth, actually exceeding Mi
in size in both forms.

The two molars possess somewhat more distinctive structural
characters than the premolars in characterizing Machaeroides.
In M, the metaconid is but moderately developed and pressed
close to the protoconid. The talonid is much reduced in size and
trenchant in character. In Limnocyon and Thinocyon the rela-
tively wider and more robust Mi has a better-developed metaconid
and a large, deeply basined talonid. In Apafaelvrvs the trenchant
heel persists and evidence remains of a metaconid that has not
been entirely oliliterated by the shearing function imposed on the
tooth.

344 PROCEEDINGS OF THE NATIONAL MUSEUM VOL.98

The camassial, M2, in M. eothen exhibits more noticeable modi-
fications than Ml toward a sabertooth specialization. The meta-
conid is lost, except for a slight rugosity on the posterointernal
margin of the backward sweeping protocone. The talonid is re-
duced to little more than a vestige, although its expression is
somewhat better in the specimen figured by Matthew. ^° The
tooth is almost catlike in appearance, quite unlike that in Limno-
cyon or Thinocyon where the carnassial is very much like Mj
except for its greater size and higher trigonid. In Apataeluncs
the lower carnassial has almost or quite reached the stage ex-
hibited by Smilodon in its modification for shearing, but still
preserving a vestige of its trenchant talonid.

Table 1. — Measurements {in millimeters) of skull, mandible, and dentition
of Machaeroides eothen, U. S. N. M. No. 17059

Greatest length of skull from anterior margin of premaxillae to posterior margin

of occipital condyles a 135

Distance from anterior margin of premaxillae to posterior narial aperture a 65

Greatest depth of maxillae above P' a 32

Width of palate between canine alveoli a 16

Greatest width of basicranial region across mastoid processes a 48

Length of upper dentition from I' to M= inclusive a 62

Length of upper dentition from anterior margin of canine alveolus to M' inclusive. ... a 66

Width across I' to P inclusive 76

Length of cheek tooth series, P' to M- inclusive 42

Length of diastema anterior to P' : posterior to P^ 3.6 ; 2.6

Length of premolar series, P^ to P* inclusive 30

Length of molar series. M^ to M^ ll.S

Canine — anteroposterior diameter at alveolus: transverse diameter 8.6: 6

P' — anteroposterior diameter: greatest transverse diameter 8.2: 2.1

P'^ — anteroposterior diameter: greatest transverse diameter 5.4: 2

P* — anteroposterior diameter: greatest transverse diameter 8 : 5

P* — anteroposterior diameter: greatest transverse diameter 10 : 8.2

M' — anteroposterior diameter: greatest transverse diameter 9.4: 7.5

M^ — anteroposterior diameter: greatest transverse diameter 3.5: 8.1

Length of lower jaw from anterior extremity to condyle 92

Depth of symphysis of lower jaw measured along anterior face 2<

Depth of flange of lower jaw below diastema between canine and Pi 22

Depth of lower jaw below Pj 16.6

Depth of lower jaw below Mj 16.6

Height of coronoid process above inferior margin of ramus 28

Length of lower dentition from anterior margin of canine to Mj inclusive 67.6

Length of cheek tooth series. Pi to M2 inclusive 43.5

Length of diastema anterior to Pi : posterior to Pi 8 : 2

Length of premolar series. Pi to P< inclusive 28.8

Length of molar series. Mi and M2 16.9

Canine — anteroposterior diameter at cingulum: transverse diameter 6.7: 3.6

Pi — anteroposterior diameter: greatest transverse diameter 8.6: 1.6

P2 — anteroposterior diameter: greatest transverse diameter 4.7: 2

P3 — anteroposterior diameter : greatest transverse diameter 7.6 : 3

P4 — anteroposterior diameter : greatest transver.se diameter 9.1 : 4

M, — anteroposterior diameter : greatest transverse diameter 8.6 : 4

Ma — anteroposterior diameter : greatest transverse diameter 8.7 : 4.4

Vertebrae. — Among cervical vertebrae preserved of Machae-

i» Matthew, W. D., ibid., fig. 71.

MACHAEROroES EOTHEN MATTHEW GAZIN 345

roides eotheii are the atlas, axis (badly crushed), and three others
including the seventh. The transverse processes are not complete
on the atlas, but sufficient remains to show that the groove or
notch at the anterior extremity for the anterior course of the
vertebral artery and the inferior branch of the spinal nerve is
not covered but widely open as in Thinocyon. The posterior open-
ing of the vertebrarterial foramen, however, is distinctly on the
posterior border of the transverse process, not dorsal to it as
described by Matthew for Thinocyon.

The two intermediate cervicals between the axis and the sev-
enth show a well-developed inferior lamella with the forward
ridge well separated from the forward extension of the trans-
verse process by the opening of the vertebrarterial canal. These
also show a well-developed forward projecting hyperapophysis
beginning superior to the postzygapophysis and extending for-
ward half to three-quarters of the way to articular surface of
the anterior zygapophysis. It is lower than the spine and more
compressed transversely.

The first eight dorsal vertebrae were found in articulation with
the last cervical, and the centra of seven additional vertebrae, not
in articulation, were preserved, at least four of which belong to
the lumbar series. The anterior dorsals show elongate transverse
processes for articulation with the ribs, and moderately high but
rapidly tapering spines. The spine of the first dorsal appears to
be the longest, and possibly the only one to have an anteropos-
teriorly expanded tip. The centra identified as lumbar are very
elongate, dorsoventrally flattened, and with the articular faces
markedly sloping, downward and backward with respect to the
longitudinal axis.

Limb bonpfi (pi. 46, c-f) . — The humerus of Machaeroides eothen
is distinctly larger and more robust than in Limnocyon verus. The
deltoid and ectocondylar ridges are wide and flaring, and both ex-
tend for a greater proportion of the length of the shaft than in L.
veiru?,. The development of the deltoid crest together with the high
and widely expanded acromion of the .scapula furnisher good lever-
age and indicates the importance of the deltoid muscU'. which func-
tions in raising the arm outward. The prominence and length of
the ectocondylar ridge denote good leverage for the supinator
longus in flexing and supinating the forearm. The greater and
lesser tuberosities are well developed with large rugose surfaces
for the muscles used in rotating the humerus. The distal end of the
humerus exhibits a large, oval entepicondylar foramen and well-
developed condyles, particularly the inner, which has a very rugose

346 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 95

anterodistal surface for attachment of flexor muscles for the
manus and forearm.

The radius and ulna are longer than in Limnocyon verus
and noticeably curved. Both elements are noticeably flattened
transversely and anteroposteriorly expanded in their distal por-
tions, although a part of this is recognized as due to crushing. The
olecranon of the ulna is relatively robust and bent somev^^hat
inward but is not so long as in L. verus. The shaft of the ulna is
not convex anteriorly, as described for L. verus, but, if anything,
is concave anteriorly. The radius shows a compound curve, con-
cave forward in the proximal portion and convex forward in the
medial and distal portions. The proximal portion of the radius
also has a well-developed bicipital tuberosity, turned slightly
inward from the shaft of the ulna, for insertion of the biceps.
Moreover, the distal portion of the radius is very much expanded
anteromedially above the prominent styloid process. This ex-
pansion supports the place of insertion for the distal end of the
supinator longus muscle which had its origin on the well-developed
supinator ridge of the humerus.

Remains of the manus include the scapholunar and the third
and fifth metacarpals. The scaphoid, lunar, and centrale are
fused, although a groove showing the line of separation between
the scaphoid and lunar can be seen across a portion of the radial
facet. The distal facets for articulation with the magnum, unci-
form, and trapezoid are deeply concave and separated from one
another by sharp angles. The metacarpals appear short and
stout, a characteristic noted in machairodonts in comparison with
true felids.

Table 2. — Measurevients (in millimeters) of the limb bones of Machaeroides
eothen, U. S. N. M. No. 17059

LetiBth of humerus 104

Anteroposterior diameter of proximal end of humerus ... 26.5

Transver.se diameter of proximal end of humerus across tuberosities 28

Transverse diameter of distal end of humerus 81,6

Greatest length of radius 78,g

Greatest diamster of proximal end of radius 13.5

Greatest diameter of distal end of radius ... I8.8

Lengtli of ulna 102

Anteroposterior diameter of olecranon below tuberosities 13.6

Greatest diameter of distal end of ulna 13

Greatest lenslh of scapholunar Ig

Length of metacarpal III 27

Length of metacarpal V 21

Length of femur parallel to axis of shaft 118

Diameter of proximal end of femur across head and greater trochanter, perpendicular

to axis of shaft ^ 35

Transverse diameter of shaft of femur at midsection 17.5

Transverse diameter of distal end of femur across condyles, excludine tuberosities 18

Transverse diameter of disUl end of femur across tuberosities IS

MACHAEROIDES EOTHEN MATTHEW — GAZIN 347

Both femora are preserved although the right is partly dam-
aged. This element is longer and sturdier than in Linmocyon
vents but otherwise has about the same curvature. The greater
trochanter, though exhibiting an enlarged, rugose surface, does
not project upward so far as in L. verus. The lesser trochanter
is a posterointernal knoblike process situated at about the same
height on the shaft as in L. verus, but the third trochanter is dis-
tinctly higher or more proximally located on the shaft, only
slightly lower than the lesser trochanter. The distd extremity is
relatively flattened on the dorsal surface and the patellar trochlea
not so long as in L. verus but with the depression carri'sd upward
on the shaft well beyond the termination of the articulating sur-
face.

The preserved portion of the tibia is relatively robust, and
curved, as described for L. verus, but the cnemial crest is promi-
nent and extends distally a greater distance. The proximal ex-
tremity is missing but the posterior surface shows a rather deep
groove between the ridges supporting the condyles, emphasized
in part by crushing. The distal extremity may not be entire. It
terminates in a simple, nearly flat surface v/ith a steep dorso-
lateral slope for articulation with the astragalus. The internal
mTlIcolus may not extend much below the facet for the astragalus,

t is incomplete.

^ U. S GOVERNMENT PRINTING OfFlCE ISa-S— 714083

U. S. NATIONAL MUSEUM

PROCEEDINGS. VOL 96 PLATE 45

Mnrharroidc. mtheu Matthew. Skull (U.S.N.M. No. 1T0.V..) : n, Dor.sal view;
I, iatfial view; r. ventral view; (/. occlusal view of upper cheek teeth
liiulKi'V Kocene, Wyoniintr. n-c three-fourths natural size; d, I'l: natural
si/e DrawiiiL' ('/) l>y Mrs. Ainie .M. Awl.

U. S. NATIONAL MUSEUM

PROCEEDINGS, VOL. 96 PLATE 46

-*<*-^«^r

Machaeroides eothen Matthew, a, b, Mandible (U.S.N.M. No. 17059) : a,
lingual view of left ramus; b, lateral view of right ramus, c-f, Limb bones
(U.S.N.M. No. 17059) : c, anterior view of left femur; d, posterior view of
left humerus; e, medial view of right humerus; /, lateral view of right
radius and ulna. Bridger Eocene, Wyoming, a, b natural size; c-f three-
fifths natural size.

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM
Vol. 96 Washington: 1946 No. 3203

REVIEW OF SOME CHALCIDOID GENERA RELATED TO
CEROCEPHALA WESTWOOD

By A. B. Gahan

This review of some genera related to Cerocephala Westwood
was occasioned by difficulty experienced in placing satisfactorily
specimens received for identification. It is hoped that the fol-
lowing treatment will eliminate some of the existing confusion
regarding these genera and make identification somewhat more
certain.

Cerocephala and its allies have been associated by most authors
with the genus Spalanf/ia Latreille to form a subfamily Spalangi-
inae or a tribe Spalangiini in the family Pteromalidae. In my
opinion Cerocephala and its allies are not closely related to
Spalanf/ia and should form a separate group for which I propose
the name Cerocephalinae. Although resembling each other in
many respects, the two groups have quite different host relations
and can be separated by several good characters. The Spalangi-
inae are all parasitic in dipterous puparia, while the Cerocepha-
linae so far as known are all associated with Coleoptera.

The Spalangiiiiae and Cerocei)halinae may be separated by the
following dichotomy:

Hind tibia with one calcariuni. Antennao inserted at the extreme anterior
nianfi'i of head, never scpaiated at base by a frontal carina; funicle always
7-seK-mentefl. Forewing without a callu.s or tuft of bristles at the proximal
end of the marginal vein and without transverse fuscous band.s; scutcllum
usually with a punctate cross furrow before apex Spalangiinac Westwood

Hind tibia with two calcaria. .Antennae in.serted well above extreme ante-
rior margin of hoad although frequently distinctly below ventral extremities
of eyes, always separated at base by a frontal carina or prominence;
funicle 5- or 6-8egmented, oi- rarely 7-segm«Mited in somo males. Forewing

349

350 PROCEEDINGS OF THE NATIONAL MUSEUM vol 96

always with a, callus at proximal end of marginal vein, this callus fre-
quently but not always bearing a tuft of erect bristles; usually with a
transverse fuscous band behind stigmal vein and another at juncture of
submarginal and marginal veins but occasionally without bands; scutellum
without a cross furrow before apex Cerocephalinae, new subfamily

Because of their bicalcarate hind tibiae the cerocephalines would
run to the family Miscogasteridae in Ashmead's key to families
of Chalcidoidea (Mem. Carnegie Mus., vol. 1, p. 228, 1904) , and in
that group, as constituted by Ashmead, they closely resemble
some apterous forms in the subfamily Lelapinae, from which they
differ principally by the absence of a neck on the propodeum and
by differences in the shape of the head. I do not consider the
Miscogasteridae to be separable as a family from the Pteromali-
dae. Instead, I think it should be combined with the Pteromali-
dae to form a large family made up of numerous smaller groups
including the Cerocephalinae, Spalangiinae, and Lelapinae among
numerous others.

Family PTEROMALIDAE
Cerocephalinae, new subfamily

Eight genera are here included in the Cerocephalinae. viz:
Choetospila Westwood, Theocolax Westwood, Cerocephala West-
wood, Theocolaxia Girault, Parcdaesthia Cameron, Acerocephala
new genus, Neosciatheras Masi, and Sciatherellus Masi. No
representatives of Paralaesthia, Neosciatheras, and Sciatherellvs
have been available for study, and they are placed in the generic
key solely on the basis of the descriptions. No doubt some genera
have been omitted that eventually may prove to belong here.

Four genera that have previously been associated by some
authors with Cerocephala and Spalangia have been excluded from
Cerocephalinae. These genera are discussed at the end of the
paper.

Description. — Head varying in shape from subcircular to dis-
tinctly oblong; below antennae usually concave, the margins of
concavity frequently armed with one or more sharp protuberances
or teeth on each side; if not concave then face with striae con-
verging toward clypeus ; antennae inserted at or beloM' middle of
head but always distinctly above clypeus and always separated
at base by a raised carina or narrow plate which is frequently
but not always produced anteriorly to form a sharp tooth or spine
between the antennae; malar groove absent; occiput carinately
margined above ; head attached to neck of pronotum very close to
vertex. Pronotum large, subconical, sloping from posterior
margin to neck without any abrupt angulations either dorsally or
laterally ; parapsidal grooves distinct and complete ; axillae either

CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN vSl

just touching at the median line or narrowly separated; scutellum
nearly flat, without a cross furrow; propodeum not declivous, its
dorsum lying in a plane only a little lower than that of scutellum,
without carinae or lateral folds and not produced into a neck
posteriorly; prepectus moderately large, triangular; mesopleuron
with a weak femoral impression. Wings either present or absent;
when winged, marginal vein of forewing as long as submarginal
or longer, with a distinct callus or thickening at its juncture with
submarginal, this callus often but not always bearing a tuft of
erect bristles ; stigmal vein never more than one-fourth as long as
marginal, usually much shorter; postmarginal vein never longer
than stigmal, usually weak, and sometimes absent; discal cilia
absent or vestigial but marginal cilia moderately long and dense;
hind wing unusually large with the marginellan vein usually
longer than submarginella. Forewing usually with one or two
fuscous areas or cross bands. Hind tibia with two slender,
unequal, and often very short spurs. Abdomen petiolate; oviposi-
tor usually shortly exserted, rarely half as long as abdomen;
gaster of female subcylindrical, sometimes flattened above or a
little compressed from the sides; of male compressed dorsoven-
trally and subtruncate at apex. Tegument of head and thorax
usually smooth and polished, occasionally sculptured.

KEY TO THE KNOWN GENERA OF CEROCEPHALINAE
j. Funicle 5-segTnented in the female, 6-segmented in the male; club solid
in both sexes. Forewing with a tuft of erect bristles at junction of
submarginal and marginal veins (apterous and subapterous individuals
of both sexes common) 1. Choetospila Westwood

Funicle 6-segmented in female, 6- or 7-segmented in male; club of female
either solid or indistinctly 3-segmented; club of male may be solid,
distinctly 2-8egmented or indistinctly 3-segmented. Foi-ewing with or
without a tuft of erect bristles at junction of submarginal and marginal
veins (apterous forms occur) 2

2. Head, pronotum, mesoscutum, and .scutellum sculptured; axillae obliquely

sulcate; forewing without a tuft of bristles at junction of marginal
and submarginal veins 7

Head dorsally, and mesoscutum entirely smooth and polished; pronotum
usually smooth but sometimes sculptured; axillae and scutellum usually
polished but sometimes more or less striated; forewing with or without
a tuft of bristles 3

3. Mandibles unusually long and conspicuous 6
Mandibles normal 4

4. Antennae inserted very distinctly below a line connecting lower extremi-

ties of eyes; head viewed anteriorly as long as broad or longer with
sides parallel; subapterous or fully winged; fully developed wing with
a tuft of bristles at juncture of submarginal and marginal veins, post-
marginal vein absent or represented by merely a short stub.

2. Th<?ocolax Westwood

352 PROCEEDINGS OF THE NATIONAL MUSEUM voua

Antennae inserted higher up, never more than slightly below a line con-
necting lower extremities of the eyes; head viewed anteriorly usually
narrowing below, rarely with the sides nearly parallel; forewing with
or without a tuft of bristles at juncture of submarginal and marginal
veins; postmarginal vein distinct but never longer than stigmal
vein 5

5. Foz'ewing with a distinct tuft of bristles at juncture of submarginal

and marginal veins 3. Cerocephala Westwood

Forewing with a callus but without a tuft of bristles at juncture of
submarginal and mai-ginal veins 4. Theocolaxia Girault

6. Mandibles bidentate at apex; forewing with a tuft of bristles at juncture

of submarginal and marginal veins . 5. Paralaesthia Cameron

Mandibles with four distinct teeth at apex; forewing with a callus but

without a tuft of bristles at juncture of submarginal and marginal

veins 6. Acerocephala, new genus

7. Female: Antennae clavate, inserted a little below middle of head; scape

not extending above vertex; first funicular joint about twice as long as
pedicel; postmarginal vein subequal to stigmal vein, the latter rather
short, the stigmal knob armed with a short tooth.

7. Neosciatheras Masi
Male: Antennae very long, filiform, inserted a little above middle of
head; scape extending above vertex; funicular segments and club
elongate, cylindrical; pedicel short; postmarginal vein nearly effaced;
stigmal vein long, straight, and with the stigmal knob scarcely devel-
oped and unarmed 8. Sciatherellus Masi

1. Genus CHOETOSPILA Westwood

Choetospila Westwood, Thesaurus entomologicus Oxoniensis, p, 137, 1874.
Choetospila Dalla Torre, Catalogus hymenopterorum, vol. 5, p. 207, 1898.
Spalmigiomor'pha GiRAULT, Mem. Queensland Mus., vol, 2, p. 333, 1913.

Choetospila has been treated as a synonym of Cerocephala
Westwood by Ashmead (Mem. Carnegie Mus., vol. 1, p. 369, 1904)
and others but may be easily distinguished from that genus by
the 5-segmented funicle of the female.

Spalangiomorpha was synonymized with Choetospila by Girault
(Insecutor Inscitiae Menstruus, vol. 5, p. 37, 1917) and later with
Cerocephala by the same author in one of his privately published
pamphlets [Indications (in new insects) of ruling power in
nature, p. 3, 1925]. In a still later publication Girault (Trans,
and Proc. Roy Soc. South Australia, vol. 53, p. 319. 1929) again
referred to the genotype species, using the generic name Spalan-
giomorpha but pointing out that the species agreed with Choeto-
spila as characterized in a table of genera by Masi (Nov. Zool.,
vol. 24, p. 188, 1917). I agree with Waterston (Rep. Grain Pest
Committee. No. 9, p. 25, fig. 13, 1921) that Spalangiomorpha is
certainly a synonym of Choetospila Westwood.

Tiipe of the genus. — Choetospila elegans Westwood.

CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN 353

CHOhTOSPlLA ELEGANS Wfstwood

Plate 47, Figures 1, la
Choetoapila elegaus Westwood, Thesaurus entomologicus Oxoniensis, p. 157,

pi. 25, fig. 10, 1874.-- Waterston, Rep. Grain Pe.«t Committee, No. 9, p.

25, fig. 13, 1921.
Spalangioinoi pha fasciativentris fJlRAULT, Mem. Queensland Mus., vol. 2, p.

334, 1913.
Spalaiigin metallica Fullaway, Proc. Hawaiian Ent. Soc, vol. 2, p. 286,

1013. (New synonjTny.)

Spalangiomorpha fasciativentris Girault is certainly the same
as Choetospila elegans, as was pointed out by Waterston in the
report cited above. Both the description and the habitat lead to
this conchision.

The type of Spalangia metallica Fullaway is in the collection of
the U. S. National Museum, and it differs in no way from typical
(\ ilcgaus.

Choetospila elegans is parasitic upon stored grain weevils and
has been distributed to many parts of the world in shipments of
stored products. The National Museum collection contains speci-
mens from various parts of the United States, Mexico, Puerto
Rico. Panama Canal Zone, Surinam, Peru, Hawaii. Guam, South
Australia, Java, India, Nigeria, and South Africa. Published
records include many additional areas where the species occurs.

Very little seems to be known about the actual biolog>^ of C.
elegans beyond the fact that it is usually associated with pests of
stored ^ain. Its most common host apparently is the rice weevil.
Sitophihi.'^ orifza (Linnaeus). 5?. grdnaria (Linnaeus). 5>. linearis
Herbst, Sitodrepa panicea (Linnaeus), Caulophilus latinasus Say,
Callofioh'nichus quadrinmcidatns (Fabricius), and C. chinensis
(Linnaeus) have been recorded as attacked by it. Very likely
other grain-infesting beetles may serve as hosts, but no authentic
records of such hosts have been published. A paper by Corbett
and Miller (Federated Malay States Dept. Agr. Sci.. ser. 1P>, p. 4.
H).3.''>) recording this species as a parasite of Sitotroga cerealella
Olivier is not available for judgment as to the authenticity of the
record. Squire (T)iel Rep. Dept. Agri. British Guiana for 1924,
pp. 121 124, 1925) records C\ rhftnns as parasitizing an uniden-
tiled scolytid infesting British Honduras mahogany tn^es (Sirri-
frnia /??/?«?7i,s-) in Guiana, and in the U. S. National Museum
collection are 14 specimens of C. rlninyix labeled as having been
reared by J. Zetek in 1924 in the l^anama Ganal Zone from a
scolytid-infested piece of "Amargo" wood iVntnirra sp.).

The species is sufficiently characterized by the descriptions
and figures given in the above-cited papers by Westwood and
Waterston.

354 PROCEEDINGS OF THE NATIONAL MUSEUM VOL 96

CHOETOSPILA PRATER (Girault), new combination

Spalangiomorpha frater GiRAULT, Mem. Queensland Mus., vol. 2, p. 334, 1913.

No representative of this Australian species is available for
study. According to the description it differs from elegans prin-
cipally by having the first funicular joint wider than long.

CHOETOSPILA TABIDA, new species

Plate 47, Figures 2, 2a

This species is very similar to elegans but differs by having
distinctly shorter antennae, by being entirely wingless in both
sexes (so far as shown by the specimens at hand) , by the propo-
deum being smooth and polished, by the ovipositor of the female
not extending beyond the apex of the abdomen, and by the an-
tennal club being unsegmented in both sexes.

Female. — Length 1,6 mm. Mostly smooth and polished, the
head anteriorly, below level of insertion of antennae, weakly
wrinkled, and the mesopleura reticulated.

Head viewed from in front a little longer than broad, nearly
squarely truncate at vertex and at mouth, and with the sides
practically parallel; compound eyes very small, subobsolete;
ocelli represented merely by minute punctures ; antennae inserted
low down at approximately the lower fifth of head, separated at
base by a sharp carina ; scrobes short and shallow, poorly defined ;
face slightly depressed medially, with a small toothlike projection
at the anterior margin on each side of clypeus ; mandibles rather
broad and stout. Head viewed laterally forming a rather broad
ellipse, approximately twice as long as broad.

Antenna 8-segmented, strongly clavate, short, its total length
only slightly greater than the frontal length of head ; scape short,
slightly thickened, about three times as long as broad; pedicel
about half as long as scape and half as long as funicle; funicle
5-segmented, the segments all transverse and gradually increasing
in width toward club; club ovoid, large, very nearly as long as
entire funicle, approximately twice as long as broad and appar-
ently not segmented.

Thorax a little narrower than head, approximately twice as long
as broad; prothorax subconical, rounded at the sides, more than
twice as long as mesoscutum; parapsidal grooves complete, very
oblique; scutellum as long as mesoscutum, nearly circular; meso-
pleuron weakly reticulated; metapleuron polished; propodeum
slightly longer than scutellum, weakly convex dorsally, without
grooves or carinae, smooth and polished except for some very
faint reticulation basad of the spiracles. Wings entirely absent.

Legs rather stout, the middle pair obviously somewhat shorter

CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN ^>S

and weaker than the other two pairs; anterior femur obviously
.swollen and its tibia a little thickened, each with a few moderately
longr hairs on its ventral margin and a few shorter ones on its
(^uter face; posterior femur slijjrhtly swollen, with a few short
hairs; posterior tibia slender at base but gradually increasing in
thickness toward apex, sparsely hairy and with two rather long-
and slender but unequal calcaria ; tarsi not thickened ; posterior
basitarsus equal to half the length of tibia.

Abdomen nearly as long as head and thorax combined, a little
broader than thorax ; petiole as long as broad, smooth ; gaster
elliptical in outline, twice as long as broad, smooth and polished
dorsally and ventrally, nearly devoid of hairs dorsally but with
numerous rather long slender bristles at apex beneath ; first seg-
ment of gaster emarginate medially: ovipositor not protruding
Ijeyond apex of abdomen.

Head, thorax, legs, and abdominal petiole brownish yellow;
anterior and posterior coxae whitish except dorsally; antennae
concolorous with head except that club is usually, thouph not
always, blackish; gaster wholly black or blackish.

Male. — Length 0.9 mm. Antennae uniformly testaceous, 9-
.segmented ; funicle 6-segmented. the first four segments very short
and difficult to distinguish, the last two larger; club solid, conical,
a little longer than two preceding segments combined; abdomen
about as long as thorax. Otherwise like the female and difficult
10 distinguish from it.

Type locality. — University Park, Md.

Type.—U. S. N. M. No. 57279.

fiemarks. — Described from five females (one holotype) and one
male collected by W. H. Anderson as pupae from the burrows of
an unidentified cossonine beetle infesting a dead branch of an
unidentified tree and reared to adults.

2. Genus THEOCOLAX Wcstwood

'! hfocalax Westwood, Philos. Map., ser. .3, vol. 1. p. ll!7. IhV'.
Lnrythia Halihay, Ent. .Map., vol. 1, p. 3:55, 18.3.3.

Theocolax is very similar to Crrocephala, and it has been treated
as a synonym by Walker (Ent. Mag., vol. 2, p. 148, 1833). It
may be distinguishod from Cmirrphahi, howt'vor. l\v h;;ving the
antennae inserted distinctly below a line connecting the lower
extremities of the eye.s, the head viewed anteriorly as long as
broad or a little lon^rer, parallel-sided, and never with more than
three small toothlike projections, one between the antennae and
one on each side of the facial depression just above the base of
each mandible. In some species the processes above the bases

356 PROCEEDINGS OF THE NATIONAL MUSEUM VOL 96

of the mandibles are absent. Both subapterous and fully winged
individuals occur. In the fully winged individuals there is a
distinct tuft of erect hairs at the juncture of the marginal and
submarginal veins of the forewing, exactly as in Cerocephala.
The postmarginal vein is absent or represented by merely a short
stub.

Type of the genus. — Theocolax foiiniciformis Westwood.

THEOCOLAX FORMICIFORMIS Westwood

Plate 47, Figs. 3, 3a; Plate 48, Figs. 1, la, lb

Theocolax formiciformis Westwood, Philos. Mag., ser. 3, vol. 1, p. 127, 1832;

Thesaurus entomologicus Oxoniensis, p. 138, pi. 25, fig. 11, 1874.
Laesthia vespertina Haliday, Ent. Mag., vol. 1, p. 336, 1833.
Cerocephala formiciformis (Westwood) Walker, Ent. Mag., vol. 2, p. 149,

1834.

I examined Westwood's type of this species in the Hope
Museum at Oxford, England, in 1927 and compared with it a
specimen from Blankenburg, Thuringia, previously identified by
Schmiedeknecht. This specimen, now in the U. S. National Mu-
seum, appeared to me to be homotypic. The National Museum
collection also contains one specimen identified by Ruschka and
said to have been reared from Anobium striatum Olivier at Stock-
holm, Sweden. This specimen is considerably paler in color than
the homotype but does not seem to differ otherwise. In addition
to these two specimens the collection possesses a series of 14
specimens from Auckland, New Zealand, said to have been reared
from Anobium-mf ested timber by E. Bollard and D. Spiller.
Apparently this series does not differ materially from the homo-
type except that one female in the lot has fully developed wings.
Except for the perfectly developed wings, this winged specimen
is exactly like the subapterous females.

Although the species has usually been described as wingless, as
a matter of fact, in none of the individuals I have seen are the
wings completely absent, usually being represented by small stubs
approximately twice the length of the tegula. In the fully winged
individual the postmarginal vein is represented by merely a very
short stub ; the marginal vein is a little longer than the submar-
ginal; the stigmal vein is approximately one-sixth as long as
marginal, slightly curved and not thickened at apex; the disk of
wing is nearly bare ; the marginal cilia are moderately long, and
there is an erect tuft of black bristles at the junction of marginal
and submarginal veins. There is a broad fuscous cloud embracing
the apical half of the marginal and all of the stigmal vein and
extending across the wing. The hind wing is about two-thirds as

CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN 357

wide as the forewing and rounded at its apex. The propodeum is
weakly reticulated and slightly shining.

Theocolax formiciformis appears to be a common parasite of
Anohium. having been recorded from several species of that
genus. It is also said by several authors to attack (Hylesinus)
Leperisinus fraxini (Panzer), but since it is known to have been
confused in some instances with Cerocephala corniyera Westwood.
a common parasite of this host, it seems very likely that all of
the records of Theocolax formiciformifi from Leperisiuus fraxini
actually refer to Cerocephala cornigea as was suggested by
Waterston (Rep. Grain Pests Committee, No. 9. p. 12. 1921).

The species is known to occur in England, Sweden. Germany,
and New Zealand and may be much more widely distributed. It
is not known to occur in America.

THEOCOLAX BAKERI (Crawford), new combination

'^rrocephnla bakfri CRAWFORD. Philippine Journ. Sci., vol. 0, p. 4t:10. 1914.

This Philippine species differs from formiciformis by being
distinctly smaller, by lacking the toothlike protuberances on each
side of the facial depression, and by the pedicel in the female
being as long as the first and second segments of the funicle com-
bined. It is also somewhat paler in color. All known specimens
of the species are fully winged and have a conspicuous tuft of
bristles at the .junction of the marginal and submarginal veins.
The head is oblong with parallel sides and with the antennae
inserted distinctly below the ventral extremities of the eyes.

There are eight specimens in the U. S. National Museum,
inclusive of the type series, all taken by C. F. Baker at Los Banos,
Philippine Islands. One of these bears a label "on scolytid No.
847."

THEOCOLAX LITIGIOSA (Kondani)

Laeathia liiigiosa Rondani, Ann. Soc. Nat. Modena, vol. 1, p. 23. 1866: Arch.

Zool., vol. 4, p. 191, pi. 7, fijf. 7-8, 1866; Bull. Soc. Ent. Ital., vol. 9, p. 183,

pi. 1, fig. 20, 1877.
Theocolax litigioHn Dalla Torre, Catalogus Hymenopteroi iim. vol. 5, p. 207,

1898.

It is impo.ssiblc to recognize this species from the short and
very unsatisfactory descriptions by Rondani. The alleged host
(a species of Cecidomyiidae) justifies a doubt that Htigiosa be-
longs in Theocolax.

3. (icnu.s CKKOCEl'liALA WcHtwood
Cerocephala Westwood. -Mag. Zool., vol. 2, CI. ix, pi. 4, 1832.
Kpimacrug Walker. Ent. Mag., vol. 1, p. 368, 1833.

Sciatheras RATZEBrKf;, Die Ichneumonen der Forstin.'^ectcn . . ., vol. 2, p. 209,
1848.

358 PROCEEDINGS OF THE NATIONAL MUSEUM vml 9«

Pam.-:ciatheras Masi. Nov. ZooL, vol. 24, p. 189, 1917. (New synonymy.)
Proamotura GiRAULT, Insecutor Inscitiae Menstiuus, vol. 8, p. 143, 1920 (new
synonymy).— DoDD (in part), Mem. Queensland Mus., vol. 9, p. 66, 1927.

On a succeeding- page I have indicated that the genotypes of
Epimacrus and Sciatlieras are both synonyms of Cerocephala
comigera Westwood, the genotype of Cerocephala. This generic
synonymy has been recognized by most authors for many years.

Paras ciather as was proposed as a subgenus of Cerocephala. The
characters given in Masi's key for distinguishing the subgenus
from Cerocephala do not function satisfactorily, and while there
are certain differences between the two genotypes in head char-
acters, these seem to be of no more thaii specific value. Accord-
ingly the subgenus is suppressed.

Proamotura, described by Girault as a genus of Cleonymidae
but compared by its author with Spalmigia Latreille and Spalan-
giomorpha Girault, differs from Cerocephala only in minor head
characters and some details of sculpture. Such differences as
exist are at most of specific value only. Dodd redescribed
Proa7notura aquila Girault, the genotype, and placed in the genus
three additional Australian species. All the species treated by
Dodd, except the genotype, lack the tuft of erect bristles at the
apex of the submarginal vein and are here transferred to
Theocolaxia Girault.

CEROCEPHALA CORNIGERA Westwood

Plate 47, Figs. 4, 4a; Plate 48, Fig. 3
Cerocephala comigera WESTWOOD, Mag. ZooL, vol. 2, CI. IX, pi. 4, 1832. —

Walker, Ent. Mag., vol. 2, p. 149, 1834.
Epimacrus rufus Walker, Ent. Mag., vol. 1, p. 369, 1833.
Sciatheras trichotus Ratzeburg, Die Ichneumonen der Forstinsecten . . ., vol.

2, p. 209, pi. 3, fig. 1, 1848.

Epimacrus rufus Walker was synonymized with C. comigera
by Walker, and this synonymy has been generally accepted.
Although I have not seen Walker's type, I believe the synonymy
to be correct.

Sciatheras trichotus Ratzeburg: has been considered a synonym
by most authors but Masi (Nov. ZooL, vol. 24. p. 188, 1917)
expresses the opinion that it may be different from comigera. In
1927 I examined the type of Sciatheras trichotus in the Ratzeburg
collection at the Forstliche Hochschule in Eberswalde, Germany,
and made the following notes upon it:

Female. — Head thick, occiput margined ; a prominent toothlike
carina between bases of antennae; face concave below antennae,
with two small prominences on each lateral margin of the exca-
vation ; antenna strongly clavate, flagellum gradually increasing in

CHALCIDOIDEA RELATEH TO CEROCEPHALA — OAHAN ?>59

thickness from base of fiinicle to club, the latter pointed-ovate;
rirst funicular sej2rment one and one-half times as long as broad,
second and third about as long as broad, fourth to sixth more or
less slightly broader than long: club apparently solid, a little
longer than two preceding segments; mesonotum smooth, par-
apsidal grooves deeply impressed, axillae smooth; scutellum nearly
Hat. smooth, except for some very minute and obscure reticulations
apically. Propodeum granularly rugulose. without carinae, folds,
or spiracular sulci, and nearly horizontal. Abdomen as long as
thorax or a little longer, slightly compressed, shortly petiolate;
ovipositor not quite half as long as abdomen. Marginal vein of
forewing nearly as long as submarginal, with a tuft of black hairs
at juncture of the two veins; postmarginal and stigmal veins short
and subequal.

Subsequently during a visit to the Hope Museum at Oxford,
Kngland. I saw the type of Cerocephalo cnmigera Westwood and
compared it with the above notes. This type is a male. Except
for sexual characters it agreed with the notes on the Ratzeburg
type.

Under the name C. comigera in the British Museum general
collection I found one headless female, and in the British collec-
tion one male also agreeing with my notes on the types.

The U. S. National Museum collection contains one female and
a male reared by H. L. Parker in April 1936. at Hyeres. France,
from Scolytif.^ nudtistriatvf' (Marshall) and identified by me as
C. comigera. The female agrees with my notes on the Ratzeburg
type of trichotvf! as well as with the Ratzeburg description, and
the male apparently differs from the type of comigera only by
having the thorax entirely dark ferruginous instead of more or
less fuscous or blackish on the scutellum and propodeum. This
difference is believed not to be of specific significance since in
other related species there is known to be a similar variation in
color.

There is but little doubt that Chftetospila elegants Westwood and
Thcocolax formic if or mis Westwood have both been confused with
(.'erocephala comigera in the literature. As pointed out by
Water.ston [Rep. Grain Pests Committee, No. 9. p. 12. 1921], the
records of this species as a parasite of stored grain pests almost
certainly refer to f'horfosp'la cJegans. The record in Dours (Cat.
Syn. Hym. France, p. 92, 1871) of C comigera from Anofiiurti
pertinux Fabricius very probably refers to Thencolax formici-
formis. The li.sting of comigera by Giraud and Laboulb^ne
(Ann. Soc. Ent. Fiancf. sf-r. .'>. vol. 7, p. 422. 1877) as a parasite

360 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 96

of Aphidae and of Odynerus sp. is quite certainly erroneous. The
species apparently normally attacks scolytids of the genera
Leper isinus, Scolyttis, Chaetoptelius, and Phloeotrihus . The most
complete description of the species is that by Russo (Boll. Lab.
Ent. Agr. Portici, vol. 2, pp. 206-215, figs. 105-110, 1938).

So far as known, C. cor7iigera is confined in its distribution to
the European continent and the British Isles.

CEROCEPHALA ECCOPTO«ASTRI Masi

Ceroeephala eccoptogastri Masi, Ann. Mus. Civ. Stor. Nat. Genova, ser. 3,
vol. 9, pp. 189-193, fig. 7, 1921.

Judged by Masi's description and figure as well as by the
indicated host this species is very likely a synonym of C. cornigera
Westwood. The types are said to have been reared from
Eccoptogaster (probably rugulosus Ratzeburg) taken at Bengasi,
Cyrenaica.

The figure published by Gonzales Ceballos (Las tribus de los
Himenopteros de Espano, p. 204, 1943) under the name Cero-
cephala eccoptogastri Ratzeburg does not agree completely with
Masi's description and figure. The head appears longer and
apparently has only one tooth, instead of two, on each lateral
margin of the facial depression. If the drawing is accurate it
probably represents a species different from eccoptogastri.
Ceballos has evidently confused the specific name with that of
Pachyceras eccoptogastri Ratzeburg, an insect quite different
from the one he figures.

CEROCEPHALA DINODERI Gahan

Plate 47, Figure 5
Ceroeephala (Parasciatheras) dinoderi Gahan, Philippine Journ. Sci., vol.
27, p. 100, 1925.

As was pointed out in the original description, dinoderi is ap-
parently very similar to caelehs Masi. The former was described
from a unique female and the latter from a unique male. Subse-
quent to the description of dinoderi, the C. F. Baker collection of
Hymenoptera was acquired by the U. S. National Museum, and
in it were seven specimens of this species, one of them a male.

This male apparently differs from the description of caelehs
by having the antennal flagellum much less conspicuously hairy,
the hairs being very sparse and short, none of them as long as
the segment from which it arises. The antennal club is much
less distinctly segmented than in Masi's illustration of the antenna
of caelehs, the dividing furrow being very shallow and indistinct.
The striations on the face are present but appear to be somewhat
weakei- than illustrated for caelehs. The head dorsally, pronotum

CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN .^61

posteriorly, mesoscutum, axillae, scutellum. and paster are dark
piceous with violaceous or metallic tints on the head and mesoscu-
tum. The remainder of head and thorax, propodeum for the most
part, and the abdominal petiole are rufotestaceous, the coxae all
whitish and the remainder of legs somewhat paler than the
underside of thorax but darker than the coxae. The abdominal
petiole is a little longer than the posterior coxa, paler at base and
at apex than in the middle, cylindrical, and granulosely punctate.
The antennae are about twice as long as the length of the head,
not at all clavate, consisting of a subcylindrical scape, a pedicel
not quite twice as long as broad, one strongly transverse ring
joint, seven funicular segments, and a club which is indistinctly
2-segmented, no thicker than the funicle, and about equal in length
to the two preceding funicular segments. Funicular segments
1 to 4 are subequal, each about as long as the pedicel and narrowed
at base; segments 5 to 7 are very slightly shorter and more com-
pact. The sensilla are rather coarse and extend the whole length
of the segments.

Described originally from Mount Maquiling, Luzon, Philippine
Islands, as a parasite of Dinoderns minutua (Fabricius), this
species has since been received from Buitenzorg. Java, where it
is said to have been reared from Calundra oryzae (Linnaeus) by
R. Awibowo. Other specimens in the collection, without host
records, are from Laguna and Los Baiios, Philippine Islands, and
Deli. Sumatra.

CEROCEPHALA CAELEBS Masi

Crroerphala (Paiasciathcras ) caelebs Masl Nov. Zool., vol. 24, p. 189, figs.
45-48, 1917.

This species was de.scribed from one male specimen collected
in the Seychelles Islands.

It apparently differs from dinoderi Gahan mainly in the more
distinctly hairy antennal flagellum.

CEROCEPHALA AQUILA (Girnult)

Plate 48, Figure 2
Prnamotura aqiiila GiRAULT, Insecutor Inscitiac Mcnstruus, vol. 8, p. 143,
1920.— DoDD, Mem. Queensland Mus., vol. 9, p. 67, 1927.

Cerocephala aqitila very closely resembles dinoderi but may be
distinguished from that species, as well as all others, by the com-
pletely striated dorsum of the prothorax.

Head viewed from in front nearly circular; antennae inserted
above lower extremities of eyes; face depressed medially, mostly
smooth within the depression, and with a more or h>ss distinct
carina running down the middle; lateral margins of the facial
depression with a broad, shallow, not very distinct incision oppo-

362 PROCEEDINGS OF THE NATIONAL MUSEUM vc.l %

site the base of each antenna but without any distinct teeth or
projections; laterad of the depression and below the antennae
rather strongly striated; front above antennae, vertex, temples,
and cheeks mostly smooth and polished ; scrobicular grooves short
and weakly reticulated, separated by a longitudinal carina which
originates just below the anterior ocellus and which is flattened
and produced to form a sharp prominence between the bases of
the antennae. Antennae rather distinctly clavate in the female,
filiform in the male. Pronotum dorsally finely and strongly longi-
tudinally striated; mesoscutum and axillae perfectly smooth and
polished, except in the grooves which are weakly foveolated:
scutellum smooth medially but distinctly longitudinally striated
laterally; propodeum rather strongly ruguloso-reticulate, occa-
sionally with indications of a median carina but more often with-
out. Abdominal petiole of female about half as long as posterior
coxae, of male equal in length to hind coxae ; gaster in both sexes
smooth and shining, in female about as long as thorax, in male
much shorter than thorax; basal segment of female gaster tri-
angularly emarginate medially, of male not emarginate; ovipositor
sheaths about as long as the petiole. Forewing in both sexes with
marginal vein very nearly as long as submarginal ; postmarginal
vein slightly shorter than stigmal ; discal cilia weak, marginal cilia
moderately long; tuft of hairs at apex of submarginal vein strong.
Color of head and thorax varying from mostly reddish brown to
mostly yellowish testaceous, the gaster and ovipositor sheaths
usually black or very dark brown ; antennal club black, remainder
of antenna and all legs concolorous with the head and thorax.
Anterior wing with a small fuscous cross band at apex of sub-
marginal vein, and a large one embracing the apical half of
marginal vein and all of stigmal vein.

Girault described this species from specimens reared from
beetle-infested twigs of Mallotus philippinensis collected at Bris-
bane, Australia. Dodd gave a more complete description and
reported the species from two additional Australian localities.
One specimen from Ayr, North Queensland, Australia, identified
by Dodd, is now in the U. S. National Museum.

Besides this Australian specimen the National Museum now
possesses the following material which I have identified as aquila:
Two females from Sigatona, Fiji, "ex lyctid." R. A. Lever, coll..
December 1942; one male from Mount Maquiling, Luzon, reared
from Dmoderu,s minutus (Fabricius) in bamboo by C. F. Baker;
one male from Los Baiios, Luzon, "ex bostrichid on bamboo."
reared by S. M. Candana; one male from Laguna, Luzon, collected

CHALCIDOIDEA RELATED TO CEROCEPHALA GAHAN 363

on laboratory window by D. T. FuUaway and I. Dobrosky; four
females from Santiagro de las Vetras. Cuba, reared from larvae of
D. minutus by A. Otero, January 26, 1933, under Ace. No. 9802a;
and a large series of specimens reared by E. A. Chapin from stems
of Arimdinaria longifolia (bamboo) infested with Dinoderics
miiiutns, Lyctus sp.. and two or three other species of Coleoptera.
the bamboo stems having been discovered in storage in Hoboken.
X. J., but said to have originally come from Mexico. Of the sev-
eral species of Coleoptera reared from these stems, individuals of
Dinoderus minutus were by far the most abundant and in all prob-
ability this species was the actual host of the parasite.

In view of the habit of this species of attacking larvae infesting
bamboo, it seems likely that it will eventually be found to be much
more widely distributed than the above few records show.

4. Genus THEOCOLAXIA Girault

Theocolaxia Girault, Lese Majeste, new Insecta, and robbery, p. 1, 1924.

'■ afomus DoDD (not Dalman), Trans. Royal Sec. South Australia, vol. 48, p.

170, 1924.
Proamotura (Girault) DoDD (in part), Mem. Queensland Mus., vol. 9, p.

66, 1927.

Theocolaxia was described in a single page, privately published,
pamphlet printed at Gympie, Australia. The genus is monobasic,
with T. lessiiif/i Girault as its type. The short generic description
is as follows :

"Theocolaxia nov. (Spalangiinae). Antennae 9-jointed, club
solid, larger. Wings as N eosciatheras as to shape, venation, fringes.
Pronotum square, exceeding mesuscutum. Parapsides shorter
than scutum, convex. Axillar sutures meeting cephalad. Scutel-
lum plane. Propodeum rugulose, irregular median and lateral
carinae. Petiole short, abdomen ovate, equal rest, ovipositor 44
it. Jaws 4-dentate."

Earlier in the same year A. P. Dodd described two Au.stralian
species in the genus Cratomy^s Dalman, but in 1927 he discovered
they did not belong in Dalman's genus and transferred both spe-
cies to Proamotura Girault. In the latter paper he described one
new species and two new varieties, redescribed P. aquila the
genotype of Proamotura, and gave a key for separating the spe-
cies. According to the descriptions, all three of the species
erected by Dodd differ from aquila, Girault by lacking the tuft of
erect bri.stles at the apex of the submarginal vein. Absence of this
tuft is the mo.st striking character differentiating the genus
Theocolaxia from Cprocephala of which Proamotura is here con-
sidered to be a synonym. Accordingly I have transferred the
three species de.scribed by Dodd to Theocolaxia and placed Pro-
amotura aquiln Girault in Ceroceph/ila.

364 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

Two North American species described by Ashmead in Cero-
cephala also lack the tuft of hairs on the forewing and seem to
agree, except in one respect, with the generic characters given in
the descriptions by both Girault and Dodd. The male antennae
of the only Australian species of which the male was known were
described by Dodd as being 11-segmented consisting of a scape,
pedicel, one ring joint, seven funicular segments, and an appar-
ently solid club. A specimen identified by Dodd as a variety of
that species is in the U. S. National Museum, and it agrees with
that description. In males of the single North American species
of which this sex is known, the antennae are 10-segmented, con-
sisting of scape, pedicel, six funicular segments, and a distinctly
2-segmented club. No ring segment is visible, and the seventh
segment beyond the pedicel, while closely resembling a funicular
segment, is obviously more closely united with the apical segment
than with the preceding and forms a part of the club. The an-
tennae of the females of this species apparently do not differ
from those of females of the Australian species, and the difference
in the one sex I do not believe to be of generic importance. ;

THEOCOLAXIA LESSINGI Girault

Theocolaxia lessingi Girault, Lese Majeste, new Insecta, and robbery, p. 1,
1924.

This species is known only from the original description which
is as follows: "T. lessingi nov. Black, glabrous; knees, tibia 1
beneath, tips tibiae, petiole red. Coxa 3, basal ys ovipositor, white,
also apex club. Antennae red save club, base pedicel, funicles
5-6. Fore wing with cross-stripe distal half marginal to apes
stigmal, also a central spot off bend of submarginal. Funicle 1
equal pedicel. Sand dunes. Main Beach, Southport, May 5, 1924.'

The type is probably in the Queensland Museum.

THEOCOLAXIA VIRIDINOTUM (Dodd). new combination

Cratomus viridinotum DoDD, Trans. Royal See. South Australia, vol. 48, p

171, 1924.
Proaviotiira viridinotum Dodd, Mem. Queensland Mus.. voL 9, p. 72, 1927.

According to Dodd this species is distinguished from the othei
Australian species by the presence of only one fuscous band oi
the forewing.

It was described from Lord Howe Island, Australia.

THEOCOLAXIA INSULARIS (Dodd), new combination

Cratomus insularis DoDD, Trans. Royal Soc. South Australia, vol. 48, p. 171

1924.
Proamotura insularis (Dodd) Dodd, Mem. Queensland Mus., vol. 9, p. 68, 1921

Described from specimens said to have been reared from rottei
wood, found on Lord Howe Island, Australia.

CHALCIDOIDKA RELATED TO CEROCEPHALA — GAHAN 365

THEOCOLAXIA INSl'LARIS rar. GRANDIS (Dodd). n«w combination
Proaviotura insularis fjrayidia DoDD, Mem. Queonslunu Mus.. vol. 9, pp. 64, 68,
figs. 2. 3. 1927.

Both sexes of this variety were described from North Queens-
land, Australia, where they were collected on tree trunks.

One male paratype of this form was received from its author
and is now in the National Museum collection. The antenna of
this specimen is 11-segrmented, the scape rather thick, pedicel
slightly longer than broad, ring segment fully as long as broad
and narrower than the following segment, funicle 7-segmented
with all its segments about equal in width and diminishing slightly
in length from first to last, the club solid but with slight indica-
tion of a cross furrow and about twice as long as but no wider
than the preceding segment. The head has a broad elevation
between the bases of the antennae, but this is not produced
into a prominent tooth or spine as in most other species. The face
is moderately convex and strongly striated, with only a very
shallow groove running from each antennal fossa to the clypeus,
the margin.^ of these grooves rounded and without any sharp
angles or protuberances. The scrobes are moderately deep with
sharp lateral margins and extend very nearly to the anterior ocel-
lus. The vertex is perfectly smooth but the frons between the
scrobes and the eye margins is striated like the face. The area be-
hind the eyes is sparsely covered with deep punctures. The whole
dorsum of the thorax is polished, except that the parapsidal grooves
are weakly foveolated and the axillar groo^/es strongly so. The pro-
podeum is ruguloso-reticulate without distinct carinae. The ab-
dominal petiole is cylindrical and distinctly longer than the hind
coxae. The wings are ample, with the marginal vein distinctly
much shorter than the submarginal, the postmarginal about as
long as the stigmal.

The color is nearly uniformly black with a slight metallic tinge,
the hind coxae, all tibiae and tarsi and the underside of abdomen
more or less fuscotestaceous. The wings are hyaline with a
fuscous band beneath the stigmal vein but without any indication
of a band behind the apex of submarginal vein.

Dodd states that after comparing this form with typical
infii/laris, no specific differincos could be found. On account of
the larger size (insularis 2.75 mm., grandis 4 to 4.75 mm.) and
the darker wings, he thought it was advisable to separate this
mainland form from the Lord Howe Lsland in.sect.

THEOCOLAXIA PRRPUI.CHRA (Dodd). new combination

Proamotura perpulchra Dodd. Mem. Queensland Mu.s., vol. 9, p. 70, 1927.
Described from one female collected at Mount Tambourine.

366 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 96

South Queensland, Australia. In Dodd's key to species, per-
pulchra is separated from insularis on the basis of a difference
in the length of the abdominal petiole. This is said to be plainly
longer than wide in perpulchra while females of insularis are
said to have it transverse. The scutellum of perpulchra is said
to be dull metallic green.

THEOCOLAXIA PERPULCHRA var. METALLICA (Dodd), new combination

P loaniotura perpulchra metallica Dodd, Mem. Queensland Mus., vol. 9, p. 71,
1927.

Described from collected specimens taken in Cairns district,
North Queensland, Australia. According to Dodd this form is
distinguished from typical perpulchra by its more extensive me-
tallic coloration.

THEOCOLAXIA SCOLYTTVORA (Ashnead). new combination

Plate 47, Figures 7, 7a
Cerocephala scolytivora (Ashmead MS.) Riley and Howard, Insect Life, vol.

4, p. 122, 1891. (Nomen nudum.)
Cerocephala scolytivora AsHMEAD, Proc. Ent. Sec. Washington, vol. 3, p. 33,

1894.

This species was described from southern Florida as a parasite
of the scolytid Loganius ficus Schwarz. The types are in the U. S.
National Museum.

Female. — Length 1.8 mm. Head viewed from in front very
slightly longer than broad, distinctly convex in outline dorsally
and less strongly so below; antennae inserted very nearly on a
line with ventral extremities of eyes; face below antennae dis-
tinctly concave, the depression sharply margined laterally; be-
tween bases of antennae is a strong protuberance which is flat-
tened and delicately margined above and terminates in a sharp
and slightly upturned point; margin of facial depression with a
shallow incision laterad of base of antenna, the lower angle of
this incision slightly prominent but not forming a distinct tooth ;
scrobes not extending to the front ocellus; ocelli in an obtuse
triangle, ocellocular line about equal to distance between posterior
ocelli ; eyes not prominent, equal in length to approximately half
the length of head. Vertex, upper part of frons, cheeks, and
whole area behind eyes smooth and polished; frons laterad of
scrobes and face laterad of the depression weakly rugulose; area
within facial depression mostly smooth.

Antennae consisting of scape, pedicel, six funicular segments
and club. Scape bottle-shaped, rather short, slender at base,
rather abruptly thickened beyond basal one-third ; pedicel slightly
longer than broad ; first funicular segment as long as pedicel and
a little longer than broad, slightly longer than second; segments

CHALCIDOIDEA RELATFD TO CEROCEPHALA — GAHAN 367

2-6 subequal in lenprth but successively increasing: very sli.crhtly
in width, the second about as broad as lonjr and the sixth only a
little broader than long; club indistinctly S-segrmented, ovate,
and about equal in length to two preceding segments.

Thorax mostly smooth and polislied. the neck of pronot'jm,
mesopleura. and mesosternum weakly reticulated ; pronotum
longer than mesoscutnm and as long as broad or a little longer;
parapsidal grooves not foveolate; scutellum about as long as
mesoscutum, nearly flat; axillae very nearly meeting; axillar
grooves weakly foveolated and distinct but not deeply impressed ;
propodeum finely and shallowly reticulated, dull, without lateral
folds or median carina. Anterior and posterior femora rather
broad; middle femur more slender; posterior tibia with two slen-
der, unequal calcaria. Anterior wing about three times as long
as broad, nearly devoid of discal cilia, the marginal fringe moder-
ately long; submarginal and marginal veins nearly equal, marginal
about seven times as long as stigmal, postmarginal a little shorter
than stigmal.

Abdomen nearly as long as head and thorax combined; petiole
distinctly longer than broad, about two-thirds as long as pos-
terior coxae, broader at base than at apex, flattened and smooth
dorsally; gaster a little narrower than thorax, slightlv compressed
from the sides, smooth and polished, the first tergite the longest
and deeply incised medially; ovipositor sheaths exserted approxi-
mately one-third the length of gaster and slightly com.pressed.

General color yellowish testaceous, the mesonotum slightly
darker than the pronotum, the apical two-thirds of abdomen dor-
sally, dark brown; antenna nearly uniformly reddish testaceous,
its club sometimes a little darker; legs generally very slightly
paler than thorax, their coxae often more or less whitish ; wings
hyaline with a broad fu.scous band behind apex of venation and a
narrow indistinct one at apex of submarginal vein.

Male. — Length 1.5 mm. Almost exactly like the female except
that the abdominal gaster in dried specimens is hardly longer
than broad, compressed dorsoventrally and truncate at apex;
petiole very nearly twice as long as broad ; antennae scarcely
distinguishable from those of female, its funicular segments a
little more loosely articulated and the transverse groove setting
off basal segment of club deeper and more distinct. Unlike the
male of insf/Inrii-!, the male of scoli/firora shows no visible ring
segment in the antenna, the funicle is 6-segmented and the club
distinctly 2-segmented.

Redescribed from the type .series compri.sed of nine females
and six males.

368 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 9«

THEOCOLAXIA PITVOPHTHORI (Ashmesd), new combination

Plate 47, Figure 8
CerocephcUa pityophthori (Ashmead MS.) Riley and Howard, Insect Life,

vol. 4, p. 123, 1891.
Cerocephala pityophthori Ashmead, Proc. Ent. Soc. Washington, vol. 3, p.

32, 1894.

Originally described from four specimens now in the National
Museum collection and said to have been reared from Pityoph-
thorus consimilis LeConte taken at Haw Creek, Fla.

Agrees with the description of scolytivora except in the follow-
ing particulars :

Female. — Length 1.4 mm. Frons laterad of scrobes and face
laterad of the depressed area below the antennae more weakly
sculptured, nearly smooth; antenna with the basal segments of
flagellum more slender, a little narrower than pedicel, the first
and second funicular segments subequal, each about as long as
broad and each a little shorter than pedicel ; club about as long as
three preceding segments combined. Anterior and posterior
femora more slender. Marginal vein a little longer than sub-
marginal. Abdomen about as long as thorax, the petiole only
slightly broader at base than at apex, about twice as long as
broad; ovipositor one-third to nearly one-half as long as gaster.
General color reddish testaceous, the vertex, dorsum of thorax,
and whole of gaster piceous and more or less strongly tinged with
a metallic sheen ; legs reddish piceous, with the anterior and pos-
terior coxae whitish except basally ; antenna reddish, the club and
usually the last funicular segment piceous; wings hyaline with a
broad fuscous band embracing the stigmal vein and about the
apical two-fifths of marginal vein, but without a band at apex of
submarginal vein; ovipositor yellowish basally, the apical half
blackish.

Male unknown.

Redescribed from the four females comprising the type series,
and two females from Durham, N. C, said to have been reared
from Thysanoes fimbricornis LeConte, December 13, 1942, by C.
L. Massey.

5. Genus PARALAESTHIA Cameron

Paralaesthia Cameron, Biologia Centrali-Americana, Hymenoptera, vol. 1,
p. 110, pi. 5, fig. 15, 15a, b, c, and f, 1884.

This genus, with P. mandibukiHs Cameron as its only included
species, is unknown to me except by Cameron's description and
figures. It is apparently related to Acerocephala but differs,
according to the description, by having the mandibles three-
fourths as long as the head and bidentate at apex, by having "a

CHALCIDOIDEA RELATED TO CEROCEPHALA — GAHAN 369

wide and deep furrow extending from behind the ocelli to the
antennae and carinated in the middle, the keel running down to
the centre." The marginal vein is said to be longer tiian the sub-
marginal, and there is a tuft of hairs at the junction of the two
veins.

Cameron's rigures of the male and female apparently differ so
widely in characters of the head as to raise a suspicion that they
are not congeneric. The male should be considered the holotype.

Described from Panama.

6. Genus ACEROCEPHALA. new genus

Because of the differently shaped head, greatly elongated man-
dibles, and the absence of a tuft of hairs at the juncture of the
snbmarginal and marginal veins of the forewing, it seems advis-
able to remove Cerocephala atroviolacea Crawford from Cero-
cephala and erect a new genus for it. The new genus apparently
resembles Paralaesthia Cameron, but if Cameron's description is
accurate they may be readily separated by the characters used in
the key. Acerocephala differs further by the absence of any
groove or depression down the middle of the frons.

Female. — Head large, viev^^ed from in front subrectangular.
very slightly broader at mouth than at vertex, its dorsal line
nearly straight, its ventral line (when the mandibles are closed)
squarely truncate, its sides straight and diverging slightly below:
face deeply impressed; lateral margins of facial depression
rounded, without projections ; clypeus not defined ; labrum large,
completely exposed; mandibles large, about one-third as, long as
ihe length of head, squarely truncate at apex and each with four
subequal short teeth ; eyes small, not prominent, obviously much
less than half the length of head; ocelli in a slightly obtuse tri-
angle; ocellocular line approximately equal to distance between
posterior ocelli ; scrobes deep anteriorly, shallower above, not
reaching to anterior ocellus; between the antennae a high, nar-
row, dorsally flattened plate originating some divStaii'.'e l)elow
anterioi- ocellus and either abruptly truncated or gradually
declivous from a point just below the antennal foveae. Antennae
in.serted distinctly below the ventral margins of eyes, 9-segmented :
.scape slender at base, becoming thicker in apical two-thirds, about
four times as long as broad, not compres.sed ; pedicel pyriform.
about one and one-half times as long as broad; funicle 6-seg-
mented. the segments all narrower at base than at apex, subetiual
in length but successively increasing in breadth; first funicular
.segment as long aa or longer than broad and no longer than
pedicel, sixth very slightly broader than long; club ovate, solid.

370 PROCEEDINGS OF THE NATIONAL MUSEUM voL %

with very slight indication of one transverse groove, very slightly
longer than last tw^o funicular segments combined and a little
thicker than preceding segment. Pronotum a little longer than
broad, narroM^ed anteriorly into a distinct neck; mesoscutum
broader and shorter than pronotum, about tvv^ice as broad as long;
parapsidal grooves deep, complete, and not foveolate; scutellum
about as long as mesoscutum, weakly convex ; axillae just meeting;
axillar furrow not or very weakly foveolated; propodeum not
declivous, about half as long as scutellum, without either median
carina or lateral folds and with very shallow spiracular sulci ;
mesopleuron with a very shallow, nonfoveolated, femoral impres-
sion; prepectus moderately large, triangular; anterior and pos-
terior coxae large, subequal ; middle coxae much smaller ; anterior
femora distinctly swollen, as large as or larger than posterior
pair ; posterior tibia with two slender, unequal calcaria. Abdomen
about as long as thorax, somewhat compressed from the sides,
distinctly petiolate; petiole broader at base than at apex, flat-
tened and smooth dorsally; basal segment of gaster deeply, tri-
angularly incised at apical middle; ovipositor sheaths exserted
about one-fifth to one-third the length of gaster. Forewing about
three times as long as broad ; marginal vein longer than sub-
marginal ; stigmal vein approximately twice as long as broad ;
postmarginal weak and not longer than stigmal; discal cilia
vestigial; marginal cilia short; juncture of marginal and sub-
marginal veins with a small but distinct callous but without a
tuft of bristles. Posterior wing about two-thirds as broad and
six-sevenths as long as anterior wing.

Male. — Antenna resembling that of female but 10-segmented,
the club distinctly 2-segmented and with slight indication of a
second cross furrow ; first funicular segment nearl.y as long as
pedicel. Abdomen a little shorter than thorax, compressed dor-
soventrally, subtruncate at apex, the apical segments retracted;
petiole a little broader at base than at apex; basal segment of
gaster not so deeply incised at apical middle as in female. Other-
wise like female.

Type of the genus. — Cerocephala atroviolacea Crawford.

ACEROCEPHALA ATROVIOLACEA (Crawford), new comhinolion

Plate 48, Figure 5

Cerocephala atroviolacea Crawford, Proc. U. S. Nat. Mus., vo!. 45. p. 314,

1913.

Female. — Length about 3 mm. Black, tinged with purple on

head, underside of thorax and abdomen ; thorax above faintly

tinged with green; narrow border around the mouth, mandibles,

CHALCIDOIDEA RELATED TO CEROCEPHALA GAHAN 371

scape, and pedicel rufotestaceous : flag-ellum black and shining;
abdomen with the petiole more or less rufous, the gaster black
slightly diluted with red basally: le.trs concolorous with thorax
except all tarsi and more or less of anterior tibiae reddish tes-
taceous. Wings hyaline, the forewing with a brownish trans-
verse band embracing all of stigmal vein and approximately the
apical two-fifths of marginal vein : venation brownish, the callus
black.

Body mostly smooth and polished ; head mostly smooth but with
scrobal cavity, narrow border along each side of scrobes. and the
sloping inner walls of facial depression weakly aciculated and
ventral surface of head weakly reticulated; raised plate between
antennae polished dorsally, narrowly wedge-shaped, broadest an-
teriorly, and abruptly perpendicularly truncate just below an-
tennae; labrum squarely truncate at apex; scrobal grooves deep
and extending more than half the distance from antennal foveae
to anterior ocellus; neck of pronotum weakly reticulated; meso-
pleura reticulated; propodeum for the most part very finely
transversely lineated, smooth posteriorly; abdominal petiole obvi-
ously a little longer than broad and a little broader at base than
at apex ; gaster entirely smooth and polished, not quite as long as
thorax; ovipositor slightly exserted, never more than one-fifth as
long as gaster.

Male. — Unknown.

Remarks. — Redescribed from the type series and two subse-
quently acquired specimens. The types were reared from a
scolytid infesting cones of pinyon (Piuvs edulis) at Las Vegas,
N. Mex. The two more recently acquired specimens are labeled
as having been reared from Conopkthoncs edulis Hopkins infest-
ing cones of Pinus edulis at Ute Pass, Colo., and are recorded
under Hopkins U. S. No. 9099e.

ACEROCEPHALA AENIGMA. new gpcries

Platk 47, Figures 6, 6a; Plate 48, Figures 4, 4a
The specimens upon which this species is based were at first
believed to represent merely a varietal form of ntroviolacea.
More careful examination of the few specimens available has
shown certain apparently constant characters by which they differ
from ofroviolacrn, however, and makes it appear necessary to
treat them as a separate species.

Female. — Length 2.75 mm. Agreeing with the description of
afroviolacea except in the following particulars: Body shining
black with no suggestion of green on dorsum or elsewhere; pedicel
and apex of scape piceoua; aciculations on front of head stronger

372 PROCEEDINGS OF THE NATIONAL MUSEUM VOL.96

and the sculptured area obviously more extensive, embracing most
of the area betw^een the eyes from a little below the anterior
ocellus to oral margin but smooth along inner orbits ; raised plate
between antennae very faintly aciculated dorsally, narrowly ellip-
tical anteriorly and not perpendicularly truncated but sloping
gradually from just below antennae to oral margin; labrum
broadly rounded anteriorly; scrobal grooves relatively shallow
and extending upward a little less than half the distance from
antennal foveae to anterior ocellus; neck of pronotum a little
more strongly reticulated; petiole of abdomen a little broader
than long and nearly twice as wide at base as at apex ; ovipositor
exserted approximately one-third the length of gaster.

Male. — Length 2.5 mm. Very similar to the female except
antenna somewhat longer and abdomen squarely truncate at apex.
Antenna with 10 distinct segments and with more or less indica-
tion of another division on the club; scape approximately four
times as long as thick, somewhat thickened in apical half; pedicel
longer than thick; funicle 6-segmented, the first segment very
slightly shorter than pedicel, about one and one-half times as long
as broad, following segments subequal to the first in length but
successively increasing slightly in width, the fifth and sixth a
little broader than long; club a little broader than last funicular
segment and a little longer than two preceding segments com-
bined, its first segment closely resembling a funicular segment
but a little less distinctly set off from the following segment.
Abdominal petiole about as broad as long, faintly sculptured and
broader at base than at apex; basal segment of gaster apparently
broadly but not deeply emarginate at apex.

Type Jocalitv. — Prineville, Oreg.

7^7/pe.— U.SlN.M. No. 57280.

Remarks. — Holotype female, allotype, and one female paratype
labeled as having been reared from Pinus ponderosa at Prine-
ville. Oreg., June 2, 1935. by W. J. Buckhorn under Hopkins U. S.
No. 18977D ; one female paratype from Pmus ponderosa, Haca-
more, Calif., June 6, 1931, by K. A. Salman under Hopkins U. S.
No. 20755D; and one female paratype from Phi us coulteri. Mount
Laguna, Calif., November 1940. by D. DeLeon under Hopkins
U. S. No. 32842D.

7. Genus NEOSCIATHERAS Masi

Wcosciathevas Masi, Nov. Zool., vol. 24, pp. 189-192, figs. 49, 50, 1917.

Neosciatheras is unknown to me except by the description. The
genotype, N. laticeps, described in the same paper as the genus, is
based upon a single female collected in the Seychelles Islands.

CHALCIDOIDEA RELATED TO CEROCKPHALA — GAHAN M^

The j^enus ai)pareiitly differs from all others treated here, except
Schtherellns Masi. by having the head and thorax distinctly
sculptured instead of mostly smooth and polished. From Seia-
fherfllii.s, known only in the male sex. it may be distinguished by
the characters used in the key.

The antennae are said to be clavate. inserted below the middle
of face but a little above a line connecting the lower extremities
of the eyes. 10-segmented, without a ring segment, the funicle
6-segmented and the club indistinctly 2-segmented. The head,
viewed from in front, is nearly as broad as long, the eyes promi-
nent, and the cheeks curved. The axillae are rugoso-sulcate. and
the scutellum similarly but less strongly sculptured. The propo-
deum has a deep, semielliptical. transverse excavation basally.
The forewing is without a tuft of hairs at the juncture of marginal
and submarginal veins, the marginal vein is about eight times as
long as the rather short stigmal vein, the stigmal knob is armed
with a short tooth, and the postmarginal vein is a little shorter
than the stigmal. The wing beyond the basal cell is said to be
yellowish gray ("flavido-grisescentes") with the transverse fascia
behind the juncture of marginal and submarginal veins and the
macula adjacent to the stigmal vein obscure, the apical margin
pallid. The abdominal petiole is almost as long as the propodeum,
cylindrical and finely striated, and the gaster is about as long as
thorax, smooth and shining, its first segment not incised at dorsal
margin. The ovipositor is exserted one-third the length of gaster.

8. Genus SCIATHERELLUS Masi

Sciath>'r,'lhiy Masi. Nov. Zool., vol. 24. pp. 189, 192, figs. 51. 52, 1917.

The type species, S. orncinxs Masi. described in the same paper
as the genus, was based on a single male specimen collected in the
Seychelles Islands.

This genus, like Neoscuitheras Masi, apparently differs from
the other genera in this group by having the mesonotum punctate.
It differs from Meosciatheras according to the de.scription by hav-
ing the stigmal vein about one-fourth as long as the marginal.
the stigmal knob only slightly developed and without a process,
the postmarginal vein almost effaced, the antennae inserted a
little above the middle of head, the scape extending above thf
level of vertex, the flagellum very long and not thickened, and the
propodeum without a transverse fovea at base. The forewing is
without a tuft of bristles at the juncture of marginal and sub-
marginal veins and has two transverse fuscous bands. The head,
viewed from in front, is a little broader than long, and the face is
obliquely striate without projections.

374 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

SPECIES WRONGLY PLACED IN CEROCEPHALINAE

One species described in Theocolax is quite certainly not a cero-
cephaline but a eupelmid.

EUPELMELLA CANADENSIS (Provancher), new combination

Theocolax canadensis Provancher, Nat. Canad., vol. 14, p. 35, 1883; Faune

entomologique du Canada, p. 809, 1887.
Cer-ocepkala canadensis (Provancher) Ashmead, Proc. Ent. Soc. Washington,

vol. 3, p. 83, 1894.

No representative of this species has been seen by me but the
description of the thorax cannot apply to any cerocephaline and
does agree quite well with some of the subapterous eupelmids.
0. Peck, of the Canadian Department of Agriculture, informs me
that he once examined the type of the species in the Provancher
collection and concluded it was a eupelmid but did not place it
generically. It is here referred tentatively to EupelmeUa Masi
but may eventually prove to belong in some other subapterous
genus of the Eupelmidae.

Ashmead's translation of Provancher's description is incorrect
with respect to the mesothorax. This is said to be longitudinally
hollowed out, not "longitudinally aciculated" as stated by Ash-
mead.

REMARKS ON SOME GENERA EXCLUDED FROM
CEROCEPHALINAE

The following genera, which at one time or another have been
associated by some authors with Cerocephala in the Spalangiinae,
are considered not to belong in Cerocephalinae :

Paraspalangia Ashmead (Mem. Carnegie Mus., vol. 1, p. 334,
1904) was included in a key to genera of Spalangiinae with a
manuscript species, P. ammlipes Ashmead, named as the genotype.
The type of this species is in the National Museum collection, and
it belongs in the Tetrastichinae. The type specimen was rede-
scribed by Girault (Ann. Ent. Soc. Amer., vol. 9, p. 303, 1916)
under a new generic and specific name, Stigmatotrastichus emer-
soni Girault. The generic name Stigmatotrastichus Girault is a
synonym of Paraspalangia Ashmead, and the specific name
emersoni Girault a synonym of annulipes Ashmead. This syn-
onymy was indicated in the list of type species of the genera of
chalcid flies by Gahan and Pagan (U. S. Nat. Mus. Bull. 123, p.
137, 1923) but without indication of where the genus belonged in
the classification.

Pegoscapus Cameron (Ann. Rep. Estacion Central Agronomica
Cuba, p. 275, 1906), assigned to the Spalangiinae by Cameron and
included in the treatment of that group by Schmiedeknecht (Gen-

CHALCIDOIDEA RELATED TO CEROCEPHALA — GAUAN 375

era insectorum. fasc. 97. pp. 385. 386, 1909), has not been seen
by me, but is certainly a fip: insect (Agaonidae) as stated by
Waterston (Trans. Ent. Soc. London, 1920. p. 129).

Tricorijphu3 Foerster ( Hymenoptera Studien. vol. 2. p. 46,
1856) was originally described without included species. Thom-
son (Hymenoptera Scandinaviae, vol. 4, p. 209. 1875^ redescribed
the genus and included T. fasckdus Thomson which is the geno-
type. Ashmead (Mem. Carnegie Mus.. vol. 1, p. 392, 1904) treated
the genus as a synonym of Cerocephala, but Masi (Ann. Mus. Civ.
Stor. Nat. Genova, ser. 3, vol. 9, p. 240. 1921) declared it to be a
good genus. The U. S. National Museum collection possesses two
female specimens from "Deutschland" identified by Foerster as
Tricoriiphus and bearing an unpublished Foersterian specific
name. These specimens agree with Thomson's description of
Trlconjphus fasciatus. In my opinion, based upon these speci-
mens, the genus Tricoryphus is closely related to Apterolelaps
Ashmead and SpalangioJaelaps Girault and should be referred to
the subfamily Lelapinae.

Spalangiopelta Masi (Ann. Mus. Civ. Stor. Nat. Genova. ser. 3,
vol. 10, p. 30, 1922) is unknown to me except by the description.
Masi states that it is intermediate between Asaphini and Spa-
langiini. The 13-segmented antenna with three distinct ring
segments, the presence of the malar grooves, the unexcavated
face, the subobsolete parapsidal grooves, the presence of a trans-
verse groove on the scutellum, the distinctly sculptured thorax and
the metallic green color seem to exclude it from close relationship
with Cerocephala . I am unable to place it definitely.

EXPLANATION OF PLATES

The drawings for the accompanying plates were made by
Arthur Cushman, of the Bureau of Entomology and Plant Quar-
antine, U. S. Department of Agriculture.

Plate 47

1-la. Choetospila elegans Westwood: 1, Antenna of female; la, antenna
of male.

2-2a. Choetospila tabida, new species: 2, Antenna of female; 2a, antenna
of male.

3-3a. Theocolax fonniciformis Westwood: 3, Antenna of female; 3a, an-
tenna of male.

4-4a. Cerocephala cornigera Westwood: 4, Antenna of female; 4a, antenna
of male.
5. Cerocephala dinoderi Gahan: Antenna of male.

6-6a. Acerocephala aenigma, new genus and species: 6, Antenna of female;
6a, antenna of male.

7-7a. Theocolaxia scolytivora (Ashmead) : 7, Antenna of female; 7a, an-
tenna of male.
8. Theocolaxia pityophthori (Ashmead) : Antenna of female.

Plate 48

1-lb. Theocolax forrniciformis Westwood: 1, Apterous female; la, anterior
and posterior wings from female of al ate form; lb, head of female,
anterior view.

2. Cerocephala aquila (Girault) : Thorax and wings of female.

3. Cerocephala cornigera Westwood: Head of female in semiprofile.
4-4a. Acei-ocephala aenigma, new genus and species: 4, Complete drawing

of female; 4a, anterior view of head of female.
5. Acerocephala atroviolacea (Crawford) : Anterior view of head of
female.

376

•ir U. S. GOVERNYENT PRINTING OFFICE: 1946—714084

U S NATIONAL MUSEUM

Species of Choetospila. Theocolax Cerocephala.

ACEROCEPHALA AND ThEOCOLAXIA
FOR EXPLANATION SEE PAGE 376.

U. S. NATIONAL MUSEUM

SPECIES OP THEOCOLAX. CEROCEPHALA. AND ACEROCEPHALA

FOR EXPLANATION SEE PAGE 376.

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

issued iMilVj^NSlSi ^y ^

SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM

Vol. 96 Waihington: 1946 No. 3204

A REVISION OF THE GENEKA OF MULLETS, FISHES OF
THE FAMILY MUGILIDAE, WITH DESCRIPTIONS OF
THREE NEW GENERA

By Leonard P. Schultz

During my recent studios of the mullets of Venezuela I became
interested in the gonci-ic relationships of the ^fuJ^ilidao and attempted
to define the genera of this group of fishes. To accomplish this I
examined specimens of the mugilid species that have been made
genotypes, basing my diagnoses on specimens in the United States
National Museum. The numerous genera of the family have not
been well defined, and I failed to locate any key or contribution in
which all the genera of the world were compared. This study is a
provisional one. Much more work needs to be done before the
various genera are thoroughly understood, especially in regard to
those centering around Afugil and Chelon as herein defined. My
conclusions were made after several hundred specimens from most
parts of the world were examined; the material used is summarized
under each genus. No attempt is made to place under each gemis
all the species that may belong there, since that task would rccjuirc
the n^cxamination of the types of all described species, scattered in
museums throughout the world.

Several ichtiiyologists have studied the Mugilidae, i)resenting in
keys or diagnoses their understanding of certain genera. Among the
recent ones may be mentioned Jordan and Evermann (U. S. Nat.
Mus. Bull. 47, pt. 1, p. 809, 1896); Oshima (Ann. Carnegie Mus.,
vol. 13, p. 241, 1922); Weber and de Beaufort (The Fishes of the
Indo-Australian Arehipelagr), vol. 4, p. 230, 1922); ^^ohr (Zool.
Jahrb., vol. 54, pp. 195 200, 1927); and Ro.\a>^ (Philippine Journ.
Sci., vol. 54, No. 3, pp. 393-396, 1934). Dr. J. L. B. Smith (Ann.

715201-40- 1 377

378 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 06

South African Miis., vol. 30, pp. 587-589, 1935) has reviewed in an
excellent manner the problems concerning the confusion in regard to
African mugilid species and their diagnoses, but he did not attempt to
solve the problem as to what genera are valid. His discussion of
characters used in describing mullets is worthy of considerable thought,
inasmuch as it applies in general to generic descriptions.

Despite the large amount of material examined, I have experienced
great difficulty in arranguig the species of Mugilidae into genera of
concise and of clear definition, owing mostly to the paucity of useful
taxonomic characters and somewhat to the inadequate descriptions
that abound in the literature. Nevertheless, I have constructed a key
to the genera that is practical, though artificial in itself, and that I
believe defines the natural generic units as observed.

The type of Agonostomus bryanti Bean and Weed, which I have
examined, belongs to the genus Hypseleotris Gill in the family Eleo-
tridae. It is not a mullet.

The family Mugilidae as recognized by authors is remarkably con-
stant in anatomical structures as far as investigated by me ; the num-
ber of vertebrae is usually 24 or 25, with 11 to 13 abdominal and 11 to
13 caudal. The relative positions of the fjns are uniform, and there
is no outstanding example of a great increase in number of fin rays in
any genus. Among other characters the shape of body, with depressed
head anteriorly, spiny preorbital, bi"oad scaly interorbital, large scales
on head and body, two dorsal fins well separated, the first with pecu-
liarly arranged spines, are characteristic of all mullets.

Among the characters studied it appears that the mouth parts of
mugilid fishes have evolved and specialized, whereas the other an-
atomical structures have remained more or less constant for most of
the genera. Therefore, I have studied the preorbital, nostrils, teeth,
jaws, and other mouth parts rather carefully in search of characters
suitable for defining and recognizmg mugilid genera. In most genera
the teeth remain fairly constant in structure at all sizes beyond the
very immature, but in Mugil there is evidence that some of the tips
are simple in the young, becoming bifid or even trifid in very large
adults. It is a well-established fact, also, that in certain species
young mullets have two anal spines, but half grown and adults have
three anal spines. Recent American autliors refer to the immature
stage of the mullet with two anal spmes as the "querimana stage."
The adipose eyelid is another character that develops with age, usually
appearing at standard lengths of 30 to 50 mm. and reaching the highest
development in the adults, or else being absent at all ages.

The key that follows is based on specim.ens of 40 mm. and over, but
one should be able by it to identify mullets to genera at lengths of
half that or shorter. 1 1 is a preliminary step in the definition of mugilid
genera, based on the genotypes and other species referred to the genera
recognized. •

REVISION OF THE GENERA OF MULLETS — SCHULTZ 379

la. Vpiitral side of iiiatidihlc with 4 bluntly rounded fleshy lobes, free posteriorly;
two strips of transverse, close-set, membranous lamellae along lateral edge
of lower jaw on each side externally; mandible bluntly pointed; lower jaw
included, without teeth; upper jaw with a narrow band of villiform teeth;
nostrils close together; no adipose eyelid; upper lij) forming lip of snout;
scales ctenoid (fresh and brackish waters of westiTii tropical Pacific).

Cestraeus V^alencienncs
lb. Mandible without the 4 fleshy lobes and lamellae as described al)ove.

iffi.l pper lip not forming anterior tip of head; snoit fleshy, bluntly rounded,
forming anterior tip of head, snout projecting in front of ui)per lip, the
latter narrow, inferior in position; lower jaw included; no adipose eyelid;
scales ctenoid.
Sa. I'ppt-r jaw with a band of incisorlikc multicuspid teeth inside of thick lips;
each side of lower jaw with a patch of multicuspid teeth widely sepa-
rated at symphysis; nostrils very close together in front of eye; margin
of lower jaw rounded, lower lips thick; no symphyseal knob at middle
of lower jaw and no notch in middle of upper lip; scales cliMioid (fresh

waters of Central America, Mexico, and Cuba) Joturus Poey

oh. Upper and lower jaws with fine teeth in a single row in each jaw, tips
simple, none of the teeth with bifid or nuilticuspid tijjs; nostrils at side
of head at lower level of orbit, widely spaced, the posterior closer to eye
than to anterior nostril; margin of lower jaw angular, not rounded,
with a symphyseal knob and upper lip with pit to receive it; maxillary
and premaxiUary not hooked downward, lower lip thin-edged, teeth in
both jaws eilifjrm, i)Osterior edge of pieorbital very narrow, and an-
terior edge nearly straight, not with a concavity; scali^s ctenoid (fresh

waters, rarely in brackish water, India) . Bhinoraugil Gill

ib. Upper lip forming anterior tip of head and not inferior in position l^eiow a
projecting fleshy snout.
4a. L'pper and lower jaws with thick lips, a distinct wide band of teeth
inside of lips, but no teeth on margin of lips; lower lip not directed or
folded downward; tips of teeth of inner rows on upper jaw bifid or
trifid, those on lower jaw simple or bifid; anterior margin of lower
jaw broadly rounded; maxillary reaching past front of orl)it and past
posterior tip of prcorbital bone; lower lip thick, not thin at edge and
not bearing teeth; no adipose eyelid; nostrils close together; scales
ctenoid (fre;h waters, tropical New World westward to Hawaiian
Islands and Mauritius, New Zealand, and N(!W South Wales).

Agonostomus Pvnnett
4b. Teeth and li|»rf not as in //a.

5a. Lower lip thick-edged, without thin edge and not directed forward

horizontally but folded or directed downward so that lower lip fits

more or Ich.'s snugly Ixthind ui)i)er lip when moutli is closed; margin

of lower jaw angidar.

6a. I-owcr lip bearing HCliform teeth externally on edges, these arranged

i'l a narrow or wide band.

7a. Iloth lipH with broad edges bearing ext( rnully a Ij.'uid of minute

teeth in several clo-se-eet rows, most of teeih having bifid tips;

lower lip directt^d but not folded downwani and without free

inner edge; teeth minute, slender, with bifid tips, arranged in

numerouH clo.HC-set rows on upper pharyngeals; noHtrih very

close together; no adipose eyelid; rear end of prcorbital bone

380 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

several times wider than space between nostrils; scales ctenoid
(marine, in Pacific Ocean of tropical New World).

Chaenomugil Gill
7b. Both lips with narrow edges bearing externally a narrow band of
setiform teeth in 2 or 3 rows; lower lip folded downward and
largely free along its inner edge.
8a. Margin of lips of both jaws with characteristic 3-pointed seti-
form teeth set in 2 or 3 rows; adipose eyelid obsolete or un-
developed; nostrils moderately separated, much closer to-
gether than anterior is from edge of snout, lip excluded;
maxillary not reaching past rear edge of preorbital bone;

scales cycloid (marine, Oceania) Neomyxus Steindachner

8b. Margins of both lips with a uniserial row (occasionally a few in
an outer or second row) of setiform teeth with unbranched
curved tips; adipose eyelid well developed (except nearly
absent in young) in adults, reaching almost to pupil; nostrils,
about as far apart as anterior is from edge of snout, lip ex-
cluded; scales cycloid (marine, Galdpagos Islands).

Xenomugil, new genus
6b. Lower lip bearing several rows of fleshy papillae, both externally and
internally, with their tips crenulate, becoming more so in adults a
foot long or longer; upper lip very broad, bearing papillae, the
crenulate edge continuous around rictus with crenulate lower lip;
nostrils close together, the anterior one as close to premaxillar}'
groove as to posterior nostril; front edge of preorbital with an
angular or concave notch; premaxillary and maxillary bent down-
ward posteriorly but tip of maxillary not exposed as in Chelon;
adipose eyelid undeveloped or absent; scales cycloid (marine,

Oceania and Indian Ocean) Crenimugil, new genus

6b. Lower lip with a thin edge, directed horizontally forward or nearly so,
not folded downward and not bearing teeth externally on lips al-
though fine teeth may occur along edge of lip ; upper lip with a band
of teeth or with a uniserial row of setiform or ciliform, or small
incisorlike teeth, sometimes more or less firmly set.
9a. Adipose eyelid well developed, reaching to or nearly to pupil except
on young 50 mm. in standard length or shorter, in which case
posterior edge of preorbital narrower than space between nostrils;
distance between nostrils wide, equal to or greater than width
of upper lip; anterior and posterior nostrils widely separated,
farther apart than anterior nostril is from groove that separates
upper lip from rest of snout; posterior edge of preobital narrower
(fig. 28, a-c) than distance between nostrils and its posterior tip
scarcely or not reaching past front of eye; maxillary and pre-
maxillary not hooked downward and maxillary not notably
exposed; maxillary and premaxillary in line with front edge of
preorbital; no teeth on vomer or palatines; scales cycloid (marine,
along coasts of Europe, Africa, Asia, North and South America,
West Indies, Oceania to Red Sea, islands of Atlantic and Pacific

Oceans in temperate and tropical waters) Mugil Linnaeus

9b. No adipose eyelid; distance between nostrils narrow, contained
2 or more times in width of upper lip: anterior nostril much
closer to posterior nostril than anterior is from groove that
separates upper lip from rest of snout; width of posterior edge
of preorbital wider than distance between nostrils.

REVISION OF THE GENERA OF MULLETS — SCHULTZ 381

JOa. Teeth setiform or ciliforin, in upjxT lip with simple undivided
tips.
11a. Teeth numerous in upper jaw in a single row, with simple tips
rather coarse and incisorlike, firmly set, not ciliform, and
not forming part of upper lip, but exposed; anterior edge
of preorbital straight or nearly so; maxillary nearly straight,
not notably exposed or hooked downward over the pre-
maxillary (fig. 29, a-c) and not extending past rear edge of
preorbital bone; premaxillary with its anterior margin
evenly curved, the nondontate posterior part in line with
toothed portion, not at .sharp angle to it; narrow band of
teeth on vomer and palatines; scales cycloid (marine and
fresh waters, Australian region and tropical western

Pacific) Myxus Gunther

lib. Teeth in upper jaw ciliform, flexible, extremely fine end
numerous, forming part of upper lip.
18a. Anterior edge of preorbital concave or angular; maxillary
with its posterior part notably exposed, sharply curved
downward over posterior part of premaxillary (fig. 30, a-d)
and extending below preorbital a distance greater than
width between nostrils; premaxillary with its front
margin sharply angular, nondentate posterior portion
hooked backward and downward almost at right angles
to toothed jjortion; vilUform patches of teeth on vomer
and palatines present or absent; scales cycloid (marine,
tropical and temperate Atlantic, Pacific, and Indian
Oceans of Old World, not yet found in New World).

Chelon Rose
ISb. Anterior margin of preorbital evenly curved or nearly
straight (fig. 31, a-c); maxillary with its posterior part
somewhat exposed but in line with posterior part of
premaxillary and not curved downward, but ending
about opposite posterior edge of preorbital; premaxillary
with its front margin evenly curved; a narrow band of
villiform tectli present on vomer and palatines; scales
ctenoid (fresh waters, possibly l)rackish too, Australia,

Burma, and South Africa) Trachystoma Ogilby

10b. Teeth in upper lip setiform rather firmly sot, with trifid or l)ifid
ti[)3; teeth in lower jaw ciliform and embedded more or less
in thin edged lower lip; anterior edge of preorbital concave;
maxillary with its posterior part notably expo.sed, sharply
curvcfl downward over posterior part of jircmaxillary and
extending below preorbital a distance greater than width
between nostrils; premaxillary with its front margin sharply
angular, nondentate posterior portion hooked backward and
downward at a sharp angle; villiform patches of teeth on
vomer and palatines; scales cycloid (marine. South Africa).

Heterotnugil, new genus

In speciilutiii^ on llio rolationsliips uuiou}; [\w. giMioni of Muj^iliduo,
I Hhould assiiino that tlio luost priinitivo inullot hud uiidori^'oiio the
Joast amount of spocialization. Suoli a miiUot should retain largely
unmodified tooth in htuids on jaws, vomer, and palatines. The

382 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

anterior edge of the preorbital bone would be straight, and the
maxillary and premaxillary with nearly straight contour would extend
in line with the straight preorbital edge. The lips would not be
specialized with a thin edge or with modified teeth and papillae.
Among the genera recognized herein, Agonostomus comes nearest to
fitting the above group of characters and may be considered as nearest
the ancestral stock of the family.

The accompanying figure 32 is presented to show in a graphic
manner some of the probable structural relationships common to
certain mugiiid genera. Joturus with its projecting snout and
broadly attached gill membranes may represent the most specialized
genus, more or less in the general line of descent from an Agonosto-
mus-like mullet.

Rhinomugil is an aberrant genus. The nasal openmgs are low on the
side of the head, in line with the lower edge of the orbit. No other
mullet appears to be closely related to this genus.

The remaining genera of mugiiid fishes appear to have two general
lines of specialization, w^hich could have arisen from a mugiiid stock
not greatly unlike Mugil as defined in this prelimmary revision. This
genus has the straight front edge of the preorbital and nearly straight
contour of maxillary and premaxillary in line with front edge of
preorbital. The lower lip is thin but unmodified. The first line of
specialization, as represented by Chelon, Crenimugil, and Heteromugil,
has a concave front edge of preorbital with strongly bent premaxillary
and maxillary bones posteriorly, usually exposed below the preorbital.
The lips and teeth may or may not be specialized. The second line
of specialization represents a group of genera, namely Xenomugil,
Neomyxus, Chaenomugil, and probably Cestraeus, with lower lips
folded downward with higlily specialized teeth and lips. The extreme
specialization in this group may be considered the fresh-water genus
Cestraeus. This genus has the teeth specialized into lamellae on the
external surface of the lower jaw.

All fresh-water genera of mullets have ctenoid scales and all marine
genera of mullets have cycloid scales except Chaenomugil, which has
ctenoid scales.

Genus CESTRAEUS Valenciennes

Cestraeus Valenciennes, in Cuvier and Valenciennes, Histoire naturelle des
poissons, vol. 11, p. 157, pi. 315, 1836 (genotype, Cestraeus plicatil's Valen-
ciennes) (Celebes).

Gonostomyxus MacDonald, Proc. Zool. See. London, 1869, No. 1, p. 39, pi. 1
(genotype, Gonostomyxus loa-loa MacDonald) (Rewa River, Na Vita Levu,
Fiji Islands).

Aeschrichthys Macleay, Proc. Linn. Soc. New South Wales, vol. 8, p. 5, text figs.,
pi., 1883 (genotype, Aeschrichthys goldiei Macleay) (rivers of New Guinea).

REVISION OF THE GENERA OF MILLETS — SCHULTZ 383

Tlie lips, teeth, preorbital bone, and jaws of the MugUidae seem to
have become diliVrentiated more tlian other charaeters. This genus
appears to have had its mouth struetures more complicated and
speeiahzed than any other genus of mullets.

Cesiraeus is characterized by the specialist d lower jjiw ami by Lui;
thick lips on both jaws; the lower jaw is devoid of teeth; externally
there occurs on each side of the mandible along the lateral edge a wide
strip of transverse, very close-set, membranous lamellae; posteriorly
the mandible ends iii four bluntly rounded lobes, free posteriorly;
the posterior end of the upper lip forms another pair of fleshy lobes;
premaxillary with a row of setiform teeth, tips simple; tip of lower jaw
without symphyseal knolj; scales ctenoid; front edge of preorbital
without concavity and scarcely or not serrated; maxillary and pre-
maxillary not bent downward in line with front edge of preorbital,
the former extending far behind the latter; nostrils in lino with upper
edge of orbit, close together, remote from groove behind upper lip;
adipose eyelid absent; apparently no teeth on vomer, palatines or
tongue ; posterior edge of preorbital w4de, serrated; margin of lower jaw
acutely angular; gill membranes extending far forward not broadly
joined across isthums.

The following collections were studied: U. S. N. M. No. 122819
from New Guinea; No. 1372G6 from Bouro Island, Dutch East Indies;
and No. 1:^7207 from the Philippines.

Genus JOTURUS Poey

JoliiTu.-i 1 ur.^. Memorial sobre la historia natural dc la Isla de Cuba, vol. 2,
p. 263, pi. IS, fig.s. 4-5, 1800 (genotype, Jolurus pichordi Poey) (Cuba).

Xenorhynchichthys Rega.n, Ann. Mag. Nat. Ili.st., .ser. 8, vol. 2, p. 461, 1908
(genotype, Xenorhynchichlliys stipes (Jordan and GUhcrt) = J oturvs stipes
Jordan and Gilbert) (Rfo Bayano near Panama).

I have examined the type of Joturus stipes fh)r(hin and Gilbert
(U. S. N. \I. No. ;il010 fiom Panama) and refer it to this genus.

This genus is characterized by the very thick lips in both jaws,
devoid of teeth in the lips, and with the: liesiiy snout projecting be-
yond the lips so that the mouth is somewhat iiiferioi- in position,
nearly horizontal; behind ii])p( r lip on up()er jaw occurs a band of
incisorlike multicusjjid teeth, ea<h side of lower jaw with a patch of
similar multicuspid teeth widely separated at symphysis; no symphys-
eal knob on lower jaw; scales ctenoid; the front edge of the preorbital
is nearly straight without concavity and not serrated; maxillary and
premaxillary not bent downward posteriorly, but in line with front
edge of preorbital; the nostrils are in line with u|)per edge of orbit,
very close together, remote from front of snoiil ; adipose eyelid absent;
te(>th present on vomer, palatines, and on tongue in small patches;
posterior end of preorbital ending in a point vent rally; margin of

384 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

lower jaw rounded; gill membranes somewhat broadly attached across
isthmus.

The following collections were studied : U. S. N. M. Nos. 45532 and
130878 from Mexico; Nos. 78887-9 from Panama; and No. 19915 from
Central America.

Genus RHINOMUGIL Gill

Rhinomvgil Gill, Proc. Acad. Nat. Sci. Philadelphia, 1863, p. 169 (genotype,
Mugil cors7ila Hamilton-Buchanan) (rivers Ganges and Bengal).

Squalomugil Ogilbt, Ann. Queensland Mus., pt. 1, pp. 3, 28, 1908 (genotype,
Mugil nasuius de Vis) (coast of Queensland) .

This genus is characterized by the thin lower lips, du*ected horizon-
tally forward, not curved downward, and supplied with ciliform teeth
embedded in the lip; upper lip with setiform teeth; tips of all teeth
simple; tip of lower jaw with a symphyseal knob; scales ctenoid;
front edge of preorbital nearly straight, v/ithout a conspicuous con-
cavity and with but a few serrations; maxillary and premaxillary
not strongly bent downward posteriorly but in line with front edge
of preorbital; the nostrils are in line with the lower edge of the orbit,
widely separated, and remote from tip of snout; adipose eyelid absent;
vomer, palatines, and tongue probably toothless; posterior end of
preorbital very narrow, ending in two or three spines; margin of lower
jaw angular; gill membranes extending far forward, not joined across
isthmus.

The diagnosis is based mostly on plate 9, figure 97, of Hamilton-
Buchanan, and on Hora, Journ. Bombay Nat. Hist. Soc, vol. 40,
No. 1, pp. 62-68, pi. and 3 text figs, of Mugil corsula, 1938; also on
Whitley's (Australian Zool., vol. 10, No. 1, p. 22, fig. 16, 1941) account
and description of Mugil nasutus, along with one specimen of R.
corsula, U. S. N. M. No. 44767 from Kangoon.

Genus AGONOSTOMUS Bennett

Agonoslomus Bennett, Proc. Comm. Sci. Corresp. Zool. Soc. London, No. 14,

p. 166, 1832 (genotype, Agonostomus telfairii Bennett) (Mauritius).
Nesiis Valenciennes, in Cuvier and Valenciennes, Histoire na^turelle des poissons,

vol. 11, p. 167, pi. 317, 1836 (genotype, Nestis cyprinoides Valenciennes)

(lie de France ; Bourbon) .
Dajaus Valenciennes, in Cuvier and Valenciennes, ibid., p. 164, pi. 316 (geno-

tj'pe, Dajaus monticola Valenciennes =Mi<5ft7 monticola Griffith) (rivers of

Dominican Republic, Puerto Rico, and Jamaica) .
Neomugil Vaillant, Bull. Soc. Philom. Paris, ser. 3, vol. 6, p. 72,1894 (genotype,

Mugil digued Vaillant ==MMgrz7 monticola Griffith) (Lower California).

Agonostomus hryanti Bean and Weed (U. S. N. M. No. 72582) be-
longs to the Eleotridae, and probably in the genus Hypseleotris Gill.
It is not a mullet.

This genus is characterized by the thick lips in both jaws and by a
wide band of small teeth in both jaws; the outer rows of the upper

REVISION OF THE GENERA OF MULLETS — SCHULTZ 385

jaw have simple tips, but the inner rows of teeth have bifid or trifid
tips, at least in the atlults; the teeth in the lower jaw usually have
simple tips, but occasionally some have bifid tips; no symphyseal
knob on lower jaw: scales ctenoid: front edge of preorbital. serrated
but without conspicuous concavity, nearly straight; maxillary and
promaxillary not bent downward posteriorly but mostly in line with
front edge of preorbital; nostrils in line with upper edge of orbit, very
close together, remote from groove behind upper lip; adipose eyelid
absent; villiform teeth on vomer and palatines but apparently absent
on tongue; posterior edge preorbital very wide; margin of lower jaw
rounded; gill membranes extending far forward not broadly connected
across isthmus.

The generic diagnosis is based on the description by Fontaine en-
titled, "Sur La Chitte (Agonostoma ieljairii Glinther)" in Bull. vSoc.
Zool. France, vol. 53, pp. 38G-390, figs. 1-4, 1928, and on collections
in the National Museum, too numerous to list the numbers here,
but from Guadaloupe Islands, Secas Island, Mexico, Panama, Vene-
zuela, and Vsesi Indies (largely from Old Providence Island, Jamaica,
Cuba, Puerto Rico, and Haiti). There are a few specimens from New
Zealand and New South "Wales and one from the Hawaiian Islands.

Genus CHAENOMUGIL Gill

Chacnomugil Gill, Proc. Acad. Nat. Sci. Philadelphia, 18G3, p. 169 (genotype,
Miigil proboscidens Giinther) (west coast of Central America).

This genus is characterized by the broad, thick, pliable lips bearing
externally a band of small teeth with bifid tips in several close-set
rows on both upper and lower jaws; lower lip directed downward and
fitting snugly behind the upper li[) when mouth is closed; tip of lower
jaw more or less bluntly pointed but symphyseal knob undeveloped;
scales ctenoid; front edge of preorbital flexible without concavity,
serrated only posteriorly; maxillary and premaxillary not bent down-
ward posteriorly; nostrils in line with uppr edge of orbit, very close
together, remote from groove behind broad upper lip; adipose eyehd
absent; no teeth on vomer or palatines but some on tongue; posterior
edge of preorbital wide, serrated ; margin of lower jaw acutely angular;
gill membranes extending far forward, not broadly connected across
isthmus.

I have studicfl the following collections of ('. ])roboscidens: U. S.
N. M. Nos. 465G3-4 and 125343 from Clarion Island; Nos. 54541,
56343, 07578, and 1070.-,n from Socorro Ishuid ; No. G5440 from Chat-
ham Island; No. f3544S from Culehra Islnnd; No. 128504 from tho
Pearl Islands; No. 47471 from MazatlAn; No. 101045 from Cupica
Bay, Colonihia;and Nos. 79778-81 , 797S0, 79829, 79830, and 1 2857 1-2
from Panama. All records from the Pacific.

715201-40-2

386 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Genus NEOMYXUS Steindachner

Neomyxus Steindachner, Ichthyologische Beitrage (VII), Sitz. Akad. Wiss.
Wien, vol. 78, p. 384, 1878 (genotype, Myxus {Neomyxus) sclateri Steindach-
ner) (Kingsmill and Sandwich Islands).

This genus is characterized by the thick phable hps bearing exter-
nally two or three rows of characteristically shaped, 3-pointed or
trifid setiform teeth on both jaws; lower lip with teeth and folded
downward and fitting snugly behind the upper lip when mouth is
closed; tip of lower jaw bluntly pointed with symphyseal knob small
and located internally ; scales cycloid ; front edge of preorbital straight
without convacity, serrated anteriorly; maxillary and premaxillary
not bent downward posteriorly; nostrils in line with upper edge of
orbit, moderately close together, the anterior nostril remote from
groove behind upper lip ; adipose eyelid absent ; no teeth on vomer or
palatines but probably some on the tongue; posterior edge of preor-
bital wide, and serrated; margin of lower jaw acutely angular; gill
membranes extending far forward, not broadly connected across the
isthmus.

I have studied the followmg collections in the United States Na-
tional Museum: Niuafau Island, U. S. N. M. Nos. 91854-6, 91858-9,
and 91983; Phoenix Islands, Nos. 115629 and 115632; Swains Island,
Nos. 115627-8 and 115631; Wake Island, No. 82893; Makemo Island,
No. 65969; Baker Island, No. 88153; Guam Island, No. 65968; Ellis
Island, No. 65967; Tongareva Island, No. 88152; Manga Riva Island,
No. 65596; Marquesas Islands, Nos. 89744-5; Hawaiian Islands,
Nos. 52772, 55433, 55434, 55439, 55475, 55488, 55525, 82867, 89532,
115630, and 126540. These specimens all came from the central
tropical Pacific Ocean or Oceania.

XENOMUGIL, new genus

Genotype. — Mugil thoburvi Jordan and Starks, in Jordan and Ever-
mann, U. S. Nat. Mus. Bull. 47, pt. 1, p. 812, 1896 (types, U. S. N. M.
No. 47576, Galapagos Islands).

This genus is characterized by thick lips in both jaws, each bearing,
externally, a uniserial row or in two rows anteriorly of setiform teeth
with unbranched tips; lower lip folded downward and fitting snugly
behind the upper lip when mouth is closed, tip of lower jaw bluntly
pointed, with a small symphyseal laiob somewhat developed behind
tip of jaw; scales cycloid; front edge of preorbital straight without
concavity and serrated; maxillary and premaxillary not bent down-
ward posteriorly; nostrils in line with upper edge of orbit widely
separated, the anterior one as close to groove behind upper lip as to
posterior nostril; adipose eyelid present, well developed on adult; no
teeth on vomer or palatines but probably a few on the tongue; pos-

REVISION OF THE GENERA OF MULLETS — SCHULTZ 387

teiior edge of preorbital rather narrow, serrated; margin of lower jaw
acutely angular; gill mtunbrancs extending far forward not broadly
connected across thi; istiunus.

^». I have examined the following collections, all taken from the
Galdpagos Islands: U. S. N. M. Nos. 41881, 41459, 50017, 05597,
8974S. The two types bear U. S. N. M. No. 4757G.

This new genus may be distinguished from all other genera in the
family Mugilidae by the key on pages 379 to 381.

Named Xenoniugil in reference to the "strange" teeth and lips.

CRENIMUGIL. new genus

Genotype. — Mugil crenilabis F6rskal.

Under Chelon, Oshima (Ann. Carnegie Mas., vol. 13, p. 257, pi.
13, fig. 1, 1922) lists a single species, ''Chelon crenilabis (Foi^kal),"
from the Pescadores Islands, west of Formosa, but since this species
belongs in a genus distinct from Chelan. Rose, with Mugil chela as
the tvpe, it appeai-s necessary to propose a new generic name for this
type of mullet. It is diagnosed below.

This genus is characterized by the thick lips bearing papillae ex-
ternally and internally; those forming the row along the external
edge of the Hps are crenulate, becoming more so in the adults and
even continuous around the corners of the mouth; the thick lower
lip is somewhat folded outward; the papillae apparently represent
"teeth"; inside of upper jaw near front of mouth are plicate fleshy
folds: symphyseal knob at tip of lower jaw; scales cycloid; front edge
of preorbital with concave notch to receive corner of mouth; maxillary
and premaxillary moderately bent downward, but the maxillary not
e.xpo.sed as in Chelan; nostrils on level of upper edge of orbit, moder-
ately separated, so that the anterior nostril is as close to groove
behind upper lip as to posterior nostril, the anterior nostril being
rather close to the groove; no adipose eyelid; no teeth on vomer but
teeth on palatines and a few on tongue; posterior edge of preorbital
very wide, wider than space between nostrils; margin of lower jaw
broadly angular; gill nu'inbranes extending far forward, not broadly
comiccted across the isthmus.

I have examined the following collections bearing U. S. N. M.
numbers: Indian Ocean, Xos. 44522 and 44557; Christmas Island,
No. 19218; Phoenix Islands, Nos. 115040-2; Samoan Islands, No.
115039; Tahiti, No. 87049; Marshall Islands, No. 65912; Ciuara, No.
05913.

This ncnv genus may b(^ distinguished from all olher l'-mm ra in tlic
family Mugilidae by the key on pages 379 to 381.

Named Crenimugil in reference t/> the crenul'ite lips.

388

PROCEEDINGS OF THE NATIONAL MUSEUM

Genus MUGIL Linnaeus

Figure 28, a-c

Mugil Linnaeus, Systema naturae, ed. 10, vol. 1, p. 316, 1758 (genotype, Mugil

cephalus Linnaeus) (European Ocean).
Cephalus Lacepede, Histoire naturelle des poissons, vol. 2, p. 589, 1800, new

name on Plumier MS. (genotype, Mugil cephalus Linnaeus).
Arnion Gistel, Naturgeschichte des Thierreichs, p. x, 1848, substitute name for

Mugil (genotype, Mugil cephalus Linnaeus).
Ello Gistel, Handbuch der Naturgeschichte fiir alle Stande, p. 356, ]850[=1847],

and Naturgeschichte des Thierreichs, p. 109, 1848; according to Whitley,

Austral. Zoo!., vol. 6, pt. 3, p. 251, 1930, a synonym of Mugil Linnaeus.
Querimana Jordan and Gilbert, Proc. U. S. Nat. Mus., vol. 5, p. 588, 1883

(genotype, Myxus harengus Giinther) (Pacific coast of Central America).

The diagnosis of Mugil is based on descriptions by Cuvier and
Valenciennes (Histoire naturelle des poissons, vol. 11, p. 19, pi. 307,
1836, Mediterranean Sea) and Giinther (Catalogue of the fishes in
the British Museum, vol. 3, p. 417, 1861, Mediterranean Sea), on

Figure 28. — Sketch of the maxillary, premaxillary, and preorbital bones of Mugil cephalus
(U. S. N. M. No. 45009 from Greece); a, Maxillary lying over premaxillary; b, view of
ventral contour of maxillary; c, preorbital.

U.S.N.M. Nos. 45009 and 84585 from the Mediterranean Sea, and
on numerous other specimens referable to one or more species from
many localities thi'oughout the seas of the world.

I have examined the five types of Mugil cetosus Gilbert (U.S.N.M.
Nos. 46554 and 48254) and a paratype (No. 124990) from Clarion
Island and refer them to this genus. Eight small types of Querimana
gyrans Jordan and Gilbert (U.S.N.M. No. 34966) belong to this
genus.

REVJ.SIOX OF THE GENERA OF MULLETS — SCHULTZ 389

This goiuis is cluiractorizod by a thin-otl<2:o(l lower lip projecting
fonvard horizontally, not curved or folded downward; the teeth in the
lower lip are setiform or ciliform, partly embedded or conspicuous;
teeth in upper lip similar. The outer row of teeth in both lips is
usually more piominent, with simple tips, and if inner rows occur these
are either bifid or trifitl, at least on adults (apparently the teeth in
certain species of this genus become bifid or even trifid in large adults);
a symphyseal knob present at tip of lower jaw; scales cycloid; the
preorbital has the front edge straight or nearly so without a con-
spicuous concavity; maxillary and premaxillary not bent downward
posteriorly, but in line with the front edge of preorbital; nostrils in
line with the upper edge of the orbit, widely spaced, the anterior is
closer to groove behind upper lip than to posterior nostril; adipose
e^yelid present; upi)er lip usually not so wide as distance between
nostrils; teeth probably absent on vomer, palatines, and tongue;
posterior edge of preorbital narrower than space between nostrils;
margin of lower jaw angular; gill membranes extending far forward,
not broadly connected across isthmus.

In this genus I find that in small specimens of certain species the
teeth have simple tips, but later the inner teeth have bifid tips and in
the largest adults some possess trifid tips. The teeth of the outer row
usually have simple tips, but in some large specimens these are bifid
too.

I have examined too many collections of the numerous species
referable to this genus to list them here. The localities represented
are: Europe, both coasts of Africa, both coasts of North and South
America, West Indies, Hawaiian Islands, Australia, Oceania, Japan,
coast of Asia, in temperate and tropical seas.

Genus MYXUS Giinther

FiauRK 29, a-c

Myxua GOnther, Catalogue of the fishes in the British Museum, vol. 3, pp. 409,
466, 1801 (genotype, Mugil elongatus Giinther) (Hobsons Bay and Port
Jackson, Australia).

This genus is characterized by a moderately thin lower lip i)rojecting
horizontally forward, with a single row of small, close-set, incisorliko
t<'eth rather finnly set; teeth in upper jaw similar and in one row; all
teeth with simple tips; no iiuier rows of teeth in either jaw; a.symphy.s-
eal knob at inside tip of lower jaw; scales cyoloid; front edge of pre-
orbital straight, no concave notch ; maxillary and premaxillary not bent
downward but nearly straight, in line with front edge of preoibital;
maxillary not exposed posteriorly; nostrils on level of upper edge of
orbit rather close together, the anterior much farthcT from groove
behind upper lip than from posterior nostril, no adipose e^'elid;
narrow band of teeth on vomer and palatines, and probably on tongue;

390

PROCEEDINGS OF THE NATIONAL MUSEUM

posterior edge of preorbital very wide, much wider than space between
nostrils; margin of lower jaw rounded; gill membranes extending far
forward, not broadly connected across isthmus.

The diagnosis of this genus was based on M. elongatus, U. S. N. M.
Nos. 47770 and 47773 from Lord Howe Island and Nos. 59889, 59913,
and 83052 from New South Wales.

Figure 29. — Sketches of the maxillary, premaxillary, and preorbital bones of Myxus
elongatus (U. S. N. M. No. 59912 from New South Wales): a, Maxillary lying over pre-
maxillary (a tooth is shown below at right hand side); b, view of ventral contour of
maxillary; c, preorbital.

Figure 30. — Sketches of the maxillary, premaxillary, and preorbital bones of Ckelon chelo
(U. S. N. M. No. 123002 from Europe): a, Maxillary lying over premaxillary; b, view of
ventral contour of maxillary; c, preorbital; d, posterior tips of maxillary and premaxillary.

REVISION OF THE GENERA OF MILLETS SCHULTZ 391

Genus CHELON Riise
FiGi'RE 30, a-d

Chelon Ro.sE, Petri Artedi Ant;erniania — Sueci synoiiyinia nominiini piscinm
.... ed. 2, p. IIS, 1793. — Jordan' and Evkrmann, Genera of fishes, pt. 1,
p. 52, 1917 (genotype, Mugil chelo Cuvier and Valenciennes).

Liza Jordan and Swain, Proc. U. S. Nat. Mus., vol. 7, pp. 261, 262, 1884 (geno-
type, Mugil capita Cuvier) (Mediterranean and seas of Europe).

Oedalechiius Fowlkr, Proc. Acad. Nat. Sci. Philadelphia, vol. 55, p. 748, 1903
(genotype, Mugil lahco Cuvier) (Mediterranean Sea).

EUochelon Whitley, .Australian Zool., vol. 6, pt. 3, p. 251, 1930 (genotype, Mugil
vaigiensis Quoy and Gaimanl) (Waigiou).

The genus Oedalechiius Fowlor, with Afugil labeo Cuvior as its typo,
appoars to roprcsent the most e.xtreme development of the mouth of
the genus Chelon. In M. labeo the upper Hp is nearly as broad as the
orbit and the maxillary bone extends vertically downward below the
preorbital bone where its posterior end is e.xposed. There are other
species from the Mediterranean in which the upper lip is not so broad
and the mouth is more horizontal with the maxillary similarly exposed.

Chelon Rose (loc. cit.) was listed bj^ Jordan and Evermann (loc.
cit.) as a valid genus, with "Chelon of Gesner, which is probably
Mugil chelo Cuvier and Valenciennes" as its type. Although the
generic name is given as Chelon by Rose, no species are listed in the
binomial sense, and no description of any kind is given, yet the genera
listed in the 1793 edition are definitely used in the binary sense, in-
cluding Chelon. Common names of pre-Linnaean authors with ref-
erences are cited in the synonymy, as for example: "Labeo. Gaz.
Arist. 1. c." and "ciielo. Rondel. 1. 9 c. 5, p. 266. gesner. p. 552." The
mullet "Chelo" or "Chelon" of the Mediterranean Sea is such a well-
known species that Giinther (Catalogue of the fishes in the British
Museum, vol. 3, p. 4o4, 1861) in the sj^nonymy under Mugil chelo
lists without question the common name "Chelon" as used by
"Rondel.," "Gesner," "Willughby," and "Ray" but does not mention
Rose. The first desciiption and binomial use of Mugil chelo appoars
to 1)0 that \)y Cuvior (in his Rogue iVriiniul).

Oshima (Aim. Canicgio Mus., vol. 13, pp. 241, 257, pi. i:!. (ig. 1,
1922) has recognized the genus Chelon Rose, 179;}, listing but one
species, Ch^'lon crenilahis (F^jrskal) under tiio genus, but this species
hjus most elaborately developed papillat^^ lips and ollioruise is nol a
Chrlon but belongs in a new genus heiein i)rop()s<'(l on page 3S7.

I have exuininod the small typo of Agonostimtus (lorsdlls Streets
from the Sumoan Islands (U. S. N. M. No. l.")ill) and refer it to
this genus. Two puratypes of Afugil canalin/lotus ,J. L. B. Smith
(IJ. S. N. M. No. [):'A\47) from Durban, South Africa, also belong to
this genus.

392 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

The genus Chelon Rose is characterized by the thiii-edged lower
lip projecting forward horizontally, not cui'ved or folded downward;
the lower lip is made up of ciliform teeth embedded in the lip, none
of which project beyond the flesh of the lip; in the upper lip the
setiform teeth project beyond the lip; tips of all teeth simple, none
bifid or trifid even in large adults ; behind outer row of setiform teeth
in upper lip occm* minute teeth in one or more rows, all with simple
tips; a symphyseal knob at tip of lower jaw; scales cycloid; the pre-
orbital has a concave notch or shallow concavity in its front margin
to accommodate the exposed maxillary, which is hooked or bent down-
ward ; the premaxillary is also bent posteriorly at a more or less sharp
angle, the posterior part of this bone lying behind the maxillary;
the nostrils are on the level of the upper edge of the orbit and are
close together, usually closer to each other than anterior is from
groove behind upper lip; no adipose eyelid present; upper lip wider
than distance between nostrils; teeth on vomer, palatines, and tongue
present or absent; the posterior edge of the preorbital is wider than
the space between the nostrils; margin of lower jaw angular; gill
membranes extending far forward, not broadly connected across
isthmus.

The collections referred to this genus are too numerous in the Na-
tional Museum to list here, but they came from the following localities
in the Pacific: Marquesas Islands, PhoenLx Islands, New Hebrides,
Sumatra, Java, Roual, Samoan Islands, Tahiti, New Guinea, Christ-
mas Island, Marshall Islands, Solomon Islands, Celebes, and Phil-
ippine Islands, China coast, Japan, Korea, Peter the Great Bay, and
New South Wales, Australia. In the Atlantic: Europe, British Isles,
Norway, Azores, and Canjiry Islands. Other localities: Mediter-
ranean Sea, Island of Mauritius, Burma and India, British East
Africa, British South Africa, and French Congo, Africa.

I did not find any specimen from the New World that was referable
to this apparently Old World genus.

Genus TRACHYSTOMA Ogilby

Figure 31, a-c

Trachysloma Ogilby, Proc. Zool. Soc, London, 1887, p. 614 (genotype, Trachy-
sioma multidens Ogilby) (Port Stevens at mouth of Keruah River) ( = Mtigil
breviceps Steindachner=MwgfiZ petardi Castelnau).

Sicamugil Fowler, Not. Nat., No. 17, p. 9, fig. 1, 1939 (genotype, Mugil hamiltoni
Day) (fresh waters of Burma).

Gracilimugil Whitley, Australian Zool., vol. 10, pt. 1, p. 19, fig. 14, 1941 (geno-
type, Grac.ilimugil ramsayi (Macleay) = Mwgri7 ramsayi Macleay, 1883)
(Burdekin River, Queensland).

Although I have not seen a specimen of Gracilimugil ramsayi
(Macleay) I am unable to find any statement in Whitley's diagnosis
that definitely separates it from Trachy stoma.

HEVISIOX OF THE GENEKA OF MULLETS — SCHULTZ

393

This e:omis is chnniclorizotl l)y haviii!> ciliform teoth omhoddcd in
the lips of both jaws, tlicir tips not or scarcely visible; the lower lip
is thin as iii Chelon and directed forward horizontally; tip of lower jaw
v^Tth s^'Hipliyseal knob; scales ctenoid; front edjje of preorbital nearly
strai<2:ht, without conspicuous concave notcli, and its posterior edge
with or without greatly enlarged spines; maxillarj'^ and premaxillary
not bent downward but extending in nearly same line as front edge of
preorbital except when ihe spines project as in hamiltoni Day; nostrils
in line with upper edge of orbit, somewhat separated, closer together
than anterior is from edpe of groove behind ui)per lip or as far apart as
anterior is from edge of groove behind upper lip; adipose eyelid
wholly absent; a narrow band of teeth on vomer and palatines and

Figure 3L — Sketches of the maxillary, premaxillary, and preorbital bones of Trachystoma
petardi (U. S. N. M. No. 59866 from New South Wales): a. Maxillary lyin^' over pre-
maxillary; b, view of ventral contour of maxillary; c, preorbital.

pioljably on tongue; nuirgin of lower jnw rounded to soni<>what
angular; gill membranes extending fur ftMuaid. not broadl3' connected
across isthmus; anal origin notably in front of second dorsal origin.
The generic dingnosis was based on three specimens of Trnchystnma
petardi (U. S. \. M. .\o. 5986G) from Clarenee River, New South
Wales. Also I refer to this genus a specimen from Durban, South
Africa, sent to the Xationnl Museum many years ago under th<> name
" MufjU euronotvfi^' by the Alb:ui\ Museum. This specimen ngrees
with Dr. J. L. B. Smith's description (Ann. South African Mus., vol.
30, pp. 010, 013, fig. 7, pi. 10, E, H).'}.'.). which slates that it occurs
almost wholly in fresh water and only mrely in the sea. 1 have
studied a small specimen of Afugil hnmiUoni Day (U. S. N. M. No.

394 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

44795) from Rangoon that I refer to this genus. From petardi it
differs in the greater development of the preorbital spines.

HETEROMUGIL, new genus

Genotype. — Mugil tricuspidens J. L. B. Smith.

This new genus of Miigihdae is a Chelon with trifid and rarely bifid
teeth in the upper lip in a single row, as observed on specimens 60 mm.
and longer. These teeth are setiform, incisorlike, and close-set.
The lower lip is moderately thin with embedded ciliform teeth. It
has the maxillary exposed beyond and belov/ the preorbital; the
pre maxillary is sharply angular, extending behind the maxillary bone
which turns downward; the upper lip is thick and wide, wider than the
distance between nostrils ; the posterior edge of the preorbital is wider
than the space between nostrils, and the anterior nostril is much closer
to the posterior one than to the groove separating upper lip and snout ;
no adipose eyelid is developed ; there are patches of villiform teeth on
the pterygoids, vomer, and tongue; other characters are those of the
genotype Mugil tricuspidens.

This new genus differs from all other genera of Alugilidae as dis-
tinguished in the key, pages 379 to 381.

I have examined one specimen, a paiatype of Mugil tricuspidens
J. L. B. Smith (U. S. N. M. No. 93651) from Mazeppa Bay, South
Africa. Specimens shorter than 60 mm. have not been collected
according to Dr. J. L. B. Smith.

Named Heteromugil in reference to the distinguishing (i. e., different)
teeth, with trifid or bifid tips.

REVISION OF THE GENERA OF MULLETS SCHULTZ 395

HETEHOilUJIL
(M Cy Po Ai< Lt)

CESTRAEUS
(F Ct P3 Aa Lf )

CHENIiu'UOlL
(M Cy ?c A" L:

UYXTJS
(M Cy P3 A.i Lt)

TSAOHTdTOMA

(? ct Ps fA I',)

NE0!.rEaJ3
(M Cy P3 Aa Lf)

JCTURUS
(F Ct P3 Aa Lw)

UDGILID STOCK
Figure 32. — Diagram of the possible relationships of genera of the Mugilidae. Letters
in the parentheses under each genus indicate some of the characters used in forming an
opinion as to the general evolutionary trends. (Aa. — Adipose eyelid absent; Ap. — Adi-
pose eyelid present; Ct. — Ctenoid scales; Cy. — Cycloid scales; F. — Fresh-water habitat;
Lf. — Lower lip folded downward; Lt. — Lower lip with thin edge projecting forward;
Lw. — Lower lip thickish; M. — Marine habitat and entering brackish waters; Pc. — Front
edge of prcorbital straight and the maxillary and premaxiilary extending in the same
general line as front edge of preorbital; Ps. — Front edge of preorbital concave or deeply
notched, the maxillary and premaxiilary bent at an abrupt angle posteriorly, and exposed
below prcorbital.)

U t- COVCHNIItNT PRIMTINS OfFICIi II4«

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

isjueJ l^tiTviL Sll^i by the

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM

Vol. 96 Wwhington: 1947 No. 3205

THE rHORID FLIES OF GUAM

By G. E. BoHART

The primary purpose of this paper is to make available the names
of several species of Phoridae (Diptera) whose biology and life history
will be di!^cu.ssod in a paper (to be published by the Navy) on the filth-
inluibiting flics of Guam. In it will :ippear biolog^ical notes and de-
scriptions and illustrations of the early stages of many of the phorids
herein described.

Except for those species reared from filthy environments no par-
ticular effort was made to collect phorids. Consequently, several of
the species described are represented by unique specimens, and it may
be inferred that numerous species remain to be discovered on the island.
Because of the paucity of material I felt it advisable to illustrate the
species as fully as possible, so that if some of these described as new
prove to be already named, or if the discovery of many more S|)ecic3
reduces the value of the key, the figures will still prove useful. In
most cases, in view of the full illustrations, descriptions have been kept
at a minimum size.

The lii'ld work in connection with this study was done on Guam at
the field laboratories of Naval Medical R'search Unit No. 2 under the
diifction of Conuiiodore Thomas J. Rivers. The taxonomic work and
the preparation of the paper were done at the U. S. National Museum
under the auspices of the Research Division of the Bureau of Medicine
and Surgery. I am much indebted to Lt. (jg) J. L. Gressitt, who
worked with mo on Guam, for some of tlic rearing and collecting of
specimens and for carrying on the work after I left the island. The
men at the National MuHCum have done their usual utmost to provide
facilities and as.sist me whenever necessary.

716067—47 1 397

398 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 86

Phorids, in relation to the insect fauna as a whole, are prominent on
Guam both as to species and individuals. The cosmopolitan Megaselia
scalaris (Loew) was abundant in all inhabited areas and bred freely in
such diverse materials as green cornstalks, rotting coconuts, carrion,
and human excrement. It was almost impossible to keep it from con-
taminating our cultures of other flies, and it bred freely in fresh stools
under examination for intestinal parasites by the parasitology labora-
tory. The highly degenerate Ghonocephalus subglaber had the habit
of swarming on decaying fruit by thousands, and it could nearly always
be taken in decaying wood. Collections of rotting shells buried just
beneath the ground surface furnished breeding material for six species
of phorids, including two of the wingless genus Puliciphora.

The figures are from drawings by the author. For subjects I used
specimens in dilute alcohol after softening (but not decolorizing) in
KOH. The genitalia were drawn from dissected specimens in alcohol
and checked again for accuracy of details with the same specimens
mounted in euparol.

The holotypes and allotypes of the new species are in the collection
of the U. S. National Museum. Paratypes will be deposited in the
California Academy of Sciences, the collection of the Hawaiian Sugar
Planters' Association, and the author's collection.

Genus MEGASELIA Rondani

MEGASELIA (MEGASELIA) SETIFEMUR, new species

Figure 38

Holotype, male. — Length 1.2 mm.; length of wing 1 mm.; frons,
dorsum of thorax, abdomen, and halteres uniformly piceous-brown ;
pleura, venter of abdomen, and legs pale testaceous except for darker
apices of hind fermoa. Head: Frontal bristles thick but not long
(not more than two-thirds as long as width of frons) ; antiales and
lower frontals close together and both lower than upper supra-anten-
nals, which are about twice as long and far apart as the lower supra-
antennals; cheeks with two strong bristles at lower angle and two
moderate ones at level of antennal insertion ; frontal hair rather sparse
and long. Thorax: Dorsal hairs short but strong, becoming sparse
ap.d bristly toward scutellum; posterior edge of scutum with four
bristles, the outer ones about one-half longer than the irnier ; scutellum
with a strong inner, apical pair of bristles about as strong as the outer
scutal ones, and a very weak outer basal pair less than half as long as
the inner ones ; pleura bare except for three bristles on ventral margin
of propleuron ; anterior spiracles included in separate sclerites. Legs :
Of moderate length, the posterior tibiae and tarsi about as long as
wings; posterior tibiae with a single dorsal row of setae inside the
dorsal hair fringe ; posterior metatarsi with two inner bristles some-

THE PHORID FLIES OF GUAM — BOHART 399

what loiijier than the others along the same row; ventral margin of
posterior femora with about four basal haii-s, much longer than any
toward the apex. Wings: ^Membrane lightly infuscated, costal bris-
tles long and sparse"; wings otherwise bare except for a few bristles
basally on posterior margin; veins placed as shown in figure 38.
Abdomen: Tergites ahnost uniformly brown, slightly paler basally
and medially and very sparsely pubescent, the hairs mostly in patches
laterally and along posterior margins; genitalia small, the apical
lamella divided into two parts, the upper part with scattered hairs as
long as the pair of terminal bristles on the lower part.

Holotype (unique) (U. S. N. M. No. 57990) : Pago River Valley,
Guam, June 1945, swept from dense jungle vegetation on a steep si )pe
(G. E. Bohart).

Remarks. — This species is similar in coloration and costal fringe of
the wing to paraha>iiseta but lacks mesopleural bristles and any
bristles on the third longitudinal vein. It runs close to equiseta
Brues and luiisctosa Brues in his key to the Philippine species (1936)
but differs from the former in having lighter color, longer and sparser
costal :wing fringe, and shorter frontal bristles. It diflfers from
nnisttosa in lacking a bristle on the base of the radial sector vein and
having four bristles on the posterior scutal margin. It runs to couplet
55 in Malloch's key to the Megaselia in the U. S. National Museum
(1912) but differs from either conspicualis Malloch or inornata Mal-
loch by having a normal costa and brown halteres. The male geni-
talia and basal ventral hairs of the hind femora will probably serve to
characterize this species.

MEGASELIA SUIS, new Hpeciei
i'^OUBE 36

Uolotype^ male. — Length 1.5 mm.; length of wing 1.4 mm.; body
yellow; frons somewhat darkened above; scutum testaceous; abdomen
yellow, with fourth tcrgitc dark except vontrally, second and tiiird
tergites dark apically and laterally; genitalia dark basally; hind
femora strongly darkened apically; wings almost hyaline. Head:
Frontal bristles excei)tiomilly long and slender; antialos as close to
j-uj)ra-antennals as to lower frontals; inner supra-antennals only half
as long as outer; antennal arista no longer than head height; palpi
with weak bristles basally in addition to four outwardly directed
apical or sui)apical ones and an inwardly directed one; genal angles
with two strong l)ristles. Thorax: Dorsal pulx-srence short and close,
becoming bristly posteriorly; posterior scutal margin with a pair of
inner bristles over half as long as the usual outer rmes; bristles along
outer margin of scutum as long as free end of K,.l. ; scutcllum with a
pair of long posterior bristles and a pair of very reduced hairlike ones

400

PROCEEDINGS OF THE NATIONAL MUSEUM

anteriorly ; pleura bare except for four ventral bristles and a postero-
dorsal patch of pubescence on the propleura. Legs: Hind tibiae and
tarsi together a little longer than wings and with relative lengths of

36 Megoeella stuntzl

Figures 33-35. — Phoridae of Guam: 33, Female of Megaselia scalaris (Loew); 34, male
of Megaselia scalaris: 35, Megaselia stuntzi, new species,

THE PIIORID FLIES OF GUAM — BOHART

401

2:3:5; hind tibiae with a simple dorsal row of 11 sotae along their inner
sides; midtibiae with a weak dorsal row of setae; fore coxae with three
strong apical bristles, the longest as long as scutellar bristles; ventral

Figures 36-38. — Phoridae of Guam: 36, Megaselia suit, new species; 37, Megaselia
parabaiiseta, new ipeciei; 38, Megaselia teti/emur, new species.

402 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

margins of hind femora with hairs of apical third longer than else-
where. Wings: About two-fifths as broad as long; setae along costa
moderately long, becoming more upright beyond point midway from
humeral cross vein to apex of Ri ; costa nearly two-thirds as long as
wings ; placement of veins as shown in figure 36 ; second vein without
setae basally. Ahdcnnen: Lateral areas of first tergite rather closely
covered with minute hairs ; tergites 3 to 5 each with lateral patches of
a few small hairs; sides of tergites 5 and 6 uniformly haired, with,
distinct apical fringe, especially on tergite 6 ; sternites 3 to 6 ventrally
with rather numerous setae or coarse hairs ; genitalia of moderate size,
the lamella large, creamy in color, with terminal bristles long and
delicate.

Holotype (U. S. N. M. No. 57991) and three paratype males : Agana,
Guam, June 1945, reared from fresh pig dung (G. E. Bohart). One
paratype male : Pago River, Guam, June 1945, swept from dense foliage
near a garbage dump (G. E. Bohart).

Reinarhs. — On Guam this species resembles only scalaris Loew in
general appearance but can be separated by its greater extent of yellow
on the abdomen, more confined propleural pubescence, and more deli-
cate terminal bristles on the genitalia. It is close to sauteri Brues from
the Philippines (Brues, 1936) but has longer costal setae, darker palpi
and legs (grayish instead of whitish), and weaker supra-antennal
bristles. It runs close to safunae Malloch from Samoa (Malloch, 1935)
but differs by having yellow f rons and antennae and more yellow on the
abdomen. In Malloch 's key to North American species (1912) it runs
to subflava but has narrower costal cells, more bristles on the pro-
pleura, and weaker hairs on the sides of second abdominal tergite. It
is much larger than hisecta Brues to which it runs in Brues's key, is
a deeper yellow and has a longer costa.

MEGASELIA STUNTZI, new species

Figure 35

Holotype^ male. — Length 1.7 mm.; length of wing 1.4 mm.; body
dark brown above, brown laterally and ventrally, without yellow or
creamy areas ; hind femora apically darkened. Head : Frontal bristles
relatively short but very thick and arranged as in figure 35 ; lower mar-
gin of head, seen from the side, with six bristles. Thorax: Dorsal pu-
bescence short and close ; lateral margin of mesonotum with four stout
bristles before the wing base ; propleura with about 12 scattered small
hairs in the upper posterior third and two small bristles at the ventral
corners; posterodorsal portion of mesopleura with a patch of about
12 short hairs a little stronger than propleural ones ; scutellum with
two pairs of strong subequal bristles; posterior scutal margin with
two pairs of bristles, the inner nearly as long as the outer. Legs:

THE PHORID FLIES OF GUAM — BOH ART 403

Hind tibiae and tarsi together distinctly longer than wings; inner
posterior margins of hind tibiae with eight bristles; hind femora with
basal ventral bristles longer than apical ventral ones; mid coxae with
three strong bristles. Wings: Slightly yellowish, with costal fringe
and placement of veins as shown in figure 35; second vein without
M'tae. Abdomcit: Uniformly blackish brown; segments nearly bare
except for apical fringes and for small lateral patches on first two
segments; sternal hairs small and few; genitalia small, with a short
lamella, bearing slender lamellar bristles; gcnitalic details as in fig-
ure 35.

Holotype (U. S. X. M. No. 57902) and one paratype male: Point
Oca, Guam, June 21, 1945, reared from dead mollusks (G. E. Bohart
and J. R. Stuntz).

Remarks. — This species resembles setifemur but is larger and has
nR':-opleural vestiture. It also resembles paraba.nseta but has no
dominant mesopleural bristle and has a much shorter costal wing
fringe. In Malloch's key to the Phoridae of Samoa (1935) it runs to
M. paciiica but has the first longitudinal vein ending closer to the
second than to the humeral cross-vein.

The fly is named for J. R. Stuntz, wlio did much of the rearing work
on Guam for our studies on filth-inhabiting flies.

MEGASELIA PARABASISETA, new species

Figure 37

Holotype^ male. — Length 1.1 mm. ; length of wing 0.9 mm., body
color a uniform dark brown, palpi, pleura, sides of abdomen, and legs
light brown but not yellow; wings distinctly brownish, frons blackish
brown; apex of hind femora scarcely darkened. Head: Frontal
bristles moderately long and exceptionally thick; lower supra-
antennals weaker than but extending as far forward as upper ones;
antiales nmch closer to lower frontals than to upper supra-antennals;
j)alpi very broad, with five roughly similar major bristles extending
outward from outer, forward margin; labium greatly exi)anded in
softened specimen; antennal arista no longer than heigiit of head;
cheeks apparently with only one strong bristle at genal angle and with
three weak ones farther forward. Thoraa': Dorsal hairs short but
strong, becoming sparse near wing and bristly toward posterior nuir-
gin ; scutum and posterior edge of scutellum each with only one pair of
bristles; propleura with a single dorsal posterior row of setae and two
ventral bristles; mesopleura with dorsal posterior corners bearing a
l)at«h of small setae in addition to a stronger bristle: anterior spiracles
enclosed in a separate sclerite. Legs: Moderately long, the posterior
tibiae and tarsi about as long as the wings and with relative proportions
of 3:4; posterior tibiae with a single dorsal row of only four or fi\e

404

PROCEEDINGS OF THE NATIONAL MUSEUM

setae; ventral margin of posterior femora with hairs of about equal
length throughout, a little weaker in the middle; metatarsi without
outstanding setae on the fringes. Wings : Rather narrow, about one-

Figures 39-41. — Phoridae of Guam: 39, Diploneura cornuta (Bigot); 40, male of Chon-
ocephalus subglaber, new species; 41, male of Chonocephalus hirsutus, new species.

THE PHOHID FLIES OF (JUAM — BOIIAHT

4Uo

tliiitl as broad as loup: ami wiili hairs of costal frinjre s[)ar c, about II
in each row; basal portion of R^^ boarin<jj on the inner sido a seta
ai)out as lonjjf as the costal setae; veins i>laceil as shown in ll;_'ure •57.

Fk.i RES 42— 15. — rhorida" of (Juam: 41, Female of Puliciphota nigTivtntri>, new species;
43, male of /*tt/iV I'pAora nigriventris; 44, fcm-'l'- ' ■> Puliri:,hnrii rrytr.ini m-w Kpcrir, 4r«
male of Puliciphora toymani.
710567—40 2

406 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ae

Abdomen: Tergites uniformly dark brown, sparsely hairy, the apical
fringes not longer than hairs on other portions except on sixth tergite ;
genitalia small, aedeagus with a complex group of hornlike appendices;
lamella single, with short terminal setae.

Holotype (unique) (U. S. N. M. No. 57989) : Pilgo River, Guam,
June 1945 (G. E. Bohart).

Reraarhs. — This species resembles sctifemur in general appearance
but has mesopleural vestiture. It runs to arizonensis Malloch or
monticola 'M^illoQh. in his key (1912) but has a longer costa than
arizonensis and differs from monticola in having the first two divisions
of the costa subequal instead of 2 : 1. It also differs from monticola
in having the outer supra-antennals only one-third as far apart as the
width of the f rons. It runs to galugensis Brues or laluvensis Brues in
the key to Philippine species (1936) but has narrower wings than
lalvwensis and the seta on the base of Ri+o, which is missing in the
latter. It differs from galugensis in its sparse costal bristles and
darker color. It is apparently similar to hasiseta from Samoa (Mal-
loch, 1935) (hence its name) but has mesopleural vestiture.

MEGASELIA SCALARIS (Loew)

Figures 33, 34
Phora scalaris Loew, Berlin Ent. Zeitschr. (Centuria VII), vol. 10, y. 53, 1866.

Length 1.6 to 3 mm. ; length of wing of a 3-mm. specimen 2.8 mm.,
body yellowish (frons of male sometimes largely black) with indistinct
brownish markings on the scutum, forming a pair of dorsal longi-
tudinal stripes, another pair of more lateral and posterior stripes, and
several angular spots anterior to these ; abdomen marked as illustrated
but sometimes more extensively dark so tliat the yellow is constricted
to basal and apical bands on the second and third tergites and basal
central spots on the fourth and fifth tergites. The first tergite and
the apical lobes of the last always remain largely yellow; palpi,
halteres, pleura, and legs (except for apices of hind femora) creamy
yellow. Head: Frontal bristles long and moderately heavy, the
supra-antennals about equidistant and the outer ones only a little
stronger than the inner; antiales a little higher than the outer supra-
antennals and much lower than but nearly as far apart as lower
f rentals; proboscis of females short and fleshy. Thorax: Propleura
with numerous scattered hairs in addition to a dorsal posterior fringe
and two to four ventral bristles; pleura otherwise bare; posterior
margin of scutum with single pair of long slender bristles and scutel-
lum with two pairs of subequal bristles ; scutal pubescence dense and
composed of very small hairs ; halteres yellow, sometimes with darker
apical spot. Wings: Costa distinctly longer than succeeding length
of wing; veins bare except for moderately short and dense costal
fringe and the usual bristles on the basal posterior margin ; costa from

THE riLOHlD FLIES OF GUAM — BOHAiiT -407

humeral cross vein to second longitudinal sli<ihtly lonrjer than from
the latter to apex of third lonijitudin;!]. L </.<: Miiltil)iae \Yith a weak
fringe of setae outside the dorsal hair fringe as well as a stronger
one inside; hind tibiae with an outer fringe composed of 9 to 12
rather strong setae; fore tibiae setose dorsally but without a definite
low of setae; hind tibial spur of male nearly as long as the metatai"sus.
Ahdoincn: Tergitcs sparsely covered with short, stiff hairs laterally
and with long ones apically on tergites 5 to 7. Genitalia as figured,
the lamella yellow with strong terminal bristles.

This is a cosmopolitan species that breeds in a wide variety of
materials. It is easily dislinguisheil from the other species on Guam
by its color pattern, types of pleural pubescence, and the male
genitalia.

Genus DIPLONEURA Lioy

Dn»LONEURA CORNUTA (Bigot)
FlQUBE 39

Fhora cornula Bigot, in de la Sagra's "Histoire physique, politique ct uaturelle
lie rile (le Cuba" (French eil.). p. 827, 1857.

Length 1.4 to 2.1 nun.; length of wing of a 3-mm. specimen 0.95
mm. ; color yellowish brown ; f rons blackish above ; thorax with median
and lateral brown areas; scutellum brown; abdomen yellow with
blackish areas laterally and apically on tergites 2 to 5 and covering
most of 6 and 7; palpi, proboscis, pleura, and legs rather clear yellow;
wings hyaline. Head: Frons with long slender spines, averaging over
half as long as width of frons; spacing as shown in figure Ii9 ; antennal
arista slightly longer than head width; proboscis of female narrow,
chitinized, extending one-third its length beyond palpi ; palpi with five
long apical and subapical spines, fourth from apex longer than antial
bristleii and directed downward; checks with two long bristles near
genal angles and a single hjiig one at level of antennal inseitions.
Thorax: Dorsum broader than long, with live bristles along each lal-
( ral margin, the longest on posterolateral corner of scutum; apii-al
margin of .s<Mitum olherwisi; with one pair of long bristles and two
mMcii shorter inner pairs of hairlike ones; scutellum with one pair of
I'Mig bri.stles on the posterolateral corners. Pubescence of pleura as
shown in figure )59. Lerjn: Posterior tibial' with dense dorsal fringe of
hair and u scries of weak setae inside and hidden by the dorsal fringe;
anterior tibiae with four setae on outer margins; inner sides of mid-
tibiae with hairs arranged in transver.se series and with apical spurs
nearly as long as nu^tatarsi; posterior melatarsi with two outstanding
setae before the middle along ventral margins. Witkjh: With whitish
microtrichia; placement of Atins and cosliil fringe as shown in fiijnrc

408

PROCEEDINGS OF THE NATIONAL MUSEUM

Parafonnio molluscovoro

SL aajjro - ontQnncI

LF Ictvsf ffontol

UF upper fronfcl

PO pre-ooeiiar

OG ocel'cr

Chonocepholus subglober

•*S Cho^ocep^o;us hirsutus

Figures 46-48. — Phoridae of Guam: 46, Parafannia molluscovora, new species; 47, female
of Chonocephalus subglaber, new species; 48, female of Chonocephaluf hirsutus, new
species.

THE PHORID FLIES OF GUAM — BOHART 409

39. Abdomen: Tergites 1 to 5 practically bare, G and 7 with sparse
general pubescence and long apical fringe hairs; sternitos distinctly
setose, the longest ones on st+rnites 5 to 7 nearly as long as frontal
bristles; ovipositor reddish with dorsal and ventral pairs of hairs as
long as frontal bristles.

Tliis tropicopolitan species is the only representative of its genus
so far found on Guam, It has been recorded from southern Kurope,
Asia, and southeastern United States. The figures of the male geni-
Talia should serve to distinguish it from other species, if they should
be found.

It was reared on Griam from decaying mollusks and swept from
vegetation along the seacoast.

Genus CHONOCEPHALUS Wandolleck

CHONOCEPHALUS HIRSUTUS. new species

Figures 41, 48

Ilolotype^ female. — Length 1 nun. ; body brown above, testaceous be-
neath, with a tine transverse subapical dark brown streak extending
about three- fourths the distance across each abdominal tergite. Head:
Eyes with about 12 facets; sides of head with four bristles anterior
to eyes, the posteriormost about one-third as long as head ; palpi about
as broad as antennae and with terminal bristles as long as palpi.
Thorax: Propleura with numerous minute hairs but no dominant one;
notal pubescence short and erect. Legs: Sparsely hairy : forelegs with
first tarsal segment a little shorter than next two combined and about
half as long as first hind tarsal segment. Abdomen: Only slightly
darker above than below; tergites 2 to 5 with two transverse rows of
setae, excei)t laterally where there are about 10 setae on each segment
as strong as the ones in the transverse rows; fifth and sixth sternite.s
with small setae laterally; sternites 4 to 7 with ventral setae, the
apical ones on sternite 7 longer than setae in the tergal rows; spatula-
shaped sternal strnrture with "handle" about three times as long as
"blade'' and with "blade" about as wide as long.

Allotype^ male. — Length 1.1 inm.; length of wing 1 mm.; blackish
brown, somewhat velvety; auteimae, palpi. ])leurae, and legs brown:
wings lightly infuscated. Tlcad: Bristles weak, the postooellars
aud lowermost frontals the strongest; median groove of frons
)iarrf)\v, not dividing the lower portion into two widely separated
rid'/es. Thorax: I)oi-sal pubescence sparse and fine: posterior scutal
margin with four bristles, the basal ])air about half as long as the
apical. Lateral vestiture as in figure 41. 'Winrjx: Veins very delicate;
costal fringe composed of many .short, weak setae; humeral cross
vein and sub^osta absent. Abdomen: First tergite as long as second

410 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9*

and sixth twice as long as fifth ; tergal pubescence minute ; genitalia
large, the right side with a long bootlike process; the left side with
its hair fringe mainly confined to a single row close to posterior
border.

Holotype female and allotype male (U. S. N. M. No. 57994) : F. E. A.
Farm, near Agana, Guam, October 1945, female taken from pig dung,
male swept from above mud in the pig pen (G. E. Bohart) . Paratype
male : From light trap, Point Oca, Guam, June 1945 (G. E. Bohart and
J. L. Gressitt) .

Remarks. — The two specimens of males (both mounted in euparol)
vary in the extension of the aedeagus and the expansion of the wings,
but this is due to mounting techniques. The allotype has one meso-
pleural bristle whereas the paratype has two. The holotype is
mounted in euparol but is unfortunately somewhat collapsed and
shrunken. This species is close to subgldber^ but the female has
stronger and more numerous abdominal setae and a more uniform
abdominal color. The male genitalia also differentiates it from
subgldber and from other species for which genitalia have been
described or figured.

CHONOCEPHALUS SUBGLABER, new species
FiGtJBES 40, 47

Holotype^ female. — Length 0.9 mm. ; dark brown to blackish brown
above, brown laterally on thorax and creamy white laterally and
ventrally on abdomen ; abdominal tergites with broad, complete trans-
verse preapical black bands containing pale circles at the bases of the
fringe setae. Head: Eyes with about 15 facets; sides of head with
five bristles anterior to eyes, the posteriormost about one-fourth as
long as head; palpi narrower than antennae. Thorax: Propleura
with a dorsal bristle in addition to scattered small setae. Abdomen:
Tergal pubescence scattered and minute except in preapical fringes;
bases of fringe hairs completely enclosed in the broad preapical black
stripes ; sternites 4 to 6 with a few minute setae medially and sternite
7 with a median sclerite bearing short setae; spatula-shaped sternal
structure with "handle" less than three times as long as "blade."

Allotype^ male. — ^Length of wing 1 mm. ; body color as in hirsutus.
Head: Bristles moderately strong, those extending from the eye along
the anterior margin of the frons and along the forward prolongation
of frons gradually diminishing in strength but most of them as strong
as postocellars or postverticals ; median groove broad and shallow,
separating forward prolongation of frons into widely separate
ridges. Thorax: Hairs of propleura scattered and mesopleura with
three setae. Legs: As in hirsutus but not so sparsely pubescent.
"Wings: Costal fringe short, with numerous hairs, those on basal half

THE PHORID FLIES OF GUAM — BOHART 411

distinctly lonirer and sparser than elsewhere. Abdomen: Sixth
tergite only one and one-half times as long as fifth and with notice-
able hairs in addition to apical fringe: genitalia without a strong
bootlike process on left side but with a fingerlike process on the right
side bearing an ai)ical peg and a thick spine beyond the middle.

Holotype female (U. S. N. M. No, 57995) and 15 paratype females:
Point Oca, Guam, June 1945, from breadfruit (G. E. Bohart and
J.L. Gressitt).

Allotype male (U. S. N. M. No. 57995) : Point Oca, Guam, June
1945 (G. E. Bohart and J. L. Gressitt); 3 paratype males: Point
Oca, December 1945 (reared from bananas by J. L. Gressitt).

Remarks. — The female differs from hirmtus by its weaker abdomi-
nal pubescence and stronger transverse dark bands on the abdomen.
It is almost identical with buccaUs Malloch in the female but males are
readily distinguishable by the genitalia. The male differs from
hirsutus chiefly in the stronger frontal bristles and in the genitalia.
It differs from the closely related buccalis Malloch (1912) and from
any of the species described from the Bismarck Archipelago by
Schmitz (1929a, pi. 1) by the genitalia processes mentioned in the
above description.

Genus PULICIPHORA Dahl

PULICIPHORA WYALANI, new specic.i

FiouBEs 44, 45

Holotype, female. — Length 1.1 mm. ; brown above and pale testaceous
beneath; frons usually with indefinite median and oblique sublateral
darker bars; thoracic pleura without dark areas. Head: Frons with
only one pair of lower frontals. Thorax: Proplcura with a few scat-
tered setae in addition to the dorsal bristle; dorsum with three pairs
of bristles, the lateralmost inserted near posterior corners of meso-
pleura. Legs: Hind tiljiae with hairs of dorsal margin longer and
stouter than elsewheroon legs. Ahdoynen: Tergite 6 with a small, lunu-
lar sclerite at the base; sternitc 5 uniformly ringed with setae; sternite
2 with a group of setae just below the tergite.

Allotype., male. — Length 1.2 mm.; length of wing 1.1 mm.; thorax
dark brown above, light brown beneath ; dorsum of abdomen and frons
blarkish brown. Head: Antialos aljout as far apart as the lower
frontals; palpi with five strong bristles. Thorax: Proplcura with scat-
tered setae in addition to dorsal bristle; posterior border of thorax
with one pair of bristles near the lateral corners; .scutelhnn with four
bristles, the inner ones about two-thirds as long as the outer. Tegs:
Midtarsi only slightly longer than midtibiae, about equal to fore tarsi.
Winijs: Very broad, witli radial vein ending at about middle. ^6-
c/omcn.' Genitalia large, witli un aiilerioily developed ventral loop and

412 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

an extensible process, apical lamella elongate and with a slender
ventral lobe.

Holotype female, allotype male (U. S. N. M. No. 57997), 6 paratype
males, and 10 paratype females: Point Oca, Guam, June 1945, from
dead shellfish, and December 1945 from canned salmon (G. E. Bohart
and J. L. Gressitt).

Remarks. — Females are somewhat variable in the distinctness of the
head markings and in the color of the haustellum, which may be light
or dark. Males sometimes have the terminal process (aedeagus) of
the genitalia retracted.

This species can be distinguished in the female from nigriventris by
the paler dorsal color, lack of dark pleural areas, and more complete
abdominal setosity. The male can be told from nigriventris and other
species by its large aedeagus and the anterior loop of its associated
genitalic structures.

The fly is named for C. L. Wyman, who worked with me many
months overseas and in Washington, D. C.

PUUCIPHORA NIGRIVENTRIS. new species
FiGXJBES 42, 43

Holotype^ female. — ^Length 1 mm. ; clypeus, f rons, antennal grooves,
sides of pronotum, anterior margins of propleura, anterior band, and
ventral surface of sternopleura black ; legs, antennae, palpi, proboscis,
cheeks, occiput, central portions of thoracic pleura and metanotum,
basal two-thirds of first abdominal segment, and basal lunule of fifth
segment yellowish brown or testaceous; abdominal sternites and all
of abdomen beyond segment 5 creamy to translucent whitish. Head:
With two pairs of lower frontals and with supra-antennals about as
long as postocellars. Thorax: Propleura with only one distinct seta;
posterior margin of scutum with two pairs of subequal bristles. Legs :
Hind tibiae with hairs of dorsal margin rather long but no stouter
than elsewhere on the tibiae. Abdomen: Segment 5 (with dorsal
gland) with rows of setae which are incomplete laterally ; segment 2
without lateral setae ; segment 6 without a dorsal sclerite.

Allotype^ male. — Length 1.2 mm. (expanded), length of wing 0.9
mm. (incompletely formed) ; blackish brown, thorax somewhat lighter.
Head: Antiales much closer together than lower frontals; palpi with
four strong bristles. Thorax: Propleura with only one distinct hair
or bristle: prothoracic spiracle in a di'^tinct sclerite. Legs: Midtarsi
considerably longer than foretarsi or midtibiae. Wi7igs: Poorly de-
veloped in holotype because of teneral condition. Abdomen: Genitalia
not particularly large ; apical lamella short and with prominent ven-
tral lobe; ventral portion of genitalia not produced basally into ab-
domen.

THE PHORID FLIES OF GUAM — BOHART 413

Holotypo. allotype (I'.S.N.M. No. rM'.t'.H')), and 8 paratypos (iiu-liul-
ing one male) : Point Oca, (ruani, June 11)45, reared from dead
shellfish (G. E. Bohart nn<l J. L. Gressitt).

Rtr/uirl-f<. — This specios resembles wynunii but can be distinguished
by its blacker color and fewer abdominal setae in the female and by
the smaller, simpler appearing genitalia in the male. It comes close
to tol'i/oensii* Kinoshita (see 8chmitz, in Lindner, llKiS) but lacks the
mesopleural bristle and scattered propleural setae of the latter.

PARAFANNIA, new genus

Head: Clypeus convex and strongly projecting, separated from
antennal scrobes by membranous area; third antennal segment sub-
spherical, with arista inserted dorsally and over twice as long as
frons; checks produced ventrally and posteriorly, the genal angles
considerably below and behind the nearest margin of the eyes and
bearing a single long bristle; cheeks otherwise with only a few short
bristles at inner angles, palpi with only two long bristles near apices,
with two rows of lesser ones extending along entire outer and ventral
sides ; proboscis fleshy, broadly and deeply bilobed ; frons much broader
than long, with one pair of erect supra-antennals, one pair of out-
wardly directed antiales, two pairs of laterals, the lowermost only
slightly below the level of preocellars; upper verticals directed inward
and postverticals directed outward; ocelli rather small and far
separated. Thorax: Scutum about two-thirds as long as broad, with
six pairs of lateral bristles, one pair along truncate posterior margin ;
scutellum posteriorly with one pair of bristles and with strongly con-
vex posterior margin ; propleura lateral, about twice as high seen from
the side as posterior portion of pronotum; anterior spiracles not above
dorsal margins of pro- and mesopleura; mesopleura^^.not divided into
anterior and posterior sections, more than twice as high as dorsal
length, entirely bare; a distinct 5-sided sclcrite separated from the
hypopleura and interposed between the latter and the pteropleura;
postnotum shorter than scutellum. Legs: Dorsal side of foretibiae
with a single subbasal seta, which is somewhat stronger than hairs
along doi-sal margin; midtibiae with a subbasal pair of strong bristles
on dorsal side, with two strong bristles near apex; hind tibiae without
dorsal hair fringes, with a single strong biistle on outer side one-third
the distance from base to apex, and apically with a short subdorsal
bristle in addition to two moderate ventral bristles. Wings: M-^.s {sec-
ond longitudinal) not separated apically from M^^b (third longitudi-
nal) ; Rj + Sc (first longitudinal) short ; base of Rs on inner side with
long bristle; veins not swollen and pterostigmae lacking. Ahdom-rn:
Nearly bare dorsally except for apical fringe of sixth tergite, laterally
and ventrally with short setose hairs set in distinct sockets (as in Pull-

414 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ge

ciphora) ; ovipositor, when exserted, composing seventh and eighth
segments, which are tubular and have fine chitinized striae, and
terminal structures composed of a dorsal piece connected to a pair
of ventral sclerites and a pair of spatulate distal processes.

This genus is in the subfamily Phorinae of Schmitz and runs close
to Gitrago Schmitz in his key to the world genera (1929a). It differs
from this, however, in the possession of an anterodorsal bristle on the
hind tibiae and in the strongly produced clypeus. It differs from
Hypocera in having bare mesopleura and from Diploneura in lacking
a free end to the radial vein.

Type of the genus : Parafannia moUuscovora^ new species.

PARAFANNIA MOLLUSCOVORA, new species
Figure 46

Holotype^ female. — Length (expanded) 2.2 mm.; length of wing
1.6 mm.; sooty black, antennae, palpi, legs, pleura dark yellowish
brown to dark brown or nearly black, halteres smoky brown, sides
and venter of abdomen smoky yellow, becoming sooty apically ; wings
slightly infuscated, veins brown. Head: Frons about twice as broad
as long; antiales over twice as close to supra-antennals as to eyes;
lower frontals and preocellars nearly in line; clypeus with a dark
basal band. Thorax: Scutum with rather sparse brownish pubes-
cence; scutellar bristles longer than scutellum and bending inward;
hindmost pair of lateral scutal bristles about as long as distance from
humeral cross vein to apex of the radius ; propleura with one slender
seta on posterior angles, two or three small hairs near the posterior
margins, and three small setae near the ventroposterior corners.
Legs: Hind tibiae and tarsi together about four-fifths as long as
wings, the tibiae four-ninths as long as tarsi ; subbasal bristle on hind
tibiae about as long as tibial width at that level ; posterior metatarsi
one-fourth as broad as long; dorsal margins of midtibiae mostly
smoky yellow but with broad dark band in region of subbasal pair of
bristles ; subbasal bristle on inner dorsal margins of f oretibiae shorter
than tibiae width. Wings: Setae fringe of costa short, dense, and reg-
ular; prehumeral portion of costa with two long bristles; bristle on
base of Rs about as long as lateral bristles on frons; placement of
veins as in figure 46. Abdomen: First tergite about half, third about
two-thirds as long as second ; tergites 1 to 5 practically bare except for
tiny apical fringes; sides and venter of abdomen with scattered
setae, which become longer and more numerous toward apical seg-
ments and have black pigmentation at their bases ; processes at tip of
ovipositor with a few long hairs ; seventh and eighth abdominal seg-
ments (forming "tube" of ovipositor) finely striated with black, the
latter with a pair of dorsal longitudinal chitinous rods.

THE PHORID FLIES OF GUAM — BOHART 415

Ilolotype (U. S. X. M. No. 57993) and 15 female paratypes: Point
0>"i, A^'ana, Guam, Juno 1945, reared from decaying mollusks buried
in ant colonies just beneath the ground surface (G. E. Bohart and
J.L. Gressitt).

/Remarks. — This fly is so distinctive that a new genus was erected
for it. Other species, if found, may show that some of the characters
used in the generic description are of specific vahie only.

Ah hough over 30 specimens were reared, no males were obtained.

KKY TO TIIK rilORIDAE OF GUAM

1. Winpless 2

Winged 5

2. Head strongly flattened dorsoventrally and with front produced beyond

antennae; d'Tsum of thorax without strong bristles; eye smaller than

antenna. (Chorioccphalus) 3

Head neither flattened nor conspicuously produced; dorsum of thorax
with several pairs of strong bristles; eye larger than third antennal
segment. (PuUciphora) 4

3. Abdomen, view from side, with tergites uniformly haired and without

distinct apical fringes; venter of abdomen light brown, almost as dark

as dorsum Chonocephalus hirsutus

Abdomen with very fine sparse hairs except for apical fringes of tergites;

blackish dorsum of abdomen contrasting strongly with creamy white
venter Chonocephalus subglaber

4. Frons, mesonotum, and first 5 abdominal tergites uniformly blackish;

thoracic pleurnn strongly marked with dark gray along most sutures;
hind tibia without noticeable fringe of bristles or stiff hairs; fifth
abdominal segment not uniformly ringed with several rows of hairs or

short bristles Puliciphora nigriventris

Most of frons and much of mesonotum and abdominal tergites testaceous
to bnnvti, not strongly different from thoracic pleural color; fifth and
sixth abdominul si-gments uniformly ringed with short bristles or hairs;
hind tibia with distinct posterior fringe of strong hairs or bristles.

Puliciphora wymani

5. Frons covered with long, strong bristles and not produced forward be-

tween antennal bases 7

Frons with iit most 2 or ?, pairs of strong bristles and produced forward
between antennal bases. (Chonocephalus) 6

6. Frons broadly imprfs.sfd mfdinlly, fmni above appearing like a pair of

ridgt-s b<?tween antcimal ba.sfs, and with anterior border near eyes
having a series of 3 or 4 moderate bristles; me.soijlcuroti with several

setae Chonocephalus subglaber

Frons narrowly impressed along midline and with bristles along anterior
border scattered and weak; me.sopleuron wltli a single seta.

Chonocephalus hirsutus

7. Radial sector unliranehed 8

Radial sector branclied near ajM'x 10

8. Clyp<'us strongly produeefj forward at oral margin; hind tibia with a

spine on outer surface toward the base; midtihiu wlili a pair of bristles

ffiward ba.se Parafannia molluscovora

Clypetis not prfKlurrd at oral margin; tibiae without basal or sub-basal
bristles. {Puliciphora)

416 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 93

9. Propleuron with about 5 small setae in additicm to the dorsal bristle ;

genitalia with long anteroventral extension Puliciphora wymani

Propleuron with dorsal bristle only ; genitalia without conspicuous ante-
roventral extension Puliciphora nigriventris

10. Preantennal bristles (on anteromedian margin of frons) directed back-

ward ; midtibia with a pair of subbasal bristles Diploneura cornuta

Preantennal bristles directed forward; midtibia without bristles before
middle. (Megaselia) 11

11. Abdomen with extensive yellow areas anteromedially on most tergites 12

Abdomen blacli or dark brown above 13

12. Fourth and fifth abdominal tergites entirely pale; frons with inner

preapical bristles much shorter than outer ones Megaselia suis

Fourth and fifth abdominal tergites partially dark; frons with inner
preapical bristles nearly as long as outer ones Megaselia scalaris

13. Posterodorsal corner of mesopleuron with a patch of small setae 14

Posterodorsal corner of mesopleuron without setae ; basal half of ventral

margin of hind femur with 4 or 5 long hairs Megaselia setifemur

14. Posteroventral corner of mesopleuron with a small bristle in addition to

setae ; costal margin of wing with about 11 bristles, which are longer

than distance between costa and radius Megaselia parabasiseta

Posteroventral corner of mesopleuron with uniform setae only ; costal
margin with about 15 short bristles Megaselia stuntzi

LITERATURE CITED

BORGMETER, T.

1925. Novos subsidies para o conhecimento da familia Phoridae (Dipt.).

Arch. Mus. Nac. Rio de Janeiro, vol. 25, pp. 85-269, 17 pis.
Brues, Charles T.

1903. A monograph of the North American Phoridae. Trans. Amer. Eut,

Soc, vol. 29, pp. 331-404, 9 pis.
1936. Philippine Phoridae from the Mt. Apo region in Mindanao. Proc.

Amer. Acad. Arts and Sci., vol. 70, pp. 365-466.
Malloch, John R.

1912. The insects of the dipterous family Phoridae in the United States

National Museum. Proc. U. S. Nat. Mus., vol. 43, pp. 411-529, 7 pis.
1935. Insects of Samoa, pt. 6, fasc. 9, pp. 329-366. British Museum.

SCHMITZ, H.

1929a. Revision der Phoridae, 212 pp. Berlin and Bonn.

1929b. Zur naheren Kenntnis einiger von Dahl beschreibener Phoriden des

Bismarck-Archipels, Konowia, vol. 8, pp. 111-122.
1938. Phoridae. In Lindner, Die Fliegen der palaearktischen Region, voL

4, pt. 33, pp. 1-64.

U. E COVEriNMCNT PRiNTlNG OFFICE. 1?47

PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM

iuued Kfllfv vl U?*^D ^y '^*

SMITHSONIAN INSTITUTION

U. S. NATIONAL MUSEUM

Vol. 96 Washington: 1947 No. 3206

A GENERIC REVISION OF THE ICHNEUMON-FLIES OF
THE TRIBE OPHIONINl

By R. A. CusHMAN

A NUMBER of years ago I planned a revision of the North American
species of the hymcnoptcrous genus Ophion and to that end assembled
from many sources a great number of specimens. Study of this
material made it evident that Ophion as we had known it was made
up of several more or less discordant elements and indicated the need
of a generic revision of the tribe Ophionini. This was undertaken,
with the results set forth in the following pages.

Unfortunately, representatives of a considerable number of the
described genera have been unavailable to me, and some of these I
have found impossible to place in my arrangement because of the
inadequacy of the descriptions. Several, however, 1 have been
enabled to place through the generous cooperation of J. F. Perkins,
of the British Museum. Such characters as the possession or lack of
(he postpccU>ral carina, the position of the second abdominal spiracle,
the presence or absence of a fenestra when scleromes are lacking, and
even the form of the mandibles and the course of the radius are, in
many cases, not referred to in generic descriptions; and every descrip-
tion omits mention of one or more of the series of (characters which I
have chosen for use as generic group characters. Most of the genera
of which no identified specimens arc available and of which I have
been unable to identify any species in the large amount of material
bi.'fore me are not included in the key to genera. However, the
probable aflinitics of each arc discussed.

417

418 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Study of such genera as have been available to me has led to the
discovery of a number of characters not previously employed in the
classification of the group. Using these characters, together with the
best of the old characters and others gleaned from generic and specific
descriptions, I have attempted to produce a more nearly natural classi-
fication than any pubUshed heretofore.

SOURCES OF MATERIAL

This study is based principally on material in the collection of the
United States National Museum, which contains specimens of
Ophionini from all the zoogeographic regions of the earth, including
representatives of a majority of the previously described genera and of
all the new genera described in this revision. A vast amount of
North American material, mostly referable to Ophion and Enicospilus
secured on loan from the collections of many institutions and private
collectors, has been studied. Particularly important, with respect to
the present revision, are the collections received from the CaUfornia
Academy of Sciences, the American Museum of Natural History, the
University of Kansas, Cornell University, and the private collection
of Henry K. and Marjorie C. Townes, since these contain material
of many more of the Nearctic genera other than Ophion and Enicospilus
than do any of the others. To these and to other institutions from
which material was secured I hereby extend my thanks.

ILLUSTRATIONS

All the figures, except plate 55, figure 71, were drawn by Arthur D.
Cuslmian, scientific illustrator, U. S. Bureau of Entomology and Plant
Quarantine. I am grateful to him for the accuracy of his drawings.

TRIBAL CHARACTERS AND RELATED TRIBES

Among the tribes of the Ophioninae are three, Ophionini, Hell-
wigiini, and Anomalini (Nototrachini) , that have traditionally been
separated from the rest of the tribes in tribal keys by the single
character of the position of the second recurrent vein basad of the
intercubitus. This has resulted in the placing in the Ophionini of
genera, especially certain genera of Therionini, that obviously do not
belong there when other characters are taken into consideration.

Some authors have included Hellwigia Gravenhorst in the Ophionini,
while Morley unites the Anomalini with Ophionini and excludes
Hellwigia. The genera Hellwigia and Hellwigiella Sz^pligeti are
obviously not at all closely related. Since this paper was sent to
press I have, through the kindness of Dr. H. K. Townes, been able
to examine specimens of both genera. Hellwigiella I find to belong
to the Ophionini, but Hellwigia I believe to belong to the
Campoplegini, as indicated on a later page.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 419

Tho jrenus Ophiopferus Brull6 has consistently been referred to the
Ophionini because of the strong^ly antifurcal second recurrent vein,
but in all other respects it is therionine.

Another tribe that comes into the complex is the Ophionellini, made
up of tho venationally anomalous OphioneUus Westwood and
flymennpharsalia Morley (Pharsalia Cresson), the latter referred by
Ashmead to the Anomalini because of the single calcarium on the
middle tibia. OphioneUus is unknown to me, but, in my opinion, all
the other characters of Hymenophfirsalia ally it too closely with such
therionine genera as Atrometus Focrster and Podogasier Brull6 to
justify either placing it in the Anomalini or maintaining it as a dis-
tinct tribe.

The following diagnosis in kej' form should present no difficulty in
the placing of a genus in its proper tribe:

KEY TO THREE OF THE TRIBES OF THE SUBFAMILY OPHIONINAE

1. Mandible, when closed and in cephalic view, with lower tooth not directly
posterior to upper tooth, but at a lower level and visible; eyes parallel or
divergent below, never distinctly convergent (i. e., face not narrower than
frons), always at least shallowly emarginate opposite antennal foramina;
occipital carina medially far below lovel of posterior ocelli, fading out below or
joining hypostomal carina at a considerable distance from lower articulation
of mandible, rarely entirely absent; cpomia entirely lacking; propodeal fpiracle
lateral in position, very long and slitlike; /ron< tibia without a minute tooth
at apex opposite calcarium; middle tibia with two calcaria; claws usually
long, each with a strong, many-toothed pecten; second recurrent vein basad
of intercubitus by a distance nearly or quite as great as length of intcrcubitus;
intercubilun strongly oblique and nearly continuous with basal abscissa of
radius; nrrvellun sharply broken, discoidella distinct, rarely nervellus not
broken but interstitial at its upper end with discoidella; hind tarsus with
apical two joints not esperially small; ovipositor rarely exsertcd.. Ophionini
Mandible^ when closed and in cephalic view, with lower tooth directly posterior
to upper tooth and not visible; eyea more or less convergent and at most with
inner margins arcuately concave; occipital carina medially usually at or near
level of posterior ocelli and joining hypostomal carina very close to lower
articulation of mandible, sometimes (Anomalini, Anomalon verbosum Cresson)
far below ocelli, rarely {Podognstcr Brull6) fading out below and not reaching
hypostomal carina or (Clalha Cameron) obsolete medially; epornia strong,
originating on lower anterior margin of pronotum and running dorsad nearly
to dorsal margin; propodtal spiracle subdorsal in position and long or short
oval but not slitlike ; /ron/ tibia with a minute tooth at apex opposifc cal-
carium; middle tibia usually with two calcaria, but sometimes (Anomalini,
Hymenopharsalia, OphioneUus, and an unnamed North American genus of
Therionini) with only one; claws simple or at most weakly pectinate and
usually small; second recurrent vein, except in Anomalini and genus Ophio-
pterus, distad of intercubitus or proximad by a flistancc; much less than
length of intercubitus, ran-ly illumennpharsnlia, Ophionrltus) entirely absent;
intercubitus perpendicular or nearly so, at lea.'^t not nearly continuous with
basal abscissa of radius; nervellus broken or unbroken; hind tarsus usually
with apical two joints small; ovipositor usually distinctly cxscrted. 2

420 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

2. Second recurrent vein far basad of intercubitus; middle tibia with only one
calcarium; occipital carina medially far below level of posterior ocelli.

Anonialini
Second recurrent vein usually distad of intercubitus, occasionally slightlj' basad,
rarely {Ophioptents) far basad or (Hymenopharsalia, Ophionellus) entirely
absent; middle tibia with two calcaria, rarely {Hymenopharsalia, Ophionellus,
and an unnamed North American genus) with only one; occipital carina
medially almost at level of posterior ocelli, rarely {Anomalon verbosum
Cresson) far below Therionini

HOST RELATIONS AND LARVAL CHARACTERS

Host relations and larval characters indicate that the three tribes,
Ophionini, Anomalini, and Therionini, are distinct natural groups and
support the placing of Hymenopharsalia in the Therionini. The
Ophionini are internally parasitic in lepidopterous larvae, the Anoma-
lini in coleopterous larvae, and the Therionini in lepidopterous pupae.

My knowledge of the larvae is based largely on exuviae extracted
from cocoons or host pupae from which known specimens were reared.
The most significant characters are to be found in the sclerotization
of the margins of the head capsule and mouth parts. In the accom-
panying illustrations of these sclerotizations in typical genera of each
of the three tribes, quite different patterns v/ill be noted.

In the ophionine larva (pi. 49, figs. 1-3) the entire labium (Im)
except the areas surrounding the palpi (Ip) is lightly sclerotized, with
a more heavily sclerotized horseshoe-shaped sclerome (sd) around the
opening of the silk duct in a deep emargination on the upper margin ;
there is no sclerotic bridge across the clypeus, the sclerotization extend-
ing for only a short distance along the frontal suture (/) above the
dorsal articulation of the mandible; the hypostomal margin (hy) of
the epicranium is heavily sclerotized with a long branch (st) extending
across the maxilla and a somewhat less heavily sclerotized extension
(oc) downward along the margin of the occipital foramen; the lower
margin of the stipes of maxilla (mx) is heavily sclerotized and the
surface of the cardo (cd) of maxilla lightly so; the mandibles (md) are
extremely small.

In the therionine larva (pi. 49, figs. 5-8), on the other hand, the
labium is sclerotized only along its superior lateral margins and the
margins of the silk duct opening, its lower margin being very poorly
if at all defined; there is a strong sclerome across the cl^'^peus (cl),
fused laterally with the pleurostomal margins, thus forming with the
hypostomal margin a strong brace extending entirely across the head
above the mouth, and sometimes spreading far back from the margins,
notably so in Therion and Heteropelma; the very heavily sclerotized
hypostomal margin lacks entirely the branch across the maxilla
and the extension along the occipital foramen ; the sclerotized margin
of the stipes of the maxilla is long and slender and there is no evident

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 421

sclerotizatii)n of the canlo; tho niaiulihlos arc very stout and overlap
across tlie median line.

The anomaline larva (pi. 49, fig. 4) is, in some respects, more or less
intermediate between those of the other two tribes. The labial ring
is narrow but apparently comj)lete, although its lower mari^in peihaps
outlines the palpal areas rather than the actual basal margin of the
labium, and there i.s a median e.xten.sion between the palpal areas;
the brace across the head above the mouth is complete but slender,
and the part across the clypeus is much more arched than in the therio-
nine larva; the ma.xillar branch is present but very short; the marginal
sclerome of the nuixiilary stii)es is long and slender and the cardo is
not sclerotized, the mandible is stout at the base with a long slender
point, the two mandibles nearly meeting on the median line.

TERMINOLOGY

For the most part the terminology employed is that characteristic
of the vast bulk of the literature of the Ichneumonidac, and that of
the wing veins and cells is in general that of Rohwer and Gahan's
"Horismology of the Hymen()j)terous Wing." ^ A few new or unusual
terms, however, need definition:

Abscissula. — The basal abscissa of radiella (pi. 53, fig. 52).

Basal constrict ion of propodcnm. — The constriction dividing that
part of propodeum basad of the basal carina or its normal position
into frenum (anterior portion) and basal area (posterior portion) (pi.
52, figs. 43, 44).

Epipleura. — The lateral portions of the tergites extending as free
flaps beyond the line of attachment of the ventral membrane. On
tcrgitc 2 the ej)ipl(!ura are usually set off by carinae and arc infle.xed,
but are not so defined or inflexed in the Tliyreodon group and in a
few other genera.

Fenestra. — A hairless area in the ajiical jiortion of the discocubital
cell usually more or less outlined by sclerotic thickenings (scleromcs)
and frequently also with one or more scleromes on its surface. This
and its parts are fully described and discussed in my paper on the
Ichneumonidac of the Sauter i^'ormo.san collection ^; and the termi-
nology of the parts is indicated on figure 52 of tho present work.
This structure is characteristic of K/iiros])ilws and absent in tho
Ophion and Thyreodon groups of genera (i)l. 53, fig. 50).

Paramere. — One of tho paired outer valves of the male genitalia
(pi. 55, figs. 84, 85).

Postnervulus. -'Yha combined second and thiril abscissae of the
brachius (pi. 53, fig. 52).

I Prr)C. Knt. Soc. WtwhinRton, vol. 1«, pp. 20-70. 1010.
• Sih. niorpli.Ui. Ent.. vol. 3, p. 302, flR. M, 1M7.

422 PROCEEDINGS OF THE NATIONAL MUSEUM vol.86

Postpectus. — The incurved part of mesosternum anterior to middle
coxae, frequently defined by transverse carinate extensions of the
carinae that flank the coxal foramina (pi. 52, figs. 36-38, 40, 42).

Postpectoral carina. — The carina defining the postpectus.

Pronotal sinus. — A cleft in the humeral angle of pronotum between
the humeral angle itself and the flange by which the pronotum is
attached to mesopleuron. In Ophion and Thyreodon the sinus is
broad, exposing the spiracular sclerite, whereas in Enicospilus it is
narrow and the spiracular sclerite is hidden by the humeral angle
(pi. 55, figs. 72-74).

Spiracular sclerite. — A small, free sclerite between the base of the
tegula and the humeral angle of the pronotum, sometimes concealed
by the latter (pi. 55, figs. 72-74).

Transverse brace. — The combined intercubitus, second abscissa of
cubitus and second recurrent vein. The comparative lengths of
these elements is stated as a ratio, e. g., 1:2:4, the figures represent-
ing the veins in the same order as mentioned above (pi. 53, fig. 52).

Umbo of second tergite. — A median, convex, polished articulating
area at base of second tergite set off laterally by pits and usuaUy
dorsally by a constriction or impression. It is characteristic of the
Ophion group of genera and undeveloped in the Enicospilus and
Thyreodon groups (pi. 56, figs. 94-99).

Tribe OPHIONINI

GROUP CHARACTERS AND GENERIC CHARACTERS

In order to construct a key placing the genera in as near their
natural relationship as possible it has been necessary in most cases
to employ groups of characters rather than single characters. A vast
majority of the species of the tribe belong to the genera Ophion,
Enicospilus, and Thyreodon, and the principal couplets, Nos. 1 and
7, of the key embody the most significant characters distinguishing
typical members of those genera. Although the characters in couplet
7 are stated positively, not all the genera or, indeed, all species of
the basic genera agree with all the characters applying to the cate-
gories in which they belong. Agreement with a majority of the
characters in either alternate determines the subsequent course of a
specimen through the key.

Very few of the characters employed in the key are peculiar to
single genera or even to single groups of genera. The anomalous
hind claws (pi. 55, fig. 77) and frenulum (pi. 54, fig. 56) of Spilophion,
the notched apical front tarsal joint of Ophiogastrella (pi. 55, fig. 75),
and the remarkably long and slender Hgula of Agathophiona (pi. 50,
fig. 28) apparently occur in no other genus, and are of themselves
diagnostic.

GENERIC REVISION OF THE OPIIIOXINI — CUSHMAN 423

Of the characters in the first couplet of the key none by itself is
entirely dia<2:nostic of the members of the Therion group of genera,
with the excei)tiou of the recurved and pointed or narrowly rounded
clypeus. The narrow tapering stigma occurs in Stauropoctoiius and
even in some species of Enicospilus; the strongly reclivous nervellus
with the fracture far above the middle in several genera of the Ophion
group and even in some species of Ophion; the unseparated epipleura
of tergite 2 in Aulophion (a new genus) and in some species of Eni-
cospilus; while the broad, weakly twisted mandible differs only in
being stouter than in Ophion. Only genera possessing all these
characters properly belong in the Thyreodon group.

Of the characters of the Ophion group the gradually narrowed and
weakly twisted mandible occurs also in Spilophion and even in a few
species of Enicospilus, which genus exhibits all degrees of torsion
from that of Ophion nearly to the extreme twisting found in Stauropoc-
tonus; the triangular stigma occurs in Abanchogastra; the lack of the
fenestra in Banchogastra, Pycnophion, Abanchogastra, and Stauropoc-
tonus, all belonging to the Enicospilus group; the curved abscissula
in Spilophion and Abanchogastra.

The strong postpectoral carina of the Enicospilus group occurs in
Aglaophion of the Thyreodon group and in Clistorapha (a new genus)
of the Ophion group, and is absent in Aulophion.

In some genera of both the Ophion and Enicospilus groups the basal
abscissa of the radius is neither straight and unthickcned as in Ophion
nor thickened hi the middle and sinuate as in typical Enicospilus, but
is thickened and curved basally.

The occipital carina, normally complete or very nearly so, is entirely
lacking in a few genera of the Ophion and Enicospilus groups.

KEY TO THE GENERA OF THE OPHIONINI

1. Clyj)cii3 more or less distinctly recurved apically and acute or sharply rounded

(rarely narrowly truncate) (pi. 51, figs. 32-3o); stigma (pi. 63, fig. 50)
extremely narrow, hardly broader than apex of combined costa and sub-
costa and merging gradually into metacarpus, radius nearly at base;
nervellus (pi. 53, fig. 50) strongly reclivous and usually broken distinctly
above middle; epipleura of tergite 2 not distinctly separated from tergite;
mandible (pi. 51, figs. 32-35) very broad and very little twisted (see

couplet 7) 2

Clypeus not as deacribed above or the insect differs by one or more of tlie
other characters 6

2. Postpectoral carina incomplete or absent (pi. 52, figs. 37-39) 4

Posti>ectoral carina complete (pi. 52, fig. 30) 3

3. Radius basally, basal vein, and absci.ssula not thickened; nervellus broken

at about upper third 4. ABlaophion Cameron

liadius ba.sally, basal vein, and abscissula thickened; nervellus broken far
below ujjper third 5. Euryophion Cunieron

424 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

4. Tropi of normal form; propodeimi extending far over hind coxae (pi. 51,

figs. 29, 30) 5

Tropi anthophagous, both labium and maxillae unusually long (pi. 50, fig,
25); propodeum hardly extending beyond insertions of hind coxae (pi. 51,
fig. 31) 3. Rhynchophion Enderlein

5. Joint 2 of maxillary palpus not inflated; ocelli small, distant from eyes; hind

coxae (pi. 51, fig. 29) not or barely extending beyond apex of propodeum.

1. Thyreodon Brull6

Joint 2 of maxillary palpus inflated; ocelli very large, usually touching eyes;

hind coxae (pi. 56, fig. 87) extending beyond apex of propodeum by nearly

half their length 2. Athyreodon Ashmead

6. Hind coxae not extending beyond apex of propodeum; (mandible not strongly

twisted; tergite 3 emarginate at apex; tarsi stout).

6. Orientospilus Morley
Hind coxae extending beyond apex of propodeum 7

7. [Agreement with a majority of the characters in either alternate determines

the subsequent course of a species through the key.] Mandible gradually
narrowed from base to apex, not conspicuously twisted (pi. 60, figs. 17-21);
pronotal sinus (pi. 55, fig. 73) broad, exposing spiracular sclerite; meso-
pleuron with speculum defined below by a more or less distinct obhque
groove; postpectus not defined (pi. 52, fig. 41) ; basal constriction of propo-
deum (pi. 52, fig. 43) more or less distinctly divided into a median and two
lateral foveae; tergite 2 with umbo more or less distinct (pi. 56, figs. 91,
92, 94-99); spiracles of tergite 2 distinctly basad of apical third (pi. 56,
figs. 90-99) ; stigma triangular, tapering from radius to apex, radius at or
beyond basal third (pi. 53, fig. 51); basal abscissa of radius either straight
and unthickened (pi. 53, fig. 51) or thickened and curved basally (pi. 54,
fig. 54), but never sinuate or thickened in middle; apical abscissa of radius
(pi. 53, fig. 51; pi. 54, figs. 54, 55, 57) at most only slightly curved forward
basally; fenestra not at all defined (a hairless area below base of stigma
should not be confused with fenestra) (pi. 53, fig. 51; pi. 54, figs. 54, 55, 59);
absoissula curved basally (pi. 53, fig. 51); nervellus broken from distinctly
below middle to far above middle, upper abscissa perpendicular (pi. 54,

fig. 55a) or recUvous (pi. 54, figs. 54a, 59a); scutellum not margined 8

Mandible aliruptly narrowed between middle and base and so twisted that
teeth are in a plane perpendicular to longitudinalaxis of body, apex very
narrow (pi. 50, figs. 15, 22) ; pronotal sinus (pi. 55, fig. 74) narrow, spiracular
sclerite concealed; mesopleuron without a defined speculum; postpectus
defined, the carina usually complete and closing mesosulcus (pi. 52, fig. 40) ;
basal constriction of propodeum (pi. 52, fig. 44) undivided though some-
times coarsely foveolate; tergite 2 without umbo (pi. 56, figs. 90, 93, 100,
101); spiracles of tergite 2 at or about apical third (pi. 56, figs. 100, 101);
stigma narrow with radius very near base, usually subparallel-sided beyond
radius and rather abruptly tapering apically (pi. 53, fig. 52) ; basal abscissa
of radius m.ore or less thickened to beyond middle and usually more or less
sinuate or undulant (pi. 53, fig. 52; pi. 54, figs. 61, 62; pi. 55, figs. 63-70);
apical abscissa of radius (pi. 53, fig. 52; pi. 54, figs. 53, 56, 58, 60) strongly
curved forward basally, then decurved to apex; fenestra (pi. 53, fig. 52;
pi. 54, figs. 61, 62; pi. 55, figs. 63-70) present, with or without scleromes;
abscissula straight (pi. 53, fig. 52); nervellus broken below (rarely at)
middle, upper abscissa inclivous to perpendicular; scutellum margined,
usually to apex 22

GEXKRIC REVISIOX OF THE OPIIIOXINI — CUSHMAN 425

8. Basal abscissa of radius straight, not thickonod basally 9

Basal al)scissa of radius curved and thickened basally (pi. 54, figs. 54, 55,

59) -....-. 16

9. Mouth parts normal, maxillae and labium not abnormally long 10

Mouth parts anthophagous, both maxilhie and InI)iuM» abnormally long

(pi. 50, fiRs. 25-28) ... 15

10. Stijima triangular, radius at or beyond basal third; apical joint of front tarsus

normal, not emarginatc on outer side 11

Stigma very long and subparalkl-sided for much of its length, radius nearly
at base (pi. 54, fig. 57); apical joint of front tarsus (at least in female)
with a deep, rounded emargination on lower outer margin (pi. 55, fig.
75) 12. Ophiogastrella Brues

11. Scutelhim, metaplcuron, and subalar tubercle not intlated 12

Scutellum, metapleuron, and subalar tubercle inflated (j)!. 52, fig. 45).

11. Australophion Morley

12. Occipital carina distinct and complete (pi. 50, fig. 14) 13

Occipital carina lacking (pi. 50, fig. 12) 10. Alophophion, new genus

13. Discocubitus ascending so close to radius that discocubital coll is much

narrower at that point than at second recurrent, strongly curved both
above and below ramcUus, which is very long and clavatc (pi. 55, fig.

71) 8. Rhopalophion Seyrig

Di.-cocubitus various in form, but not as described above 14

14. Mandible abruptly rectangular at apex, teeth indicated only by a broad,

deep longitudinal furrow; tergite 1 only as long as coxa plus trochanter,
IKJstpetiole as long as petiole; tergite 2 without lateral foveae at base.

9. Apatophion Shestakov
Mandible with distinct teeth apically; tergite 1 longer than coxa plus trochan-
ter, postpetiole shorter than petiole; tergite 2 with lateral foveae at base.

7. Ophion Fabricius

15. Lobes of ligula only moderately elongate, spatulate (pi. 50, fig. 2G) ; ovipositor

long, slender, recurved, sheath fully as long as first segment of abdomen
(pi. 56, fig. 91); temples narrow, receding (])l. 50, fig. 26).

13. Potophion, new genus

Lobes of ligula extremely long and slender, nearly or quite as long as thorax

(pi. 50, fig. 28); ovipositor stout, not exserted (pi. 50. fig. 98); temples very

broad, buccate (pi. 50. figs. 1 1, 28) 14. Agathophiona Westwood

10. Mesosternum with a conspicuous tubercle on each side at npical third.

15. Eremotylus Foerster

Mesosternum without such tubercles 17

17. Kyc8 very .ehallowly emarginate (j)l. 50, fig. 23); clypeus (pi. 50, fig. 23)
very short, concavely truncate at apex, in profile forming a distinct angle
with face, finely punctate; prepectoral carina at most faijitly indicated
(pi. 52, fig. 48); upper abscissa of nervcllus perpendicular (pi. 54, fig. 55a).

IC). Simophion, new genus
Eyea deeply emarginatc (pi. 50, figs. 17-20, 24); clypeus (pi. 50, figs. 17-20,
24) less consj)icuouHly short, truncate or arcuate ut apex, in virtually same
plane as face, usually coarsely punctate; prepectoral carina usually dis-
tinct; upper abHci.s.sa of nervellus usually more r)r less rcclivous, if not the

fracture is above middle f|)l. 54, figs. 54a, 59a) 18

IS. Head very thin, j)rolongefl below eyes, mouth parts anlhopliagous, both
ligula and laoitiia abnormally long (pi. 50, figs. 14, 20, 27); hypopygium
in female cxtenrling far beyond a|)ical tergite, the ovipositor when sheathed
directed dorsoccphalad (pi. 56, fig. 99) 17. Trophophion, new genus

426 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Head neither abnormally thin nor abnormally prolonged, mouth parts normal,
not anthophilous; hypopygium of normal form (pi. 56, figs. 95, 96) 19

19. Nervellus broken at or below middle (pi. 54, fig. 54) ; abdomen not unusually

compressed or heavily sclerotized dorsally, not serrate in profile (pi. 56
fig. 96); clypeus short and very broadly truncate (pi. 50, figs. 17, 18) __ 20
Nervellus broken far above middle (pi. 54, fig. 59); abdomen very strongly
compressed along dorsal margin, serrate in profile, middle tergites being
creased and more heavily sclerotized medially and with prominent apices
(pi. 56, figs. 95, 97); clypeus large and long (pi. 50, figs. 19, 24) 21

20. Postpectus defined, mesosulcus closed (pi. 52, fig. 42),

18. Clistorapha, new genus
Postpectus not defined, mesosulcus open 19. Boethoneura, new genus

21. Apical margin of clypeus very broadly truncate, labrum broadly exposed

(pi. 50, fig. 24); lower margin of mandible evenly curved (pi. 50, fig. 24);

legs and antennae stout 20. Genophion Felt

Apical margin of clypeus arcuate or, rarely, narrowly truncate medially,
labrum narrowly exposed (pi. 50, fig. 19) ; lower margin of mandible bent
at the middle, straight before and beyond bend (pi. 50, fig. 19) ; legs and
antennae slender 21. Chilophion, new genus

22. Occipital carina absent (pi. 50, fig. 13) ; mandible very small, so twisted that

upper tooth appears to be on inner margin (pi. 50, fig. 15); scutellum not
margined laterally; mesopleuron with a distinct longitudinal furrow from
fovea to top of prepectus (pi. 52, fig. 49) ; stigma extremely narrow, tapering
from radius, which is very near base (pi. 54, figs. 53, 58) ; basal abscissa of
radius thick and curved basally, apical abscissa strongly curved forward at
base (pi. 54, figs. 53, 58), abscissula slightly thickened and weakly curved
basally; apical joint of trochanter in middle and hind leg with a sharp,

decurved tooth on outer apical margin (pi. 55, fig. 83) 23

Occipital carina present; differing also from most of the other characters
above 24

23. Postpectus completely defined; nervulus postf ureal; postnervulus broken

shortly above middle; nervellus broken at about middle; epipleura of
segment 2 distinctly separated from tergite throughout.

22. Stauropoctonus Brauns

Postpectus not defined; nervulus antefurcal to interstitial; postnervulus

broken far above middle; nervellus broken far below middle; epipleura of

segment 2 separated from tergite only basad of spiracles and there weakly

so 23. Aulophion, new genus

24. Spiracles of tergite 2 at, before, or somewhat behind middle, but always

basad of apical third (pi. 56, figs. 90, 93) ; stigma elongate triangular; basal
abscissa of radius neither thickened nor sinuate; no fenestra; second discoidal
cell pointed or very narrow at base; first abdominal segment stout (pi. 56,

figs. 90, 93); ocelli small 25

Not agreeing entirely with above 26

25. Postpectoral carina complete, mesosulcus closed; first abdominal segment

very short and broad, sternite not nearly reaching spiracles, tergite 2 with
spiracles distinctly basad of middle; ovipositor not exserted (pi. 56, fig. 93).

24. Banchogastra Ashmead

Postpectoral carina obsolete, mesosulcus open; metapleuron tumid; first

abdominal segment more slender, sternite reaching spiracles, tergite 2 with

spiracles at or beyond middle; ovipositor strongly exserted (pi. 56, fig. 90).

25. Pycnophion Ashmead

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 427

26. Claw of hind tarsus with poctcn extending beyond apex (pi. 55, fig. 77);

mandible gradually narrowed from base to apex; penultimate hook of
frenulum usually very different from others (pi. 5-1, fig. 56c); (abscissula
strongly curved basally [pi. 54, fig. 56b]; a hairless area below stigma but

no well-defined fenestra (pi. 51, fig. 56]) 26. Spilophion Cameron

Claw of hind tarsus normal (pi. 55, fig. 80); mandible usually abruptly
narrowed ; penultimate hook of frenulum normal 27

27. Stigma broad with radius near middle, basal abscissa of radius not thickened;

no fenestra (pi. 54, fig. CO) ; abscissula curved, nervellus broken at middle,

upper ab.-^ci.-^sa j)erpent!icular 27. Abanchogastra Perkins

Stigma narrow with radius near base; basal abscissa of radius more or less
thickened; fenestra distinct, with or without scleromes (pi. 53, fig. 52; pi.
54, figs. 61, 62; pi. 55, figs. 63-70); abscissula straight; nervellus broken
below middle, upper abscissa usually inclivous. 28. Enicospilus Stephens

1. Genus THYREODON BrulI6

Plate 49, Figure 1; Plate 51, Figures 29. 32; Plate 52, Figures 38, 47;
Plate 53, Figure 50; Plate 55, Figures 72, 84; Plate 50, Figure 86

Thyreodon Brull6, Histoire naturelle des in.sectes, Hym6nopt^rcs, vol. 4, p. 150,
pi. 42, fig. 3, 1846. — SzfiPLiGETi, in Wytsmaa, Genera inscctorum, fasc. 34,
p. 25, 1905.— Hooker, Trans. Amer. Ent. Soc, vol. 38, p. 106, 1912.— Mor-
let, a revision of the Ichneumonidae based on the collection in the British
Museum (Natural History), pt. 1, p. 7, 1912. — Enderlein, Zool. Anz., vol.
39, p. 626, figs. 1-4, 6, 1912. [Genotype: Thyreodon cyaneua Brulld. By
designation of Hooker, 1912.]

Oleler Shestakov, Konowia, vol. 5, p. 259, 1926. [Genotype: {Oleter selenadion
Shostakov) = Thyreodon lalicindua Cresson.] Monobasic.

A genus of very large species, exclusively American and largely
tropical in distribution. I have listed above only the more significant
contributions to literature on the genus.

Head (pi. 51, fig. 32): Narrower than thorax; temjiles buccate to
receding; occipital carina usually not reaching hypostomal carina,
genae broad below; eyes rather small, moderately emarginate opposite
antennae; malar space half or more as long as basal width of mandible;
ocelli very small, removed from the eye.s by more than their diameter;
frons with deep antennal scrobes bounded medially by a prominent
interantcrmal ridge and laterally by carinae along the eyes, the
carinac usiially turning away from the eye margins above toward the
lateral ocelli; face moderatel}' convex; clyj)eus not distinctly separated,
long, pointed at apex and in profile with apex recurved, foveae very
large and deep; labrum concealed; mandible broad and short, strong]}''
narrowed from base to apex, not strongly twisted, teeth subequal;
tropin of normal length; second joint of maxillary palpus flat, tri-
angular; antenna shorter than body, rather stout, subsetiform, scapo
very short and squarely truncate at apex.

Thorax: Stout; pronotum with a deep transverse foveolate furrow
dorsally, bounded posteriorly by a high ridge and anteriorly by the

428 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

recurved margin; pronotal sinus (pi. 55, fig. 72) broad, exposing the
spiracular sclerite; raesoscutum precipitous anteriorly, notaulices more
or less distinct, broad, usually foveolate or rugose; scutellum short,
precipitous apically, basal fovea and lateral areas very deep, lateral
carina high, extending at most to top of apical slope; prepectoral
carina (pi. 52, figs. 38, 47) ascending well up on mesopleura, where it
frequently bends abruptly toward, and usually reaches anterior mar-
gin; postpectoral carina (pi. 52, fig. 38) distinct only medially, where
it is sometimes represented only by a tubercle on each side of meso-
sulcus; metapleuron (pi. 51, fig. 29) small, triangular, flat, deeply inset;
propodeum (pi. 51, fig. 29) bulbous, without carinae, precipitous
apically, basal constriction (pi. 51, fig. 29) very deep and narrow,
spiracles very long, situated near bottom of constriction. Wings
(pi. 53, fig. 50) densely hairy, with neither a hairless area below base
of stigma nor a fenestra; stigma extremely narrow, no broader than
combined costa and subcosta, radius very close to base, stigma very
gradually tapering from radius and merging imperceptibly with
metacarpus; radial cell very narrow; basal abscissa of radius neither
strongly thickened nor curved, apical abscissa a long sigmoid curve;
basal abscissa of radius, intercubitus and apical abscissa of cubitus
forming a nearly straight line, only slightly angled at the nodes; basal
vein straight or slightly decurved at lower end; discocubitus curved,
without ramellus; intercubitus as long as or longer than second
abscissa of cubitus; discoidal cell not or barely as long as brachial cell;
lower apical angle acute; postnervulus broken above middle; nervellus
from shortly antef ureal to shortly postf ureal; frenulum with many
widely separated and frequently unevenly spaced hooks and extending
from near radius about halfway to apex of wing; abscissula straight or
weakly curved, cubitella usually not nearly reaching margin of wing;
nervellus strongly reclivous, broken far above middle, upper abscissa
strongly inclivous. Legs short, hind leg not or barely as long as
abdomen; hind coxa (pi. 51, fig. 29) not extending beyond apex of
propodeum; front and middle coxae each with a more or less tubercle-
like extension below articulation of trochanter; claws moderately
curved, pectination short and dense.

Abdomen (pi. 56, fig. 86): Alore than twice as long as head and
thorax, strongly compressed beyond first segment; first segment
slender, spiracle at about apical fifth, petiole usually compressed,
postpetiole short, convex above; tergite 2 much shorter than tergite
1 and longer than 3, without an umbo, spiracles beyond middle but
before apical third, gastrocoeli nearer to spiracles than to base,
epipleura not at all separated from tergite; sternite 2 reaching to about
middle of tergite 2, sternite 3 not or barely reaching apex of tergite 2;
tergite 3 with a distinct, subcircular gastrocoelus-like area at extreme

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 429

base on each side: midtlle terprites not'ciiinrixinate medially; ovipositor
not exsertcd; male hypopygium with apex arcuate or medially emar-
ginate, paramere (pi. 55, iig. 84) acutely pointed at apex and fre-
quently suhapically annulate on lower margin.

This genus, containing about 30 species, is divisible into two fairly
distinct groups, one mcluding the genotypes of both Thyreodon and
Older, characterized by small head, sloping temples, more distinct
notaulices, frequently with prominent rugae at their anterior ends,
and slenderer habitus; and the other consisting of T. atricolor (Olivier),
grandis Cresson, fernaldi Hooker, and their relatives with larger head,
broader temples, obsolescent notauhces and stouter habitus. These
appear to be all characters of specific rather than generic significance.

2. Genus ATHYREODON Ashmead

Plate 51, Figure 33; Plate 52, Figure 37; Plate 56, Figure 87

Athyreodon Ashmead, Proc. U. S. Nat. Mu?., vol. 23, p. 87, 1900.— Hooker,
Trans. Amer. Ent. Soc, vol. 38, p. 100, 1912. [Genotype: (Athyreodon
thorarirus A?hme&d) = Athyreodon atriventris (Cresson), according to Hooker,
1912.] Monobasic.

Tipulophion Krieciibaumer, Zeitschr. Hym. Dip., xn\. 1, p. 75, 1901. — Schulz,
ZeiLschr. Hym. Dip., vol. 3, p. 252, 1903; Spolia hymenopterologica, p. 97,
1006. [Genotype: {Tipulophion gigas Krieciibaumer) = yl//!J/reof/on atri-
ventris (Cresson), according to Hooker, 1912.] Monobasic.

Macrophion Sz^pligeti, in Wytsnum, Genera insectorum, fasc. 34, p. 32, 1905. —
ScHMiKDEKNECHT, Opuscula iclincumonologica, fasc. 18, p. 1420, 1908. —
MoRLEY, A revision of the Ichncuraonidae based on the collection in the
British Museum (Natural History), pt. 1, p. 14, 1912, [Genotype: {Macro-
phion ornalus Sz^pligeti) = (T/ij/rcor/on) Macrophion grenadensis (Ashmead),
according to .Morley, 1912. By designation of Viercck, Proc. U. S. Nat. Mus.,
vol. 42, p. 640, 1912.]

Hooker seems to have been correct in synonymizing the genotypes
of Athyreodon and Tipulophwn with atriventris (Cresson), and there
appears little doubt that both Macrophion ornatus Sz^pligeti and
Thyreodon grenadensis arc also atriventris.

This genus, also Neotropical, is closely related to Thyreodon, from
which, in the species that resemble Thyreodon most clo.sely, it is rather
weakly separated. With the acquisition of more material it may be
found neces.sary to synonymize it with Thyreodon, but on the basis of
the 9 or 10 species available to me it is distinguishable as follows:

Head (pi. 51, fig. 3.*]): Temples usually very narrow and genae
narrow below, rarely both broad; eyes larger, more deeply emarginate;
malar space usually very short, rarely as in Thyreodon', ocelli large,
ocellocular space fref|uently o})solele and rarely as long as diameter of
an ocellus; antennal scrobes small and poorly defined; clypeus sharply
rounded but not angulate at apex; second joint of maxillary palpus
inflated.

430 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

Thorax (pi. 52, fig. 37) : Propodeum usually not distinctly inflated,
not reaching apex of hind coxa, basal constriction less deep and meta-
pleuron not so deeply inset, rarely approaching the condition in
Thyreodon. Wings with discoidal cell usually longer than, rarely equal
in length to, brachial cell, lower apical angle nearly or quite a right
angle; frenulum shorter than in Thyreodon, hooks fewer and more
closely spaced; cubitclla reaching wing margin; nervellus frequently
not broken, the brachiella originating at its upper end. Hind leg longer
than abdomen, coxa extending beyond apex of propodeum by nearly
half the length of coxa; front coxa only slightly, middle coxa not at all
tuberculato apically.

Abdomen (pi. 56, fig. 87): Petiole usually terete; sternite 2 not
reaching middle of tergite 2, sternite 3 not reachmg apex of tergite 2.

The critical characters of this genus are the inflated second joint of
the maxillary palpus, the large ocelli and the extension of the hind
coxae beyond the propodeum.

3. Genus RHYNCHOPHION Enderlein

Plate 50, Figure 25; Plate 51, Figures 31, 35; Plate 52, Figure 39; Plate 55,
Figures 76, 85; Plate 66, Figure 88

Rhynchophion Enderlein, Zool. Anz., vol. 39, p. 630, figs. 5, 7, 8, 1912. [Geno-
type: Rhynchophion odontandroplax Enderlein.] Monobasic.

A third American genus of the Thyreodcn group, anomalous in the
anthophagous trophi and the entire lack of the postpectoral carina,
and differing from Thyreodon otherwise as follows:

Head (pi. 50, fig. 25; pi. 51, fig. 35): Broader across temples than
across eyes; eyes smaller than in Thyreodon and only arcuately emar-
ginate, malar space longer; antennal scrobes shallow and not cari-
nately defined; mandible longer, subparallel-sided, lower tooth the
larger; trophi anthophagous, both labium and maxillae unusually
long; second joint of maxillary palpus slender, terete; scape some-
what obliquely truncate.

Thorax: Pronotum with only a shallow groove dorsally, posterior
and anterior margins not at all elevated; notaulices obsolete; meso-
pleuron with a broad longitudinal impression at level of fovea; pre-
pectoral carina (pi. 52, fig. 39) developed only ventrally; postpectoral
carina not at aU indicated medially; metapleuron (pi. 51, fig. 31)
large, subovate, convex; propodeum (pi. 51, fig. 31) not at all inflated,
basal construction shallow medially, spiracles situated behind con-
striction. Wings very densely clothed with long, recumbent hair;
lower apical angle of discoidal cell right or slightly obtuse; frenulum
short, hooks closely and evenly spaced. Legs rather slender, hind leg
much longer than abdomen; front and middle coxae not at all tuber-

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 431

culate apically; hind coxa (pi. 51, fig. 31) extending nearly its entire
dorsal length beyond apex of propodeum.

Abdomen (pi. 56, fig. 88): Not or barely twice as long as combined
head and thorax, veiy strongly compressed and, in side view, very
deep; firet segment rather stout, slightly decurved, postpetiole not
bulbous, spiracles shortly behind apical third; tergite 2 shorter than 3,
spiracles at about middle, gastrocoeli near base; sternite 2 reaching
apex of tergite 1, sternite 3 to apex of tergite 2; male hypopygium
broad, apex deeply emarginate on each side of middle and with a
median acute tooth, paramere (pi. 55, fig. 85) rounded at apex.

This genus apparently includes comparatively few species, of which
only two besides the genotype are described. It is largely tropical
in its range but extends into the southwestern part of the United States.

RHYNCHOPHION ODONTANDROPLAX Endcrlein

lihynchophion odontandroplax Enderlein', Zool. Anz., vol. 39, p. G30, figs. 5, 7,
8, 1912.

I have identified as this species a female from Sao Paulo, Brazil, in
the collection of Hcnrj' K. and Marjoric C. Townes, which agrees
almost perfectly, except in sexual characters, with the original descrip-
tion. The clypeus in this species, while of the same type as in the other
species, is somewhat truncate medially.

RHYNCHOPHION FLAMMIPENNIS (Ashmead). new combination

Thyreodon Jlammipennis Ashmead, Proc. California Acad. Sci., ser. 2, vol. 4, p. 125,
1894. — MoRLEY, A revision of the Ichneutnonidae based on the collection
in the British Museum (Natural History), pt. 1, p. 12, 1912. — Hooker,
Trans. Amer. Ent. Soc, vol. 38, p. 129, 1912.

The range of this species, originally described from Baja California,
Mexico, is shown to extend from Ecuador to the southwestern United
States by the following specimens in the United States National
Museum: One male, Po.sorja, Ecuador, F. Campos K.; one female,
Tlahuililo, Durango, Mexico, A. Busck; one female, two males,
Douglas, Ariz., August 22, 1932, and August 2, 1933, W. W. Jones,
and one female, August 20, 1932, collector unknown; one female,
Tucson, Ariz.; and one female, northern New Mexico, A. O. \V'^eese.

RHYNCHOPHION LIGULIFEB (Morlcy), now combination

Thyreodon ligulifer Morley, A revision of the Ichneunionidao ba.sod on the col-
lection in the British Museum (Natural History), pi. 1, p. '.», I'.d'J.

Morley's descriplioii of (he elongates troj)hi, the less iiilliilcd and less
deeply constricted propodeum, and the more prominent coxae leaves no
doubt that this species belongs in Rhynchop/uon. Morlcy overlooked
the same characters in Jlammipennis, which he discussed on a later
page of the same work.

432 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

4. Genus AGLAOPHION Cameron

Plate 51, Figures 30, 34; Plate 52, Figures 36, 46; Plate 56, Figure 89

IDidyonotus Kreichbaumer, Zool. Jahrb. Syst., vol. S, p. 197, 1894. [Genotype:
Ophion (Diclyonotus) melanarius Kreichbaumer.] Monobasic.

Aglaophion Cameron, Journ. Straits Branch Roy. Asiatic Soc, No. 39, p. 131,
1903. — MoRLEY, A revision of the Ichneumonidae based on the collection
in the British Museum (Natural History), pt. 1, p. 15, 1912. [Genotype:
Aglaophion flavinervis Cameron.] Monobasic.

Hybopleurax Enderlein, Zool. Anz., vol. 39, p. 624, 1912. [Genotype: Hybopleu-
rax sumatranum Enderlein.] Monobasic. New synonymy.

Coracophion Shestakov, Konowia, vol. 5, p. 261, 1926. [Genotype: Coracophion
manganicolor Shestakov.] Monobasic. New synonymy.

This genus is the Old World representative of the Thyreodon group.
It is restricted in distribution to the eastern part of the Palearctic
region and the Oriental region.

I have not seen the genotype of Aglaophion but believe I am correct
in transferring it to the genus Thyreodon purpurascens Smith, as which
I have identified certain specimens in the United States National
Museum from China and Japan. As Hybopleurax sumatranum
Enderlein I have identified a specimen in the United States National
Museum, and on this basis synonymize Hybopleurax with Aglaophion.
These two are the only described species I know, aside from the
genotype, that are referable to the genus. The possible synonymy
of this genus with Dictyonotus Kriechbauxner is discussed under the
latter genus.

From the other genera of the Thyreodon group tliis genus may at
once be distinguished by its complete and high postpectoral carina
(pi. 52, fig. 36) and deeply emarginato tergites. From the above
description of Thyreodon it also differs as follows:

Head (pi. 51, fig. 34) : About as wide as thorax; eyes more shallowly
emarginate; malar space nearly as long as basal width of mandible;
antennal scrobes shallow and less distinctly defined; clypeus rather
distinctly separated, recurved but rounded at apex; second joint of
maxillary palpus clavate.

Thorax: Pronotum not deeply grooved dorsally, neither anterior nor
posterior margin prominent; mesoscutum hardly precipitous anteriorly,
notaulices obsolete; mesopleuron (pi. 52, fig. 46) with a longitudinal
groove or impression at top of prepectus dividing the pleuron into
approximately equal upper and lower areas; sternaulices obsolete;
metapleuron strongly convex, frequently tuberculate; propodeum
(pi. 51, fig. 30) not bulbous, basal constriction only moderatel3^ deep,
spiracles situated behind constriction. Wings with lower apical angle
of second discoidal cell right or slightly obtuse; frenulum much shorter
and with comparatively few (about 10) hooks; cubitella nearly reach-
ing margin of wing; nervellus broken at or near upper third. Legs

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 433

lonirer, hind Ice; nuich lonp-er than abdomen ; hind coxa cxtcndinp: about
half its dorsal length beyond propoilcuni; front and middle coxae not
produced beyond articulations of trochanters; claws more strongly
curved apically, with stout, well-separated teeth.

Abdomen (pi. oG, fig. 89): Barely twice as long as head and thorax;
spiracles of first segment at about apical fourth ; petiole terete; tcrgites
3-7 emarginate medially.

AGLAOPHION FLAVINERVIS Cameron

As stated above, I have not seen this species, but some notes on the
type supplied by Mr. Perkins should aid materially in its identifica-
tion:

"Occipital carina strong and complete; ocular carina on frons dis-
tinctly raised; apical margin of clypeus outwardly reflexed; numdible
sinular to that of Tlixjreodon; pronotal sinus deeply emarginate; notauli
absent; pit before postscutellum distinct; mesopleuron with a broad,
deep, smooth, transverse furrow; stcrnauli very weakly indicated
anteriorly; postpectoral carina very strong and complete; metapleuron
weakly intumescent; propodeum about as long as broad, basal con-
striction very narrow, deep, partially interrupted (by becoming
shallower) laterad ; stigma as in EnicospiliLs; radial cell narrow and in
form like that of Thyreodon, but a little broader, apically sharply
curved to the costa; fenestra absent; basal abscissa of radius of hind
wing almost straight; second tergite without umbo, spiracles beyond
middle; tergites 3-5 deeply incised dorsally."

AGLAOPHION PURPURASCENS (Smith), new combination

TItyreodon purpurascens Smith, Trans. Ent. Soc. London, p. 395, 1874. — Schclz,
S];olia hynicnoptorologica, p. 98, 1906. — Mohley, A revision of the IchntMi-
monidac based on the collection in the British Museum (Natural History),
pt. 1, p. 10, 1912. — UcHiDA, Journ. Fac. A^r. Hokkaido Imp. Univ., vol. 21,
p. 200, 1928; Iii.secta Mat,suiiiurana, vol. 14. pp. 43, 124, 1940.

Ophion metallicum Radoszkowski, HoraeSoc. Ent. Ros.sica, vol. 21, p. 433, 1887. —
ScHUi.z, .Spoiia hymenoptfroiogica, p. 97, I90G.

?lJictyonolu8 melanarius Kriechbaimkr, Zool. Jahrb. Syst., vol. 8, p. 197, 1894.

Coracophion mangani color Siiestakov, Konowia, vol. 5, p. 2G1, 192(3.

Uchida appears to be correct in synonymizing Ophion metallicum
Radoszkowski and Coracophion manqanlcolor Shestakov with jmr-
purcLscens. The doubtful .synonymy of Dictyonotus melanariiLs Kriech-
biumier is mine; for discussion of this see under Dictyonotus.

As this species I have identified the following female specimens in
the United States National Mu.seum: One, Hashimoto, Japan, June
26, 1928, T. U. Gardner; two, (jifu, Japan, Y. Nawa;one, Ruling,
Kiangsi, China, N. Gist Gee; and one without locality.

725.'>04 — 47 2

434 PROCEEDINGS OF THE NATIONAL MUSEUM vol. oa

J. F. Perkins examined the type and made the following notes
comparing the species with Aglaophion flavinervis: "Differs in having
the mandible in front of a line joining the base anteriorly to the V
between the teeth, strongly depressed; clypeus conspicuously, though
not strongly, outwardly reflexed in the apical one-third (at an angle
of about 150°) ; ocular carina of the same form as in Euryophion, but
the weak groove to the ocelli is not interrupted; propodeum and meta-
pleura of the same form as in Aglaophion, but a httle shorter, tergites
3-5 weakly excised apically."

AGLAOPHION SUMATRANUM (Enderlein), new combination

Hybopleurax sumatramim Endeelein, Zool. Anz., vol. 39, p. 625, 1912.

A female captured by H. M. Pendlebury at Nakon Sri Tamarat,

Khao Luang, Siam, on March 30, 1922, agrees perfectly with the

description.

5. Genus EURYOPHION Cameron

Euryophion Cameron, Ann. South African Mus., vol. 5, p. 83, 1906. — Morley,
A revision of the Ichneumonidae based on the collection in the British
Museum (Natural History), pt. 1, p. 15, 1912. [Genotype: Euryophion
nigripennis Cameron.] Monobasic.

This African genus is unknown to me, and I should have been
unable to place it without the assistance of J. F. Perkins. With this
assistance I place it, I think satisfactorily, in the Thyreodon group,
where it appears to be the South African analog of Aglaophion. The
following description is compounded of the characters given by
Cameron and Morley together with those furnished by Mr. Perkins:

Large insects, exceeding 25 mm. in length, entirely black wings.

Head: Strongly swollen behind eyes; occipital carina with a median
notch directed toward the neck; frons margined by carinae both be-
low and at sides, carina at side of frons extraordinarily raised and
continued to posterior ocellus by a groove which runs from eye to
front of posterior ocellus, the carina and groove interrupted shortly
before ocellus by a short, deep, transverse fiU"row; clypeus medially
reflexed and truncate; mandibles broad, not conspicuously twisted;
eyes parallel and not emarginate; ocelli small; mandibles short and
broad, only slightly twisted; antenna thick, tapering and about as
long as forewing.

Thorax: Mesoscutum with a shallow median longitudinal furrow,
notauHces obsolete; scutellum not margined; postscutellum elevated;
mesosternum flat, sternauUces shallow anteriorly, deep posteriorly;
postpectoral carina complete, but difficult to see ventrally as it
runs close to the coxae; metapleuron with a large tubercle in middle;
propodeum not longer than broad, not conspicuously intumescent

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 435

basally, basal constriction very narrow. Wings with stipiia and
radial coll very narrow, the sti<;nia nier<2;ino: imperceptibly with meta-
carpus, radius nearly at base; basal vein, basal abscissa of radius and
abscissula somewhat thickened, apical abscissa of radius bent sharply
forward apically. Legs with hind coxa extending distinctly beyond
apex of propodeum; tarsal joints 1-4 compressed.

Abdomen: First segment short and broad; tergite 2 deeper than
long, with epipleura extremely broad, spiracles shortly behind middle.

In making his notes on this genus Mr. Perkins did not have the
genotype before him, but based his observations on the tj'pes of
Eunjophion magnijicus Morley and E. superhus Morley, which he
believed properly referred to the genus.

6. Genus ORIENTOSPILUS Morley

Orientospilus Morley, A revision of tho Iclineumonidae based on the collection
in the British Museum (Natural History), pt. 1, p. 6, 1912. [Genotype:
Orientospilus individuus Morley. By designation of Vicreck, U. S. Nat.
Mus. Bull. 83, p. 107, 1914.]

This genus is unkno\vn to me. As described, it presents a confusing
combination of characters. The short, tumid propodeum overlying
the hind coxae suggests Thyreodon, the emarginate third tergite
Aglaophion, and the fenestra Enicospilus; but the character that
seems to ally it with any genus distinguishes it from each of the others.

Mr. Perkins examined the genotype and sent me the followhig notes:

"Mandible slightly twisted, but very strongly tapering, upper
tooth much longer than lower; clypeus broadly, shallowly emarginate
apically; pronotal sinus broad; speculum not set off; postpectoral
carina absent; sciitellum margined; basal constrictioji of propodeum
deep and undivided, intumescent behind this (similar to Thyreodon,
])ut laterally less sharply constricted at base); umbo present; spiracles
of second tergite (which has extremely narrow epipleura) a little
beyond middle; stigma as in Enicospilus; basal abscissa of riidius
thickened basally and sharply curved; apical abscissa weakly, evenly
arched towards the anterior margin, fenestra absent; abscissula sharply
curved at base."

It is particularly noteworthy that Perkins' observation on the lack
of the fenestra is diametrically opposite that of the author of tho
genus, who described the fenestra as present and with scleromes.

Enicospilvs reticvlnfus Cameron, 1899 (not 1902), which Morley
refers to this genus difTers in nearly eveiy chnrocter from the original
description of tiie gfinis. As I liiive identified it tiiis species is an
Enicospilus, anomalous only in its broad and weukly twisted mandibles.
It is closely allied to E. flavoplagiatus Cusiinuin.

436 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

7. Genus OPHION Fabricius ^

Plate 49, Figure 3; Plate 50, Figure 21; Plate 52, Figures 41, 43; Plate 53,
Figure 51; Plate 55, Figures 73, 81; Plate 56, Figure 94

Ophion Fabricius, Entoraologia systematica . . ,, Suppl., pp. 210, 235, 1798;

Systenia Piezatorum, p. 130, 1804. [Genotype: Ichneumon luteus Linnaeus.

By designation of Curtis, British entomology, vol. 13, p. 600, 1836.]
Paniscus Schrank, Fauna Boica, p. 316, 1802. [Genotype: Ichneumon luieus

Linnaeus.] Monobasic.
Neophion Morley, A revision of the Ichneumonidae based on the collection in

the British Museum (Natural History), pt. 1, p. 30, 1912. [Genotype:

Neophion crassus Morley. By designation of Viereck, U. S. Nat. Mus.

Bull. 83, p. 100, 1914.J

To give a bibliography of this genus would be to list the works,
dealing with the genus, of virtually all the more prominent students
of the Ichneumonidae since the time of Gravenhorst. Suffice it here
to cite a few of the more useful works for the identification of species:
European species, Schmiedeknecht, Opuscula iclmeumonologica, fasc.
4, pp. 1334-1348, 1908, and supplement, fasc. 24, pp. 24-47, 1935.
American species, Hooker, Trans. Amer. Ent. Soc, vol. 38, pp. 21-50,
1912. Species of the world as represented in the British Museum of
Natural History, Morley, A revision of the Ichneumonidae based on
the collection in the British Museum (Natural History), pt. 1, pp.
53-66, 1912. Eastern Asiatic species, Uchida, Journ. Fac. Agr.
Hokkaido Imp. Univ., vol. 21, pt. 2, pp. 204-211, 1928. New Neo-
tropical species, Szepligeti, Ann. Mus. Nat. Hungarici, vol. 4, p. 133,
1906. Central American species, Cameron, Biologia Centrali-
Americana, Hymenoptera, vol. 1, p. 293, 1906. The keys in these
works fall far short of including all the described species, and recourse
must be had to the many scattered descriptions of species.

Under strict interpretation of the International Code of Zoological
Nomenclature, Paniscus Schrank is isogenotypic, and therefore
synonymous, with Ophion Fabricius. Believing, however, that
stability of the nomenclature of the genera mvolved would be best
served by the preservation of the names in their traditional senses,
I some years ago * attempted, I fear rather unsuccessfully, to show
that Schrank's conception of Paniscus was based on the excellent
figures in DeGeer's "Memoires pour Servir a I'Histoire des Insectes,"
1771, which almost certainly illustrate the insect long knowTi as
Paniscus cephalotes Holmgren. The alternative was to synonymize
Paniscus with Ophion and substitute for Paniscus in the traditional

' Since this synonymy was written Dr. Henry K. Townes has published his monumental "A Catalogue
and Reclassification of the Nearctic Ichneumonidae" (Mem. Amer. Ent. Soc, No. 11, 1944^5), on p. 730 of
which, in addition to the synonyms listed here, he also synonymizes Oenophion Felt and Ophiogastrella
Brues. In my opinion the venational characters by which I have separated these two genera from Ophion
are of much more than specific significance.

* Proc. U. S. Nat. Mus., vol. 64, art. 20, p. 21, 1924.

GENERIC REVISION OF THE OPIIIONIXI — CUSHMAN 437

sense N'fielia Gray. The Inst-montioned iinme is monobasic, with
Paniscus inquinatiLS Gravonhorst as type, anci was orii!:inally defined
1)V the sinplo character of the number of liooks in the frenulum. This
species has not otherwise been recoirnized since its description in 1829,
and the correctness of the identification of Gray's specimen is ex-
tremely doubtful. Townes, in 1938,* synonymized Paniscus Schrank
with Ophion and used A^etilia Gray for Paniscns of authors. Although
1 fi'cl that this action was too precipitate and will perhaps not receive
the support and following of a majority of the contemporary specialists
on the Ichneumonidae, his action seems to have blocked the preserva-
tion of Paniscus in the sense sanctioned by well over a hundred years
of usage, and I shall follow him in the use of Netelia. There is,
however, still the threat of instability in the nomenclature of the genus
if the type of Paniscus inquinatus Gravenhoj'st is still in existence,
for there is no assurance that examination of the type will show it to
be congeneric with the species now to be known as Netelia, especially
those of the subgenus Netelia as identified by Townes.

Ophion is a genus of world-wide distribution and contains many
hundreds of species differing widely among themselves but each
possessing virtually all the characters in the foregoing key that apply
to the Ophion group of genera. In this somewhat heterogeneous
assemblage are included a number of groups of sj)ecies which are more
or less well distinguished from the general mass of more typical forms
but are not, I think, generically distinct. Such species as bifoveolatus
Brulld, colnradensis (Felt), slossonae Davis, and others in the North
American fauna are representatives of such groups, and one is perhaps
justified in suspecting that some of the generic names that have been
proposed apply to such groups. It may be possible and advisal)le to
employ some of these names, and others not yet proposed, in the
subgeneric sense.

The synonymy of Neophion Morley is on the authority of J. F.
Perkins, who examined the tyi)e of A'^. crassus Morley and pronounced
it a species of Ophion.

The genus is characterized as follows:

Head (pi. 50, fig. 21): Various in form, but ahva^'s with trophi
normal, with none of the elements greatly elongate; mandible gradually
narrowed from base to apex and not conspicuously twisted, teeth
equal or lower tooth the longer; clypeus truncate or broadly arcuate
at apex; occipital carina distinct.

Thorax: Pronotal sinus (i)l. /jo, fig. 73) broad, exposing spirncular
sclcrite; notaulices more or less distinct at least anteriorly; meso-
pleuron with speculum distinct; prcpectoral carina (pi. 52, fig. 41)

• Lloy.lla, vol. I. [.p. Ifi8-231, 1038.

438 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

present; postpectus not defined; scutelliim, subalar tubercles, and
metapleura more or less convex but not inflated; scutellum rarely
completely margined; propodeum (pi. 52, fig. 43) excarinate to com-
pletely areolated, not produced over hind coxae, basal constriction
neither especially deep nor especially long, divided into a median and
two lateral foveae; frenum much shorter medially than basal area.
Wings (pi. 53, fig. 51) without fenestra or scleromes, though fre-
quently with a hairless area below base of stigma; stigma elongately
triangular, tapering evenly from radius to apex, radius at or beyond
basal third; basal vein and basal portion of discocubitus usually
convergent toward anterior margin; basal abscissa of radius straight,
not thickened basally, apical abscissa sinuous or straight, rarely curving
forward at base; abscissula curved basally; nervellus broken from
slightly below to distinctly above middle, upper abscissa perpendic-
ular. Legs stout to very slender; apical joint of front tarsus and all
claws (pi. 55, fig. 81) normal in form.

Abdomen (pi. 56, fig. 94) : Tergite 2 with umbo distinct, spiracles at
or near middle, rarely far before or distinctly behind middle, epipleura
distinctly separated throughout; ovipositor not exserted.

8. Genus RHOPALOPHION Seyrig

Plate 55, Figure 71

Rhopalophion Seyrig, Mus. Nat. Hist. Nat., Mission Scientifique de I'Omo,
vol. 3, Zool., fasc. 18, Hym. 2, Ichneumonidae, p. 49, 1935. [Genotype:
Rhopalophion curvus Seyrig.] Monobasic and designated.

Described from four African species, only one of which is named.
This genus is, I think, not distinct from Ophion, though lacking a
specimen I prefer not to synonymize it. The most conspicuous char-
acter and the one on which the genus was principally founded, the
course of the discocubitus (pi. 55, fig. 71), is very doubtfully of generic
significance. The one character mentioned that deters me from syn-
onymizing the genus is described as follows: *'2e abscisse radiale des
ailes posterieures droites ou tres legerement sinuee (contrairement au
genre Ophion oil elle est fortement recourbee a la base)." It seems
probable, since the second abscissa of radiella in Ophion is always
straight, that the "2" is a typographical error for "l" and that the
description applies to the abscissula, which, in Ophion, is more or less
distinctly curved basally. The form of the mandible, as described,
and of the stigma and basal abscissa of radius and the lack of the
fenestra as shown in the figure of the wing, indicate close relationship
to Ophion.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 439

9. Genus APATOPHION Shostakov »•

Apatophion Shestakov, Konowia, vol. 5, p. 2G2, 192C. — Mkyer, Konowia, vol.
IC, p. IS, 1937. (Genotype: Apatophion mirsa Shestakov.) Monobasic.

Despite the pniicity of characters pven in the oriidnal description
that would aid in placiuc; tliis genus in the foreccoinf^ key, it seems evi-
dent that it belonp^s in the Ophion group and very close to Ophion.
In fact, Meyer treats it as a subgenus of Ophion, distinguishing it only
by the character of the postpetiole being as long as the petiole. Meyer
is probably correct in this treatment.

The genus should be readily distinguished by the combination char-
acter of a very short first abdominal segment with the postpetiole as
long as the petiole, broad temples, rather stout and basally attenuate
antennae, peculiarly formed mandibles in which the teeth are separated
only by a deep groove on the outer surface (this may be due to the
wearing away of the teeth), exareolate propodeum, lack of foveae at
the base of torgite 2, broad stigma, sinuate apical abscissa of radius.
The lack of foveae at the base of tergite 2, if this implies the absence
of the umbo, is very unusual for the Ophion group.

The genotype (from Persia) is a rather small ophionine (12 mm.),
black with ferruginous markings and legs and flavous wings.

10. Genus ALOPHOPHION,« new genus
Plate 50, Figure 12

Remarkable in its entire lack of the occipital carina and differing
only by that character from all species of Ophion, this group of species
should perhaps be considered only a geographical subgenus of Ophion.
But the several species before me, all from southern .South America,
present such uniformity of structure as to form a compact group more
conveniently treated here as a genus.

•» After this pQpor wa." w>nl fo tho printer a sp<«imon Identinol by Dr. Towncs a.s Iltllirigiella timilit
Rz6pliKPti WM rpccivcd from him. This specimen Is crrtnlnly not referable to the same trlt>o a.s Ilellicigia
bat beloDR.'' to the f)phionlnl. In the foreRoInK key to genera it runs directly to Apalophion Shestakov, and,
despite the very evident tcoth on tho mandibles, I believe it belongs to that genus. At my request Dr.
Townes has compared It with the original description and reiMirts that It npireea very well, not only with tho
Ronerlc description but also with the description of the genotype, except In details of coloration, especially
the color of the wing.i, which in //. timilit are blackish and in A. mirta are described a-S flavous.

If we accept Dr. Townes's Identification of //. Mimillii as correct, the following should replace "0. Oonus
Apatophion Shestakov" end Its citation, and llilluittella Si/>pllKctl should replace Apatophion Shestakov
in the key to genera:

9. r.cnua nKI-LWIGIF;i.I,A S7/pIlgotl

ntlltrigiellii Pzftn Ioeti. Ornera Inscctortim. fa.sc. 34, p. 23, 10H.'>. |Ocnotyi>e. IltUwigitUa nigrlpennU Sf.fipll-

getl. ny desicnntlon of VIereck, IJ. R. Nat. Mus. null. 83, p. 07, I»M.|
Apotophion finit."rrAKOV, Konowia, vol B, p. 202, 192fi.— Meter, Konowia, vol. 10, p. 18, 19.17. [aonolype:
Apatophion mima .Sh&itakov.) Monoba.'lc.
• From a-X4*oi-wltbout a ridge, in refcrenoo to the lack of the occipital carina.

440 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

In addition to the lack of the occipital carina, all the species, of
which only the genotype is identified specifically, conform to the
following description:

Head (pi. 50, fig. 12): Broad behind, temples nearly reaching out-
side tangents of eyes; stemmaticum high and sharply defined by a
surromiding groove; malar space short; mandible rather long and little
narrower at apex than at base; clypeus broadly truncate.

Thorax: Notaulices rather deep and foveolate anteriorly; pre-
pectoral carina complete; scutellum not at all margined; propodeum
long, in profile with a gradual, weakly convex slope. Wings with a
hairless area below base of stigma; discocubitus broken but with at
most a veiy short ramelius, basal portion nearly parallel to basal
vein; second discoidal cell rather long and narrow; postnervulus and
nervellus broken distinctly above middle.

Abdomen: rather slender.

Genotype. — Ophion chilensis (Spinola),

11. Genus AUSTRALOPHION Morley

Plate 52, Figure 45

Australophion Morley, A revision of the Ichneumonidae based on the collection
in the British Museum (Natural History), pt. 1, pp. 4, 30, 1912. [Genotype:
Ophion peregrinus Smith.] Monobasic.

I have seen no specimen of the genotype, but there appears no
doubt that the species described below is properly referred to the
genus. Despite the remarkable inflation of the scutellum, subalar
tubercles, and metapleura, this species is hardly more than subgeneri-
cally distinct from Ophion. It agrees with all the key characters
leading to Ophion.

AUSTRALOPHION INFLATUS. new species

Male. — Length 17 mm., antenna 20 mm.

Head: Temples strongly convex but not reaching outside tangents
of eyes; occipital carina complete, joining hypostomal carina far above
ventral articulation of mandible; ocelli not touching eyes, stenmiaticum
only moderately high and defined by rather weak grooves; face very-
little broader than frons, clypeus broadly truncate; malar space
distinct but short; mandible considerably narrowing toward apex;
antennae slightly longer than body and very slender, all joints much
longer than thick.

Thorax (pi. 52, fig. 45): Polished, finely and sparsely punctate;
notaulices broad and shallow anteriorly; scutellum high, almost
hemispherical, fovea deep; frena of scutellum and postscutellum not
at all foveolate; prepectoral carina complete; subalar tubercle and
metapleuron very large and convex; propodeum rather short and

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 441

convex, its surfaco iincvon but not (lelinitcly rug:osc, its posterior
angles rather prominent with two carinae diveriring hackwaicl from
eacli. ^Vings with a snnill liairless area below base of stigma; apical
abscissa of ratlins curving slightly forward fiom base; discocubitus
sharply broken by a distinct ramelliis, basal abscissa strougly con-
vergent with basal vein; postnervulus broken distinctlj- above middle;
nervellus broken slightly IjcIow middle. Legs ver}'' slender; claws
(male) ver}' densely pectinate.

Abdomen: Slender.

Brown, with legs to apices of femora coiicolorous, tibiae and tarsi
stramineous, antennae black, and the following markings lemon
yellow: inner orbits, an elongate spot in posterior orbit, a spot on
lower anterior margin of pronotum, scutellum, postscutellum, tegulae,
subalar tul)ercle, a large spot in lower posterior angle of niesopleuron,
and the metapleuron; wings slightly infumate, venation dark brown,
paler basally.

Type locality. — Egmont, New Zealand.

Tj/pg— U.S.N. M. No. 57G03.

One male taken February 29, 1920, by R. J. Tillyard.

12. Genus OPHIOGASTRELLA Brucs

Plate 54, Figure 57; Plate 55, Figurk 75

Ophiogostrella Brues, Ann. Ent. Soc. Amer., vol. 5. p. 201, pi. 27, fig. 1, 1912.
[Genotype: Ophiogostrella maculilhorax Brues.] Monobasic and designated.

Through the kindness of C. T. Brues I have been able to study the
two paratypes of the genotype, and one of these is now deposited in
the United States National Museum.

The peculiarly formed apicnl joint of the front tarsus (pi. 54, fig. 57)
with its notched lower outer margin and the venation of (he forewing
(pi. 55, fig. 75), especially the long narrow stigma with radius near
base, the strongly curved apicnl abscissa of radius, and the very short
intercubitus distinguish tliis genus immediately. Brucs described the
vertex as "not margined behind," but I find that the occipital carina,
though not stronir, is complete. The second disroidal cell is short
and broad, with the lower posterior angle shari)ly acute, the ba.se very
short and the di.scocubitus very strongly curved, but witliout a ramel-
lus; the nervulus dislinftly antefurcal; the n<'rvellus broken at about
its lower third. The prep(!ctoral carina is obsolete above and weak
elsewhere. The speculum is indistinctly defined because of the very
weak groove and the fact that the entire Tn(>sopl<'uron is poli.^hed; the
scutellum is carinately margined to apex. Otherwise the genus agrees
with all the key characters leading to the Ophion group of genera.

The genotype is the smallest (7-8 mm. long) species of the Ophionini
known to mo.

442 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

13. POTOPHION,' new genus

Plate 50, Figure 26; Plate 56, Figure 91

Essentially Ophion in all the group characters, this genus is remark-
able for the anthophagous trophi (pi. 50, fig. 26) and exserted oviposi-
tor (pi. 56, fig. 91). Those two characters should sufiice to distin-
guish it. Some of the other characters mentioned in the following
description of the genotype, especially the form of the head, may also
be of generic significance.

Genotype. — Potophion caudatus, new species.

POTOPmON CAUDATUS. new species

Head (pi. 50, fig. 26) : About as long as broad; eyes long and narrow;
combined face and clypeus much longer than broad, slightly broader
than frons; temples sharply receding; occipital carina weak, especially
medially and at lower extremities; cheeks and malar space very nar-
row; ocelli large, but not touching eyes; mandible much broader at
base than at apex ; trophi, especially the labium, unusually long, lobes
of Hgula about three times as long as broad.

Thorax: Slender; notaulices weak; scutellum not at all margined;
prepectoral carina strong, complete, very strongly sinuate; postpecto-
ral carina lacking; propodeum long, only moderately convex, with
complete basal and medially interrupted apical carinae. Wings with
radius at basal third of stigma, its apical abscissa gently recurved;
a sharply defined hairless area below base of stigma; discocubitus
sharply broken by ^a long ramellus, its basal abscissa strongly con-
vergent with basal vein; lower apical angle of second discoidal cell
acute, postnervulus broken a little above middle ; veins of transverse
brace in about the ratio of 1:2:4, abscissula long, strongly curved at
base, gently sinuate beyond; nervellus broken a little below middle.

Abdomen (pi. 56, fig. 91): Strongly compressed beyond tergite 2;
postpetiole depressed, more than half as long as petiole; tergite 2 with
spiracles very sHghtly basad of middle, umbo distinct, epipleura
separated throughout; ovipositor recurved, sheath about as long as
first tergite.

Ferruginous, with head largely, thorax laterally, parapsidal stripes
and lateral margins of mesoscutum, postscutellum, and petiole yellow;
dorsal edge of abdomen somewhat brownish; sheath blackish; wings
hyaline, costa and stigma yellow, other veins brown.

Type locality. — Mount Omei, Szechwan, China, 11,000 feet.

Type.— v. S. N. M. No. 57604.

Two females, both captured by D. C. Graham, the holotype in
July 1936, and the paratype at 12,000 feet, 30 miles north of Tatseinlu,
Szechwan, on July 9, 1923.

' From roHis = drinking, in reference to the long trophL

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 443

14. Genus AGATHOPHIONA Westwood

Plate 50, Figures U, 28; Plate 56, Figuke 98

Agathophiona Westwood, Tijdschr. Ent., vol. 25, p. 20, pi. 4, figs. 5-13, 1882. —
Cameron, liiolotria Centrnli-Americana, Hynienoptcra, vol. 1, p. 297, i>\. 12,
fig. 11, ISSC— AsHMEAD, Proc. U. S. Nat. Mils., vol. 23, p. 87, 1900.—
Schmiedeknecht, Opuscula ichneumonologica, fasc. 18, p. 1419, 1908.
[Genotype: Agathophiona jidvicornis Westwood.] Monobasic.

An anomalous genus, in respect to both the mouth parts and the
abdominal structure, but exhibiting most of the characters of the
Ophion group.

Head (pi. 50, figs. 11, 28): Temples buccate and extending far
beyond outside tangents of eyes; occipital carina distinct but not
reaching hypostomal carina; eyes and ocelli small, eyes shallowly
emarginate; occllocular line and malar space distinct; clypeus deeply
separated, broad. Hat, broadly truncate at apex; labrum exposed;
antenna stout, tapering both basally and apically, much shorter than
bod}'; mandible only slightly narrower at apex than at base, some-
what twisted, teeth bluntly rounded; trophi elongate, especially the
ligula, which is prolonged into two slender processes each grooved on
the imier side and fitting together to form a slender tube extending,
when at rest, posteriorly to or beyond apex of propodeum, galea of
maxilla elongately oval and thin, both maxillary and labial palpi
ver}' slender.

Thorax: Short-ovate; pronotum with scrobes weakly defined, pro-
notal sinus broad, exposing spiracularsclerite; mesoscutum precipitous
anteriorly, notaulices deep but short; scutellum broad, precipitate
posteriorly, margined only basally; mesopleuron and sternum convex,
prepectoral carina complete, sj)eculum defined, fovea deep, post-
pectus not defined; mctapleuron evenly convex, lower marginal
carina not especially strong; propodeum very short, precipitate and
Bomcwliat concave behind, barely overla[)i)ing hind coxae, basal
carina absent, apical carina i)resent only laterally, basal constriction
indistinctly interrupted, very narrow doi-sally except medially where
it is in the form rjf a pit, frenum extremely short; spiracle large, elon-
gate, well behind bottom of constriction. Wings dniscly hair}' both
dorsally and ventrally and without fciicstru or hairless area in dis-
cocubital cell; stigma narrtjwly triiiiigular, radius at about basal
fourth; basal absci.ssa of radius not thickeneil, weakly curved basally,
apical absci.ssa nearly straight; veins of transvereo brace in about
the {jroportions of l:l.o:3; discocubitus strongly curved, usually
without trace of ramellus; second discoidal cell twt)-thirds as broad
apically as long posteriorly, posteiior angle acute, base broad, the
postnervulus being broken at about middle, nervulus distinctly post-
f ureal; abscissula strongly curved basally; nervellus broken at or

444 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

slightly above middle, reclivous, upper abscissa perpendicular. Legs
stout and rather short; hind coxa rather short-ovate, flattened dor-
sally; hind femur little longer than combined coxa and trochanter;
tibia somewhat expanded apically, inner calcariumx barely half as long
as basitarsus, nearly uniform in thickness and without a conspicuous
fringe of long hairs along inner margin ; basitarsus much shorter than
other joints combined, apical joint much longer than fourth, rather
slender and decurved, claws rather wealdy curved, coarsely pectinate
in female, closely so in male.

Abdomen (pi. 56, fig. 98) : Kather weakly compressed and in
profile slender; first tergite broad and stout with spiracles at about
apical two-fifths, sternite barely reaching a point ventrad of spiracles;
tergites 1-3 subequal in length; tergite 2 with umbo, gastrocoeli
near base, indistinct, spiracles about at middle; tergite 6 in female
deeply cleft medially; hypopygium in female extremely long, directed
downward (in museum specimens) and wrapped around the ovipositor
which is directed upward with its tip enclosed in the short sheath,
which does not extend above the apical tergite, ovipositor rather
long and stout with the deep round dorsal notch near the apex ; male
genitaha unusually long.

The genotype is the only known species. It is shining black with
pale reddish antennae and tarsi and deeply infumate wings. It is
known only from Mexico.

15. Genus EREMOTYLUS Foerster

Eremotylus Foerpter, Verh. naturh. Ver. preuss. Rheinlande, vol. 25, p. 150,
1868 (no species included). — Thomson, Opuscula entomologica, fasc. 12,
p. 1193, 1888. — Bratjns, Arch. Ver. Freunde Naturg. Mecklenburg, vol. 43,
p. 98 (1889), 1890. — Sz^pligeti, in Wytsman, Genera insectorum, fasc.
34, p. 35, 1905 (part). — Schmiedeknecht, Opuscula ichneumonologica,
fasc. 18, p. 1450, 1908; suppl. fasc. 24, p. 49, 1935. [Genotype: Anomalon
marginatus (Gravenhorst) Jurine. First species included.]

Camptoneura Kriechbaumer, Zeitschr. Hj'm. Dip., vol. 1, pp. 22, 23, 1901.
[Genotype; Anomalon margincdus (Gravenhorst) Jurine. By designation of
Viereck, U. S. Nat. Mus. Bull. 83, p. 27, 1914.] Isogenotypic with Eremotylus
(Foerster) Thomson.

I have seen no specim.en representing this genus. Certain North
American species have been referred to it, but according to the
descriptions by Thomson and Brauns and the additional characters
communicated by J. F. Perkins such reference is incorrect. They are
properly referable to Enicospilus despite their lack of alar scleromes.

In erecting Eremotylus it was Foerster's intention to include in it
those species of Ophion in the broad sense, in which the discocubitus
is unbroken and the discocubital cell lacks scleromes. In his un-
pubhshed manuscript he assigned to it Ophion bombycivorus Graven-
horst and Ophion undulatus Gravenhorst. Ophion marginatus he

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 445

li'ft in Ophion, but from descriptions of that spocies it is apparent
that it agrees with the original description of Eremohjlus and can
stand as the genotype.

Subsequent to its original description Lremotyhis has had three
somewhat ditferent interpretations. The first, and the one here
followed, was that of Thomson, with Ophion marginatus Gravenhorst
as type. Ashmead (lS9t)) gave it a second interpretation when he
placed in it his E. arctiae, an American species of Enicospilus without
scleromes; while Kriechbaumer referred to it Ophion undulatus
Gravenhorst, genotype of AUocamptiis Thomson and Oi/matoneura
Kriechbaumer on a later page of the same paper in which he had
erected Cymaioneura for that species. The last two interpretations
I consider synonymous with Enicospilus Stephens, Mr. Perkins
writes, "the only species I recognize as belonging to Eremotylus is
marginatus Grav.," and " Camptoneura curvinervis Kriechbaumer,
I consider, belongs to a distinct genus."

The following generic description of Eremotylus is compiled from
those by Thomson and Brauns, with several characters (indicated
by quotation marks) furnished i)y Mr. Perkins interpolated:

Head: Temples broad; occipital carina distinct; ocelli small, not
touching eyes; clypeus not separated, apex rounded; mandible stout,
"not conspicuously twisted," teeth ec[ual.

Thorax: "Pronotal sinus broad"; prepectoral carina above and
notaulices obsolete; postpectoral carina absent; "speculum sharply
defined l^clow, strongly transcostate"; mesosternum with a conspicuous
tubercle on each side at posterior third; scutellum acutely margined;
propodeum short, basal constriction divided (Perkms says, "sharply
excavated and undivided"), basal carina acutely elevated in middle,
apical carina interrupted. Wings with "stigma narrowly lanceolate";
radius basally thickened and abruptly curved, "apical abscissa
evenly curverl"; "fenestra absent"; discocubitus not convergent with
basal, ramellus absent; brachial cell not narrowed apically, upper
margin straight; nervulus slightly antefurcal; postnervuhis broken
far al>ovo middle; a})scisHula strongly curved basally; ncrvellus
broken below middle, nearly ])erpendicular.

Abdomen: Membrane of tcTgite 1 reaching spiracle of tergite 2;
tergite 2 with umbo distinct and "wit!i a deep elongate furrow on
each side, which is subparallel with the lateral margin and extends to
about one-half the distance from the base to the spiracle, the spiracle
at middle of segment."

Such of the key characters as are included in the above description
indicate that Eremotylus belongs in the Ophion group and probably
near Boethoneura, new genus.

446 PROCEEDINGS OF THE NATIONAL MUSEUM vol.86

16. SIMOPHION.s new genus

Plate 50, Figube 23; Plate 52, Figure 48; Plate 54, Figure 55; Plate 56,

Figure 92

This and the next five genera form a series separable from the rest
of the Ophion group of genera by the form and course of the basal
abscissa of radius, which is distinctly thickened at the extreme base
and rather abruptly bent or curved immediately beyond the thicken-
ing, whence it is straight to the interception of the intercubitus. From
all the other genera the present genus differs in the pecuhar form of
the clypeus, the obsolete or lacking prepectoral carina, and the weakly
emarginate eyes.

Head (pi. 50, fig. 23) : Thick, temples strongly convex and nearly
or quite reaching outside tangents of eyes, occiput and frons not or
very shallowly concave; occipital carina rather weak but complete;
frontal orbits shghtly tumid; eyes parallel, very weakly emarginate;
stemmaticum sharply defined, ocelh not nearly touching eyes; malar
space short; clypeus very short, broadly truncate, usually concavely
so, at apex, protruding in such manner as to form a distinct angle with
face; mandible long and narrow, lower margin bent at nearly a right
angle, teeth narrow, upper one the longer; palpi very slender; antenna
slender, filiform, much shorter than body.

Thorax: Pronotal sinus distinct but unusually shallow, spiracular
sclerite visible; mesoscutum very strongly convex and precipitous
anteriorly, notaulices obsolete or wanting; scutellum narrow, margined
at most basally, its frenum and that of postscutellum shallow; pre-
pectoral carina (pi. 52, fig. 48) very weak or absent ; postpectoral carina
absent; speculum defined; propodeum excarinate, basal constriction
shallow. Wings (pi. 54, fig. 55) with discal cihation short and sparse,
a hairless area below stigma; stigma nearly as in Enicospilus, radius
at about basal fourth; basal abscissa of radius thickened and curved
at extreme base, apical abscissa somewhat reciu-ved from base; basal
vein decurved at base; discocubitus with first recurrent portion
sinuous, cubitus portion rather strongly arched, the two portions
joining in a rather sharp curve or a slight angle, sometimes with a
trace of ramellus, first recurrent and basal vein nearly parallel; inter-
cubitus half or more as long as second abscissa of cubitus; nervulus
interstitial or postf ureal; postnervulus broken shortly above, nervellus
shortly below, middle; frenulum short, with 6-9 slender hooks spaced
rather far apart. Legs very slender; claws w^eakly curved, pecten
(9) with short, fine teeth.

Abdomen (pi. 56, fig. 92) : Slender, tergite 1 decurved, petiole terete,
postpetiole depressed; tergite 2 with distinct umbo, spiracles at or
shghtly beyond middle, epipleura completely separated.

> From o-tj«A»=snubnosed, in reference to the short, projecting clypeus"

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 447

Oenotype. — Simophion ejcarinatus , new species.

SIMOPHION EXCARINATl'S, new speciea

Female. — Head: Temples hardly reaching outside tangents of ej'es;
posterior tangent of eyes bisecting posterior ocelli; malar space less
than half as long as basal width of mandible; cl3'peus concavely trun-
cate, more than twice as broad as long; antennae extremely slender,
45-jointed, all flageUar joints longer than thick.

Thorax: Densely and finely pubescent and laterally minutely
punctate and mat, shining dorsally and ventrally; propodeum finely
nigulose posteriorly, shining basally. Legs extremely slender, hind
femur about 10 times as long as deep, of nearly- uniform depth through-
out; inner hind calcarium hardly one-fourth as long as basitarsus.

Abdomen: Pohshed, very minutely pubescent; tergites 2 and 3
subequal in length and nearly as long as 1, 4 and 5 subequal and fully
three-fourths as long as 3, 6 two-thirds as long as 5, 7 and 8 together
about equal to G.

Dark brown; orbits so broadly yellowish white as to make the head
appear largely of that color; mesoscutum somewhat darker than rest
of thorax, sometimes with prescutum paler or with faint traces of
parapsidal stripes; wings hyaline, stigma and veins dark brown,
metacarpus, a spot at apex of costa and radices pale, tegulae stram-
ineous.

Type locality. — Tempe, Ariz.

Type—U. S. X. M. No. 57G05.

Paratypes. — ^CaUfornia Academy of Sciences; collection of Henry K.
and Marjorie C. Townes.

Seven specimens, including the holotype, collected at light at
Tempe, Ariz., by E. V. Walter and M. Martinez in February and
March, 1023 to 192G, under Tempo No. 4916; 1 female, Gila Bend,
Ariz., March 1, 1930, C. D. Lebert; 2 females, Phoenix, Ariz., 1933,
R. H. Crandall; and 2 females, Needles, Calif., February-March
1922, J. A. Kusche.

In addition to the genotype, representatives of 4 other species arc
before me. All are from the arid regions of the southwestern United
States. All of the 40 specimens are females.

17. TROPHOPHION." new genus

Plate 50, Fiourks 14, 20, 27; 1'i.ati. .'').'■), FunuE 78; Plate 56, FinuuE 99

A remarkable genus differing in many respects from tho more
typical genera of the Ophion group, notably in the form of tho head,
the anthopiiagous mouth i)arts, the bnsally curved and thickened
radius jiiid tho very little compressed abdomen with the female

• From rpo^i)<-food. In referonoe to the lonx trophl.

448 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 96

hypopygium elongate and protruding. In the last-mentioned char-
acter and the anthophagous mouth parts it resembles Agathophiona,
but whereas in the latter genus it is the labium that is most highly
developed, in the present genus it is the maxillae and the prolongation
is much less. Trophophion is most closely related to Genophion Felt,
from wliich it can be at once distinguished by its long tliin head,
subclavate antennae, the form of the abdomen, the unusually large
and coarsely pectinate claws, and by many other characters mentioned
in the following description.

Head (pi. 50, figs. 14, 20, 27): Extremely thin and long, occiput
deeply concave; temples strongly convex; malar space very long;
occipital carina fading out below and not reaching hypostomal carina;
face and clypeus wealdy convex, clypeus broadly truncate; labrum
semicircularly exposed, maxillae and labium very long, especially the
galea, palpi unusually short; mandible long and narrow, scarcely nar-
rowed toward apex, strongly curved, only slightly twisted; eyes
narrow; ocelli very small, stemmaticum not elevated and not defined
by a groove; antenna much shorter than body, flagellum slender at
base, gradually thicker toward apex.

Thorax: Stout; pronotum long and flat medially, with no transverse
groove; pronotal sinus broad, exposing spiracular sclerite; mesoscutum
moderately convex, notaulices absent; scutellum evenly convex, mar-
gined only at base; speculum wealdy defined; prepectus narrow, carina
obsolete above, postpectus not defined, propodeum very short, apical
slope precipitous, not at all overlapping hind coxae, median and lateral
foveae of basal constriction deep and widely separated. Wings with
coarse veins and dense, short discal ciliation, a small hairless area
below base of stigma; stigma narrowly triangular, with radius at
basal third; basal abscissa of radius tliickened and curved at base,
apical abscissa straight; second discoidal cell short and broad, lower
apical angle acute, discocubitus strongly curved; frenulum with 6 or 7
hooks; abscissula rather wealdy curved at base; nerveUus broken
above middle, upper abscissa perpendicular to mediella. Legs very
stout and short; apical tarsal joints long, claws (pi. 55, fig. 78) long
and coarsely pectinate, with few teeth.

Abdomen (pi. 56, fig. 99): Wealdy compressed; tergite 1 decurved;
tergite 2 with umbo short and rather wealdy defined, spiracles at
middle; apical tergites in female deeply emarginate medially; hypo-
pygium in female very long and protrudmg, ovipositor when sheathed
pointing dorsocephalad.

Genotype.— Trophophion tenuiceps, new species.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 449

TKOPHOPHION TKNUICEPS, new species

Female. — Ltnif^th 12 mm., antonna 8 mm.

Head: Moro than twice as broad as thick; temple extendinj^ slightly
beyond outside tangent of eye ; diameter of ocellus shorter than ocelloc-
ular line and hardly half as long as postocellar line; eye more than
twice as long as broad, shallowly emarginate within; malar space
distinctl}' longer than basal width of mandible; galea three times as
long as broaii ; antenna 30-jointed, subapical joints of llagellum thicker
than long and nearly twice as thick as basal joint.

Thorax: Evenly and rather densely punctate laterally (including
speculum) and ventrally, more sparsely and finely so and more polished
dorsally; propodeum with basal carina rather distant from base, dis-
tinct on each side of middle but fading out medially and laterally,
posterior face precipitate from basal carina, surface before carina
polished and punctate, apical slope indistinctly transversely rugulose
with traces of apical carina laterally. Wings with basal abscissa
of radius half as long as apical abscissa; veins of transvei*se brace in
about the proportion of 2:5:6; second discoidal cell shorter than
brachial cell, second recurrent more than half as long as basal
abscissa of subdiscoidcus; basal vein and base of discocubitus sub-
parallel; ncrvulus interstitial and inclivous; postnervuhis broken
shghtly above middle; nervellus broken at about upper third. Legs
stout and short, hind femur barely reaching apex of tergite 2 and less
than four times as long as deep; apical tarsal joints slender, decurved,
each as long as the corresponding second joint, claw two-thirds as
long as apical joint, pecten with five long, strong teeth.

Abdomen: Polished and virtually impunctate.

Bright ferruginous; wings hyaline with a brownish longitudinal
stain along apical anterior margin of each and one along basal vein;
venation black, costa and base of stigma yellowish; tarsal claws and
ovipositor sheath black.

Male. — Essentially like the female, except that tergites are not
emarginate.

Type local ily. — Phoenix, Ariz,

Holotype and allotype. ~\]. S. N. M. No. 57(5013.

Paratyj)es. — California Academy of Sciences; Citrus Experiment
Station, Riverside, Calif.; collection of Henry K. and Marjorie C.
Townes.

Described from four females and two males, (he holotype and nude
and female paratypes collected at Phoenix, Ariz., April 12, \\y.i[),
by K. H. Crandall; the allotype and a female j)aratyi)e at Palm
Springs, Calif., March 22, 19 Hi, by C. L. Fox; and a female in Palm
Canyon, Calif.. March 25, IWA'A, by P. H. Timberlake,

723504 — 47 8

450 PROCEEDINGS OF THE NATIONAL MUSEUM vol, 96

18. CLISTORAPHA,"' new genus

Plate 50, Figure 18; Plate 62, Figure 42; Plate 54, Figure 54; Plate 56,

Figure 96

From all the other genera of the Ophion group this genus is immedi-
ately distinguishable b}^ its possession of a complete postpectoral carina.

Head (pi. 50, fig. 18): Thick; temples strongly convex; occiput
deeply concave, carina complete; eyes and ocelli large, malar space
and ocellocular line much reduced, eyes deeply emarginate and nearly
parallel within (rarely in species with very strongly buccate temples,
the eyes and ocelli are smaller, the malar space and ocellocular line
rather long, and the face is considerably broader than the frons);
clypeus short, broadly truncate, coarsely punctate; labrum conspicu-
ously protruding beyond clypeus; mandible not strongly twisted,
narrow and strongly curved, little narrower at apex than at base;
antemia filiform, not or barely as long as body.

Thorax: Pronotal sinus broad, spiracular sclerite exposed; meso-
scutum sloping anteriorly, notaulices complete; scutellum margined at
least basally; speculum distinct; prepectoral and postpectoral carinae
(pi. 52, fig. 42) complete; propodeum moderately convex in profile.
Wings (pi. 54, fig. 54) densely, finely pilose, with a hairless area below
stigma; stigma elongate triangular, with radius at about basal third;
basal abscissa of radius curved and thickened at base, apical abscissa
shghtly curved ; discocubitus strongly curved, without ramellus, basal
portion nearly parallel to basal vein; second discoidal cell much
narrowed at base, its lower apical angle acute; nervulus distinctly
antef ureal to shortly postfurcal; frenulum short, with only 5 or 6
hooks; abscissula curved, somewhat thickened basally; nervellus
weakly broken at or below middle, upper abscissa rechvous. Legs
short and rather stout, hind femur barely reaching beyond apex
of tergite 2 ; claws closely pectinate, not more closely so in male than
in female.

Abdomen (pi. 56, fig. 96): Strongly compressed, tergite 2 with
spiracles shortly beyond middle, umbo distinct, epipleura separated
throughout; male genitalia with paramere deep, its ventral margin
evenly curved to the subacute apex.

Genotype. — Ophion subfuliginosus Ashmead.

Several other species occur in the arid southwestern United States,
all undescribed.

" From »<Xti<rT6i=closed, and ^¥>^=suture. In reference to the closing of the mesosulcus by the postpecto-
ral carina.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 451

CUSTORAPHA SUBFUUGINOSA (ABluncad), new combination

Opf ion subfuUginosvs Asiimead, Proc. California Acad. Sci., ser. 2, vol. 4, p. 126,
1894.— HooKKR, Traii.s. Amer. Eiit. Soc, vol. 38, p. 29, 1912. [Lcctotype
(hereby i!t\sigiiatcd) : The female specimen from Kl Ta.stc, Baja California,
Mexico, in the collection of the California Academj- of Sciences.]

One of the El Taste cotypos of tliis species (a female) is in the
United States National Museum and the other two and the ElChincho
specimen are in the California Academy. Through the kindness of
the Academy and of E. Gorton Linsley I have had the opportunity
to examine the male and the female from El Taste.

In addition to the types I have seen the following specimens:
One female from near San Fernando, Ariz., August 28, 1925, R.
Budlong; one of each sex, St. Xavier Mission, Tucson, Ariz., July 29,
1924, E. P. Van Duzee; two females and three males, San Pedro
River, Fairbanks, Ariz., September 6, 1927, J. A. Kuschc; one female,
Washington Mountains, near Nogales, Ariz., September 7, 1927,
J, A. Kuschc; Baboquivari Mountains, Ariz., September 15, 1928,
O. C. Poling; three females, 30 miles east of Quijotoa, Pima County,
Ariz., August 28-29, 1927, Cornell University lot No. 542 sub. 33G;
Cobabi Mountains, 80 miles west-southwest of Tucson, Ariz., Sep-
tember 3, 1921, E. R. Tinkham.

In his second reference to tliis species (Proc. California Acad. Sci.,
ser. 2, vol. 5, p. 547, 1895) Ashmead recorded two specimens, a female
and a male, from San Jos6 del Cabo, Baja California. These speci-
mens, both females though one is labeled male by Ashmead, are in
the National Museum. They are misidentified and are not congeneric
with subfuliginosus, but belong to the genotype species of the new
genus Boeihoneura described below.

19. BOETHONELRA," new genus

Plate 50, Figdhe 17

Closely related to Cliatorapha, but lacks the postpcctoral carina.
Agrees otherwise with the above description of Clutorapha, although
the temples are narrower, tlie clypeus relatively longer, the stigma
broader, the discocubitus more sharplycuived in the middle and
frequently with a trace of the ramellus.

(knoiype .— Bodhoneura arida, new species.

Several other species, all undescribed, occur in the and southwestern
part of the United States.

■■ From fioTiOiu^TcintoToc, and ffypor-neno, in reference to tho tblclccncd base of radius.

725504—47-

452 PROCEEDINGS OF THE NATIONAL MUSEUM vol. oe

BOETHONEURA ARIDA, new species

Ophion subfuliginosus Ashmead, Proc. California Acad. Sci., ser. 2, vol. 5, p. 547
1895 (not Proc. California Acad. Sci., ser. 2, vol. 4, p. 126, 1894).^ Mis-
identification. No description.

For explanation of the above citation see Clistorapha subfuliginosus
(Ashmead) .

Female. — Length 13 mm.

Head (pi. 50, fig. 17): Occiput deeply concave; temple rather
strongly convex, but not nearly reaching outside tangent of eye;
eye nearly twice as long as broad, rather shallowly emarginate;
ocelli narrowly but distinctly separated from eyes; malar space
short but distinct; face and clypeus nearly flat, coarsely punctate,
the face more densely and finely so medially and polished and nearly
impunctate laterally, the head elsewhere impunctate and polished;
clypeus twice as broad as long, very broadly truncate at apex ; exposed
portion of labrum nearly as long as clypeus, heavily sclerotized,
punctate, with long rather dense hair; mandible more than twice
as long as broad basally, tapering to apex, slightly twisted, punctate;
antenna slightly shorter than body rt:40-jointed, slender, of nearly
uniform diameter except for a slight apical taper.

Thorax: Stout, shining, punctate, mesoscutum less distinctly so,
notaulices briefly impressed anteriorly; scutellum high, polished and
sparsely punctate, margined basally, its frenum more coarsely and
densely punctate; frenum of postscutellum striate; propodeum with
basal carina medially and the apical carina laterally distinct, the
latter sometimes complete, basal area polished and sparsely punctate,
the rest of the surface finely rugoso-punctate ; spiracles large, elongate
reniform. Wings rather densely ciliate with bare areas below base of
stigma and in base of radiellan cell; stigma narrowly triangular with
radius distinctly basad of middle; thickening of basal abscissa of
radius extending nearly to middle; apical abscissa slightly curved;
basal vein slightly curved for most of its length with a sharper reverse
curve above its junction with median vein; discocubitus bent just
basad of bulla, the basal portion a weakly sigmoid curve; veins of the
transverse brace about in the ratio 1:2:3; second recurrent about half
as long as basal abscissa of subdiscoideus, the angle between the two
veins slightly acute; nervulus interstitial to slightly antefurcal;
postnervulus broken at about upper third; radiella strongly curved
and somewhat thickened basally; nervellus broken at or slightly below
middle, its upper abscissa distinctly reclivous; frenulum composed of
5 or 6 closely spaced hooks. Legs rather short and stout, hind femur
barely reaching apex of tergite 2; claws rather coarsely pectinate.

Abdomen: Stout, hardly twice as long as head and thorax; first

GENERIC REVISION OF THE OPHIONINI CUSHMAN 453

segment slightly decurved, spiracle at about apical third; tergite 2 a
little more than three times as long as broad at base.

Ferruginous; vertex, frons, subalar tubercles and scutellum, and
sometimes the notaulices, 3'ello\vish; wings hyaline, radial cell in-
fumate basally and along anterior margin, stigma and costa ferru-
ginous, other veins blackish.

Type locality. — San Jos6 del Cabo, Baja California, Mexico.

IloUdype and poraiypes. — U. S. N. JM. No. 57G07.

Paratypes. — Califurnia Academy of Sciences; American Museum of
Natural History (New York); Cornell University.

Nine females, the holotype and two others from the tj^pe locality;
two from Phoenix, Ariz., April 19, 1933, R. II. Crandall; one from 14
miles east of Oracle, Ariz., July 27, 1924, J. O. Martin; Kits Peak,
Rincon, Baboquivari Mountains, Ariz., August 1, 1946, 4,050 ft.;
one from Congress, Ariz., May 6, 1902, Oslar; one from Yerington,
Lyon County. Nev., July 29, 1909. A headless male from Yerington,
Nev., in the collection of Cornell University probably belongs here,
but because of its poor condition it is not included in the typo series.

20. Genus GENOPHION Felt

Plate 50, Figure 24; Plate 55, Figure 79; Plate 56, Figure 95

Genophion Felt, New York State Mus. Bull. 76 (19th Rep. State Ent.), p. 123,
1904. [Genotype: (Genophion gilletli Fe\t) = Genophion costalis (Cresson).
By original designation.]

Hooker '* synonymized Genophion gilleffi Felt with Ophion costalis
Cresson, thereby inferentially synonymizing the genus with Ophion
although he did not list it in the synonymy of Ophion. In my opinion
the genus is distinct.

Head (pi. 50, fig. 24): Thick; temples strongly convex; occiput
concave, carina usually complete though rarely obsolete below; eyes
narrow, not bulging, shallowly, broadly emarginate; ocelli small to
rather large, but never nearly touching eyes; malar space distinct,
frequently very long; clyj)eus rather flat, large, broadly truncate,
labnim prominent; mandible narrow, evenly curved, not much
broader at base than at apex; antenna much shorter than body,
slightly thickened before apex.

Thorax: Stout; pronotum rather long medially, with transverse
groove shallow, pronotal sinus broad, exposing spiracular sclerite;
mososcutum strongly convex in profdc, notaulices obsolete; scutellum
large, subquadratc, strongly convex, usually margined basally; meso-
ple\iron with speculum weakly defined; prepeeloral carina strong

» Tmns. Amcr. Ent. Poc., vol. 38, p. 2fl, 1912.

454 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

though usually not quite reaching anterior margin; postpectoral
carina absent; propodeum strongly convex in profile, entirely without
carinae. Wings densely hairy with a hairless area below base of
stigma, stigma triangular, with radius at about basal third; basal
abscissa of radius thickened and curved basally, apical abscissa straight
or weakly curved; discocubitus strongly curved, without ramellus,
basaUy subparallel with basal vein; second discoidal cell narrow at
base, lower apical angle acute; nervulus postfurcal; abscissula curved
basally; nervellus broken far above middle, upper abscissa rechvous;
frenulum very short, with only 5 or 6 hooks. Legs rather short and
stout, hind femur reaching httle beyond apex of tergite 2; claws (pi.
55, fig. 79) closely pectinate, no more closely so in male than in female.

Abdomen (pi. 56, fig. 97) : Very strongly compressed, its upper
margin serrate in profile due to the heavily sclerotized and acute
apices of the compressed tergites; segments 1 and 2 stout; tergite 2
with spiracles at about middle, umbo very distinct, epipleura com-
pletely separated; male genitaha with paramere slender, its ventral
margin straight basally then bent upward and rising obhquely to the
acute apex.

This is another genus of the arid southwestern part of the United
States, being represented there by five or six species, of which only
the genotype is described.

GENOPHION COSTAUS (Cresson). new combination

Ophion costale Cresson, Proc. Acad. Nat. Sci. Philadelphia, 1878, p. 366. —

Hooker, Trans. Amer. Ent. See, vol. 38, p. 26, 1912.
Ophion costalis Cresson, Dalla Torre, Catalogus hymenopterorum, vol. 3, pt.

1, p. 189, 1901. — MoRLEY, A revision of the Ichneumonidae based on the

collection in the British Museum (Natural History), pt. 1, pp. 58, 60, 1912.
Genophion gilletti Felt, New York State Mus. Bull. 76 (19th Rep. State Ent.),

p. 123, 1904.

I have examined the types of both costalis and gilletti and have seen
three additional specimens, a male from Colorado (C. F. Baker), a
female from Boulder, Colo., June 7, 1922, about 6,650 feet (American
Museum of Natural History), and a male from Mount Diablo, Calif.,
April 20, 1935 (G. E. and R. M. Bohart).

21. CHILOPHION," new genus

Plate 50, Figure 19; Plate 54, Figure 59; Plate 55, Figure 82; Plate 56,

Figure 97

Most closely related to Genophion in the basally bent and curved
radius, in the strongly reclivous nervellus with the fracture far above
middle and the upper abscissa also rechvous and in the serrate abdo-
men; but differing in the form of the clypeus, which is thick, flat, and

13 From xe'Xoi=liPi in reference to the large clypeus.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 455

apically ftrcuato, the lone:, slender letrs and antennae and in some of
tlie other characters detaik'd in the followinij; description:

Head (pi. 50, fig. 19): Thick; temples strongly convex but not
especially broad; occiput concave, carina complete; eyes rather
narrow, not bulging, broadly emarguiate; ocelli rather large, but not
contiguous with eyes, stemmaticum prominent and sharply defined
by grooves; malar space distinct but not long; clypeus large, thick,
flat, apex broadly arcuate, labrum very narrowly exposed; mandible
rather long and narrow, its lower margin sharply curved in middle;
antenna slender, filiform.

Thorax: Rather slender, pronotal sinus broad, exposing spiracular
sclerite; mesoscutum moderately convex in profile, notaulices obso-
lete; scutellum subtriangular, strongly convex, not margined; specu-
lum defined; prepectoral carina distinct except at upper end and
rarely obsolescent tlu'oughout; postpectoral carina absent; propodeum
weakly convex in profile, entirely without carinae. Wings (pi. 54,
i\^. 59) rather sparsely hairy, a hairless area below base of stigma;
stigma elongate triangular, radius at or slightly basad of basal third;
basal abscissa of radius thickened and curved at base, apical abscissa
somewhat curved; basal vein rather strongly decurved at lower end;
nervulus postf ureal ; second discoidal cell rather long, its lower apical
angle acute, base not especially narrow; discocubitus straight basally,
curved apically, without ramellus, basal portion nearly parallel \vith
basal vein; frenulum very short, with G or 7 hooks; abscissula curved
basally; nervellus broken far above middle, upper abscissa reclivous.
Legs slender; claws (pi. 55, fig. 82) rather closely pectinate.

Abdomen (pi. 56, fig. 97): Slender basally, strongly compressed
beyond tergite 2, serrate in profile, the compressed tergites heavily
sclerotized along dorsal margins and acute at apices.

Genohjpe. — Ophion abnormis Felt.

Another genus of the arid southwestern part of the United States,
whence I hav(! seen several species, of which onl}' the genotype is
described.

CIlILOi'IlION AUNOKMIS (Felt), new combination

Ophion ahnormum Felt, New York State Miis. Bull. 7G (lOtli Ucp. State Ent,),

p. 121, pi. 2, fiR. f), 1904.
ErymolyluK felli Vierkck, Trans. Kan.sas Acad. Sci., vol. 19, p. 312, 190.'i.
Ophion ahnormin IIooKEU, Trans. Amer. Ent. Soc, vol. 38, p. 47, pi. 3, fig. \U,

1912, female, not male.

Hooker is correct in synonyniizing/f//i with nbnonnU. Except for
the abnormal discocubitus of the type of the latter, the two types are
virtually identical, and I have labeled a apecinuMi in the Nafional
Museum that I compared with both types as a homotype of both
species. The males that Hooker referred to abnormis and the typo of

456 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Ophion abnormis magniceps Hooker have nothing to do \vith the
present genus, but are Ophion.

In addition to the types I have examined the following specimens:
Six females, Denver, Colo., April 26; two females, Denver, Colo.,
June 10, 1901, Dyar and Crandell; one female, Anniston Camp,
Colo., June 1933, Cockerell; two females, Fort Collins, Colo., June
24, 1913; two females, Hamilton County, Kans., 3,350 feet, F. H.
Snow; one female, Stanton County, Kans., 3,000 feet, S. J. Hunter.

22. Genus STAUROPOCTONUS Brauns

Plate 50, Figure 15; Plate 54, Figure 53; Plate 55, Figure 83

Stauropodonus Brauns, Arch. Ver. Freunde Naturg. Mecklenburg, vol. 43, p. 93,
(1889) 1890.— Kriechbaumer, Zeitschr. Hym. Dip., vol. 1, p. 22, 1901. —

Sz^PLiGbTi, in Wytsman, Genera insectorum, fasc. 34, p. 35, 1905 (as synonym
of Eremotylus Foerster). — Schmiedeknecht, Opuscula ichneumonoiogica,
fasc. 18, p. 1448, 1908; suppl., fasc. 24, p. 47, 1935. [Genotype: Ophion
bombycivorus Gravenhorst.] Monobasic.

Stauropododonus [sic!] Brauns, Morley, A revision of the Ichneumonidae based
on the collection in the British Museum (Natural History), pt. 1, p. 16,
1912 (part). — Uchida, Journ. Fac. Agr. Hokkaido Imp. Univ., vol. 21, p.
213, 1928 (part).

Nipponophion Uchida, Journ. Fac. Agr. Hokkaido Imp. Univ., vol. 21, p. 201,
1928. [Genotype: Nipponophion variegatus Uchida. By original designa-
tion.] New synonymy.

This and the next genus form a distinct group characterized espe-
cially by the entire lack of the occipital carina; very smxail mandible
with the upper tooth shifted to the inner margin, so that when closed
the mandible appears edentate; a longitudinal groove dividing the
mesopleuron into nearly equal upper and lower parts; extremely
narrow, attenuate stigma with radius near base, basally thickened
and strongly curved radius; and the presence on the outer apical
margin of the trochanter in middle and hind legs of a sharp decurved
tooth. Of these, the peculiarly formed mandible and the trochan-
teral tooth occur only in these two genera and in no other genus do
the other characters listed occur in the same combination.

Neither Szephgeti nor Morley nor Uchida understood this genus,
for Sz^pligeti synonymized it with Eremotylus Foerster, Morley
synonymized with it Spilophion Cameron, and Uchida accepted
Morley's interpretation while redescribing it as Nipponophion.
Both Eremotylus and Spilophion have the occipital carina distinct,
whereas the lack of this carina is one of the most anomalous characters
of Stauropoctonus.

Head (pi. 50, fig. 15): Unusually small; temples somewhat convex
but narrow and sharply receding; occipital carina entirely absent,
postvertex descending perpendicularly from ocelli, which are very
large and contiguous with eyes; eyes large, bulging, deeply emarginate,

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 457

parallel below autcnmic; face strongl}'- convex, narrow, but broader
than frons, clypeus almost continuous with face, convex, broadly
rounJetl at apex, narrowly exposing; labruni; nuilar space extremely
short; mandible small, so strongly twisted that upper tooth is on
inner margin; antenna long, very slender, attenuate.

Thorax: Pronotal sinus moderately broad, exposing spiracular
sclcrite; mesoscutum moderately convex, notaulices obsolete; scutel-
lum margined only basally; mesopleuron longitudinally impressed
from fovea to top of prepcctus; postpectoral carina complete and
high; propodeum briefly prolonged over hind coxae, basal carina very
strong, apical carina not defined, basal constriction divided into
median and lateral foveae. Wings (pi. 54, fig. 53) with dense long hair,
a hairless area below base of stigma, but without fenestra; stigma very
narrow, merging imperceptibly with metacarpus, radius near base;
basal abscissa of radius conspicuously^ thickened and curved basally,
apical abscissa strongly curved; discocubitus concavely curved
basally, convexly curved apically, basal portion slightly convergent
with basal vein; lower outer angle of second discoidal cell right;
nervulus interstitial or postfurcal; postnervulus broken at or above
middle; abscissula slightly thickened and weakly curved at base;
nervellus rectangularly broken at about middle; legs extremely
slender; apical joint of trochanter in middle and hing legs (pi. 55, fig.
83) with an acute tooth on outer side at apex; claws closely pectinate.

Abdomen: Slender, strongly compressed; petiole convex dorsally;
tergite 2 broadly constricted basally, with a deep longitudinal groove
on each side, but with no umbo, spiracles at apical two-fifths, epiplcura
separated and sharply infolded.

1 have not seen the genotype, an<l the above descrij)tion is drawn
from a damaged specimen of Nipponophion variegatus Uchida and the
new species described below. J. F. Perkins says (in Hit.): '^S.
bombycivorus Gr. agrees entirely with the generic diagnoisis of the
Chinese species."

STAL'ROPOCTONUS VARIEGATUS (IVhlda), new conililnnlion

S'ipponophton vririrgalus Uchida, Joiirn. Fac. Agr. Hokkaido Imp. Univ., vol. 21,
p. 201, 1928.

STAUROPOCTONIIS CIIEZANUS, new apcclca

Distinct from all the other described species in its almost uniformly
ferruginous color.

Female. — Length 25 mm., antenmi incomplete.

Temples very narrow; cominned face and clypeus twice as long us
width of face; eye less than twice as long as broad; thorax sparsely and
minutely punctate, mesopleuron below longitudinal furrow and meta-
pleuron more coarsely and densely so, mesopleuron rugoso-puuctate

458 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 9a

before middle coxa; basal carina of propodeum only moderately high,
basal area sloping into basal constriction, not precipitate, apical slope
flat with rugae radiating from foramen, and outlined laterally by
high curved rugae, below which the sides are vertically rugose.

Ferruginous; head largely yellow, stemmaticum black; scape,
pedicel and anellus piceous above, fiagellum yellowish ferruginous
slightly darkened basally; mesoscu turn with three brownish longitudi-
nal fasciae, the median one darker and more distinctly defined;
tergite 3 dorsally brownish, laterally together with entire fourth
tergite yellowish; tergites beyond fourth dark ferruginous; wings
faintly infumate, hairless area hyaline, base of costa and stigma and
metacarpus reddish, venation elsewhere blackish; legs concolorous
with body.

Type locality. — Beh Luh Din, 30 miles north of Chengtu, Szechwan,
China.

Type.—V. S. N. M. No. 57608.

One specimen captured July 27, 1933, at 6,000 feet altitude by
D. C. Graham.

23. AULOPHION,!^ new genus
Plate 50, Figuee 13; Plate 52, Figure 49; Plate 54, Figure 58

This is the American analog of Siauropoctonus Brauns, with which
it shares all the most anomalous characters, but it is immediately
distinguishable by its entire lack of the postpectoral carina and differs
further as follows:

Temples (pi. 50, fig. 13) weakly convex and exceedingly narrow;
eyes strongly convergent below emargination ; clypeus more strongly
rounded at apex, not or barely exposing labrum; pronotal sinus very
broad; scutellum subquadrate, not margined basally; longitudinal
impression of mesopleuron (pi. 52, fig. 49) m the form of a narrow
groove; postpectoral carma entirely lacking; propodeum with basal
and apical carinae complete to entirely lacking; forewing (pi. 54,
fig. 58) with hairless area extending along base of radius, extreme
base of radial cell also hau-less; discocubitus sharply curved or sub-
angulate at about middle, straight basally and weakly curved apically,
basal portion strongly convergent with basal vein; lov/er outer angle
of second discoidal cell sharply acute; nervulus antefurcal to inter-
stitial; postnervulus broken far above, nervelius far below, middle;
petiole flattened dorsally; tergite 2 with lateral grooves broad and
shallow, epipleura not infolded, separated only basally and there
only by thinner texture. Agreeing otherwise with the foregomg
description of Siauropoctonus .

Genotype. — Aulophion bicarinatus, new species.

" From auXAi= groove, referring to the groove on the mesopleuron.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 459

ALLOPHION BICAUINATUS. new iipccics

Female. — Loiifrth 20 mm., antenna 20 mm.

Head: Unsculpturccl e.xcept for very fine, sparse punctures on face;
face at level of clypeal foveae distinctly narrower than frons; antenna
as lonf^ as body, with 60 joints, all flajrellar joints much lonp:er than
thick.

Thorax: Pronotum slightly tumid along dorsal and anterior lateral
margins; mesosrutum minutely punctate; scutellum polished, its
frenum and that of postscutellum lijiely punctate, not at all foveolate;
mesopleuron polished above longitudinal groove, very minutely
shagreened, finely punctate and mat below, as are also the meso-
sternum and metapleuron; propodoum with short, dense, erect pile,
both basal and apical carinae complete and a smgle median carina
from basal carina to apex, transverse carinae arcuate, higher laterally,
median fovea of basal constriction longitudinally rugose, basal area
minutely sculptured, surface behind carinae finely rugulose. Wings
with basal abscissa of radius, measured across the curve, less than a
third as long as apical abscissa: intcrcubitus slightly longer than
second abscissa of cubitus; second discoidal cell nearly two-thirds as
broad apically as long on subdiscoideus, postnervidus broken at
upper third; intercubitella more than half as long as abscissula.
Legs very long and slender, hind tibia very nearly reaching apex of
abdomen.

Ferruginous; head (discolored but probably yellow), pronotum,
pleura, anterolateral margins of mesoscutum, and front coxae yellow;
antennae and abdomen from near base of tergite 3 infuscate; wings
hyaline, venation brown.

Type locality. — Costa Kica.

Type.— v. S. N. M. No. 57009.

One specimen received from the Paris Museum and captured in
1884 by de Lafon. It lacks most of the right antenna, parts of the
left front and middle legs, and the apical three joints of both hind
tarsi.

AULOPHION EXCARINATUS, new 8pecl«s

Differs from bicnnnntu.<i Cushman as follows:

Male. — Length 20 mm., antennae 22 mm.

Face at level of clypeal foveae hardly narrower than frons; antenna
slightly longer than body; pronotum not tumid along margins;
propodoum entirely without basal and median carinae and with oidy
faint lateral traces of apical carina, ininutcly punctate-rugulose;
basal ahsci.«sa of radius fnlly a third as long as apical abscissa; second
discoidal cell hardly half as broad apically as long on subdiscoideus;
intercubitella less than half as long as abscissula.

460 PROCEEDINGS OF THE NATIONAL MUSEUM vol.98

Entire head and thorax, except three broad ferruginous fasciae on
mesoscutum, yellow, as are also all coxae and trochanters; legs else-
where and abdomen stramineous, tergite 2 and dorsal edges of com-
pressed portion of abdomen ferruginous; antenna with scape, pedicel
and annellus piceous, flagellum ferruginous above, stramineous below;
stigma, costa, metacarpus and thickened base of radius ferruginous.

Type locality. — San Esteban, near Puerto Cabello, Venezuela.

Type. — Collection of Henry K. and Marjorie C. Townes.

One specimen captured by P. J. Anduze on December 20, 1939.

24. Genus BANCHOGASTRA Ashmead «

Plate 56, Figure 93

Banchogastra Ashmead, Proc. U. S. Nat. Mus., vol. 23, p. 87, 1900; Fauna Hawai-
iensis, vol. 1, pt. 3, p. 343, pi. 9, fig. 3, 1901. — Perkins, Fauna Hawaiiensis,
Suppl. 2, p. 680, 1910; Trans. Ent. Soc. London, p. 530, (1914) 1915. [Geno-
type: Banchogastra nigra Ashmead.] Monobasic.

Despite the stout form, especially of the abdomen, and the entire
lack of the fenestra, this anomalous Hawaiian genus is more closely
allied to Enicospilus than to Ophion in most of the important char-
acters distinguishing those two genera. It is perhaps significant that
neither Ophion nor any genus closely allied to it occurs in the endemic
fauna of the Hawaiian Islands, whereas Enicospilus is represented by
numero'13 species.

Head: Thick; temples strongly convex but considerably narrower
than eyes; occiput rather deeply concave, carina complete; eyes and
ocelli small, malar space and ocellocular line long; eyes broadly emargi-
nate; face and frons broad, about equal in breadth; stemmaticum
distinctly set off and somewhat elevated; clypeus narrow, apex
broadly rounded, labrum narrowly exposed; mandible much twisted,
abruptly narrowed from base; antenna slender, filiform.

Thorax: Short and stout; pronotal sinus narrow, spii-acular sclerite
concealed; mesoscutum in profile moderately convex, notaulices
obsolete; scutellum margined to top of the abrupt apical slope;
speculum not at all defined; prepectoral carina incomplete above;
postpectoral carina complete; propodeum very short, precipitous
behind, not at all overlapping hind coxae, only basal carina present,
basal constriction not interrupted. Wings with dense setae and with-
out fenestra or hairless area; stigma narrowly triangular, radius at
about basal third ; basal abscissa of radius straight and slender, apical
abscissa curving strongly from base; second discoidal cell narrow,
nearly or quite pointed at base, lower apical angle acute, discocubitus

" In his "Catalogue and Reclassification of the Nearctic Ichneumonidae" (Mem. Amer. Ent. Soc, No.
11, p. 737, 1944-45) Townes has synonymized this genus with Enicospilus. I exclude from Enicospilus all
species lacking the fenestra. This and the unusually stout habitus are, I think, sufficient to justify the reten-
tion of Banchogastra,

GEXERIC REVISION OF THE OPHIONINI — CUSHMAN 461

gently ciirvetl. not broken, its basal portion parallel to basal vein;
ncrvulus antefurcal; frenulum short, with or 7 hooks; abscissula
straifiht; nervellus broken slif^htly below middle, upper abscissa
inclivous. Lep^s short and stout; claws closely i)ectinate.

Abdomen (pi. 56, fig. 93): Short, stout, weakly compressed; first
segment very broad, depressed, sternite not nearly reaching spiracles;
tergite 2 extremel}'^ short and broad, hardly as long as third, without
umbo, spiracles basad of middle, epipleura defined only basally.

The only two species known are from the Hawaiian Islands. They
are black insects having more the general aspect of an exceptionally
stout campoplegine than of an ophionine.

2.-). Genus PYCNOPHION Ashmead

Plate 56, Figure 90

Pycnophion Ashmead, Proc. U. S. Nat. Mus., vol. 23, p. 87, 1900; Fauna Hawai-
icnsis, vol. 1, pt. 3, p. 344, pi. 9, fig. 4, 1901. — Perkins, Fauna Hawaiicnsis,
Suppl. 2, p. 680, 1910; Trans. Ent. Sec. London, 1914, p. 530, 1915. [Geno-
type: Pycnophion molokaiensis Ashmead. 1 Monobasic.

This is finothor anomalous Hawaiian genus allied to Enicospilufi
but remarkably difFerent in general form, in its lack of the fenestra
and in the strongly exserted, recurved ovipositor and obsolete post-
pectoral carina. The wing venation is nearly the same as in Bancho-
gastra, but the form of the body is very different.

Head: Rather thick, temples narrow, weakly convex and very
sharply receding; occiput narrow and weakly concave, carina complete;
eyes large and bulging, deeply emarginate; vertex narrow; stemma-
ticum not elevated, ocelli small, not touching eyes; malar space dis-
tinct; mandible twisted, strongly narrowing from base to middle;
clypeus roujided at apex, in profile strongly convex apically, labrum
narrowly exposed; anteima very slender, filiform, not or barely as
long as body.

Thorax: Stout; pronotal sinus narrow, spiracular sclcritc concealed;
mesoscutum precipitous anteriorly, notaulices obsolete; scutellum
broad, mnrgincd nearly or quite to apex; prepectornl cnnna distinct,
not quite reaching anterior miirgin of mesopleuron; j)ostpectoral
carina obsoletely indicated only at sides; speculum not defined;
metapleuron very strongly tumid; propodeum short, prccijiitous
behind, with or without the basal carina, basal constriction not
interrupted. Wings sparsely hairy except apically, with a hairless
area below base of stigma, but without fenestra; stigma narrowly
triangular, radius at about basal third; basal abscissa of radius straight,
slender, apical abscissa strongly curved; second discoidal cell narrow,
pointed or very narrow at base, lower apical angle right or slightly
acute, disrorubitiis gently curved, basal portion parallel to basal vein;

462 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

nerviilus antefiircal; frenulum short, with 5 to 8 hooks; abscissula
straight; nervellus broken below middle, upper abscissa inclivous.
Legs with femora rather stout, tibiae and tarsi long and slender, claws
closely pectinate.

Abdomen (pi. 56, fig. 90): Stout, moderately compressed; first
segment with petiole nearly terete, postpetiole depressed; ovipositor
exserted at least the length of first segment, usually recurved, sheath
slender.

I have seen all three of the described Hawaiian species, the single
specimen of fumipennis Perkins unfortunately headless. This species
exliibits some gradation toward Enicospilus in its long, narrow second
tergite with the spiracles well beyond the middle and its straight ovi-
positor. In the other two species the second tergite is notably short
with the spiracles at about the middle, and the ovipositor is recurved.

26. Genus SPILOPHION Cameron

Plate 54, Figure 56; Plate 55, Figure 77

Spilophion Cameron, Spolia Zeylanica, vol. 3, pt. 10, p. 124, pi. B, fig. 13, 1905. —
MoRLEY, A revision of the Ichneumonidae based on the collection in the
British Museum (Natural History), pt. 1, p. 16, 1912 (as synonym of Stauro-
podoctonus [sic!] Brauns). [Genotype: Spilophion maculipennis Cameron.]
Monobasic.

Coiloneura Sz^pligeti, in Wytsman, Genera insectorum, fasc. 34, p. 35, 1905.
[Genotype: Coiloneura melanosiigma Sz^pligeti. By designation of Viereck,
U. S. Nat. Mus. Bull. 83, p. 35, 1914.]

Since Spilophion and Coiloneura were pubhshed in the same year
and apparently at almost the same time there is some question as to
which actually has precedence. The evidence that I have been able
to gather is as follows: Part 10 of volume 3 of "Spoha Zeylanica"
bears the date October 1905. Fascicle 34 of "Genera Insectorum"
is dated simply 1905, but at the bottom of the last page is printed the
notation "Budapest, 14. August, 1905," which presumably is the
date on which the manuscript was transmitted by Szepligeti. This
fascicle of the "Genera Insectorum" was received at the library of
the Smithsonian Institution on November 15, 1905, and the "SpoUa
Zeylanica" on November 23, 1905. If we assume that both were
mailed immediately on publication, a much longer time would have
been required for the "Spolia Zeylanica", published in Ceylon, to
reach its destination than for the "Genera Insectorum," published in
Brussels, and the fact that the former was received only 8 days after
the latter would seem to give it precedence in publication. Moreover,
among the recommendations under Article 28 of the International
Code of Zoological Nomenclature are two which would seem to give
preference to Spilophion over Coiloneura. One recommendation is

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 463

to give preference to that jreniis for which n genotype is specified.
No genotype was specified for oitlier, but Spilopftion is monobasic and
Coilojieura was based on two species. The second recommendation
as stated applies to species but perhaps may also be interpreted to
apply to genera. It is that a "specific name accompanied bj^ both
description and figure stands in preference to one accompanied only
by a diagnosis or only by a figure." Spilophioii maculipennis is both
described anil figured. Fur the above reasons I prefer to use Spilo-
phion as the name of this genus.

The presence in this genus of the occipital carina is sufficient to
prove Morloy's error in synonymiziug it with Stauropoctonus.

In a majority of the group characters Spilophion agrees with
Enicospilus, especially in the body characters, but differs in most of
the wing characters and in the anomalous hind claws and penultimate
hook of the frenulum.

Head: Rather thin with temples strongly receding; occipital
carina strong but not quite reaching hypostomal carina; eyes and
ocelli large, eyes deeply emarginate, slightly more widely separated
across clypeal fovcae than across frons; clypeus convex, broadly
truncate, wnth a narrow refle.xed margin, labrum exposed; mandible
broad, only slightly twisted and little narrower at apex than at base;
antenna longer than bodj^, very slender, filiform.

Thorax: Pronotal sinus very narrow, spiracular sclerite concealed;
notaulices absent; scutellum flat, strongly margined; speculum not
defined; prepectoral carina strong, postpectoral carina complete;
propodcum flattened, basal carina strong, basal constriction uninter-
rupted. Wings (pi. 54, fig, 56) densely hairy, with a large hairless
area below stigma and base of radius, but without a well-defined
fenestra and with no scleromes; stigma small, triangular, tapering
beyond radius, which is distant from base; basal abscissa of radius
thickened at base, slightly bent or curved immediately beyond thick-
ening, thence slender, apiral abscissa strongly curved; second discoidal
cell very narrow at base, its lower posterior angle slightly acute;
abscissula (pi. 54, fig. 5Gb) curved; frenulum (pi. 54, fig. 5Gc) with
5 or hooks, the penultinuite hook usually larger and of difTerent
form from others; nervcllus broken somewhat below middle at a
right angle. Legs extremoly slender, apex of tibia hardly reaching
to apex of abdomen; claw of hind tarsus (pi. 55, fig. 77) sharply bent so
that apical portion forms an acute angle with base, last tooth of pecten
beyond apex of claw.

Abdomen: Slender; postpetiolc little broiulor than petiole; tergite 2
without umbo, spiracles shortly basnd of apical third; epipleura
completely separated and infolded; ovipositor sheath not exsertcd.

464 PROCEEDINGS OF THE NATIONAL MUSEUM vol.88

SPILOPfflON MACULIPENNIS Cameron

Spilophion macuUpennis Cameron, Spolia Zeylanica, vol. 3, p. 124, pi. B, fig. 13,
1905.

Stauropodoctonus orientalis Mori.ey, A revision of the Ichneumonidae based on
the collection in the British Museum (Natural History), pt. 1, p. 18, 1912.
New name for Spilophion macufipennis Cameron, not {Enicospilus) Stauro-
podoctonus macuUpennis (Cameron), 1886.

Because of Morley's error in transferring both this species and
Enicospilus macuUpennis Cameron to Stauropodocioniis , thereby
making them secondary homonyms, it was necessary for him to
rename the later species, but since they are not congeneric and neither
is a Stauropodoctonus I here restore Cameron's name for the present
species.

S. macuUpennis is widely distributed in the Indo-Malayan Region.

27. Genus ABANCHOGASTRA Perkins »•

Plate 54, Figure 60

AlhTjreodon Ashmead, Fauna Hawaiiensis, vol. 1, pt. 3, p. 343, pi. 9, fig. 2, 1901
(not Ashmead, 1900). — Perkins, Fauna Hawaiiensis, vol. 2, pt. 6, p. 679,
1910. (Genotype: Aihyreodon hawaiiensis Ashmead.] Monobasic.

Abanchogastra Perkins, Trans. Ent. Soc. London, 1902, pt. 2, p. 141. — Schmiede-
KNECHT, Opuscula ichneumonologica, fasc. 18, p. 1421, 1908. — Cushman,
Proc. Hawaiian Ent. Soc, vol. 12, p. 53, 1944. [Genotype: {Abanchogastra
debilis Perkins) = ^. hawaiiensis (Ashmead).] Monobasic.

This genus presents a curious combination of the characters of
Ophion and Enicospilus, the venation largely that of Ophion and the
head and body characters those of Enicospilus. In the last reference
cited above I have discussed the synonymy of the genus and that
of the genotype.

Head: With narrow, strongly receding temples; occipital carina fine
but complete; eyes deeply emarginate; ocelli large, nearly (9) or quite
(cT) touching eyes; frons shghtly narrower than face; clypeus hardly
separated, apex broadly rounded, labrum rather broadly exposed;
mandible abruptly narrowed from near base, strongly twisted, upper
tooth longer than lower tooth; trophi normal; antennae very slender,
nearly or quite as long as body.

Thorax: Slender; pronotum without definite scrobes, pronotal sinus
narrow, spiracular sclerite concealed; notaulices not at all indicated;
scutellum narrow, margined to beyond middle; mesopleuron nearly
flat with fovea obsolete and without defined speculum; prepectus
not reaching anterior margin of mesopleuron but ending abruptly
just above level of lower angle of pronotum; sternaulices not at all
indicated; mesosternum evenly convex; postpectus weak or narrowly

1' Townes (Mem. Amer. Ent. Soc, No. 11, 1944-45) has synonymized this genus with Enicospilus, Be-
cause of the lack of the fenestra and the generally Ophion-like venation, I retain it as a distinct genus.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 465

interrupted medially; nietajjleuron wealdy convex, lower marginal
carina rather weak throughout; propodcum, in profile, gently convex,
overlajiprng about basal third of hind coxa, entirely without carinae,
basal constiictioii miintcrrui)tcd, frenum long, spiracles rather small
and close to bottom of constriction. Wings (pi. 54, fig. 60) densely
hairy both doi'sally and ventrally, witiiout fenestra or hairless area
in discocubital cell; stigma broad with radius shortly before middle;
radius with basal abscissa nearly straight, not thickened, apical
abscissa cur\-ing strongly forwai'd at base; veins of transverse brace
(in genot3"pe) in ratio of 1:3:3; second discoidal cell very narrow at
base, half as broad apically as long posteriorly, posterior angle acute,
discocubitus strongly curved; ncrvulus strongly antcfurcal; abscissula
rather strongly curved basally; nervellus broken at about middle,
reclivous, upper abscissa perpendicular. Legs very slender; hind
co.xa elongate; inner hind calcarium less than half as long as basitarsus,
tapering from base and with a conspicuous fringe of long hair along
inner margin; basitarsus nearly or quite as long as rest of joints
together, joints 4 and 5 subcqual and together hardly longer than 3;
claws small, veiy coarsely pectinate, the pecteu consisting of 4 or 5
long strong teeth.

Abdomen: Slender, strongly compressed aj)ically; tergite 1 gradually
broadening, postpctiolc hardly twice as broad as petiole, spiracles just
posterior to apical third; tergite 2 shghtl}'^ shorter than 1, with shallow
basal lateral grooves, but without umbo, gastrocoeli elongate, midway
between base and spiracles, which are at apical third; ovipositor sheath
hardly reaching apex.

28. Genus ENICOSPILUS Stephens

Plate 49, Fiocre 2; Plate 50, FiounES 9, 10, 16, 22; Plate 52, Figures 40, 44
Plate 53, Figtre 52; Plate 54, Figures 61, 62; Plate 55, Figures 63-70, 74,
80; Plate 56, Figures 100, 101

Enicospilus Stephens, Catalogue of British insects, p. 352, 1820 (without descrip-
tion or ."^pccics) ; Illustrations of British cntonioIoRy, vol. 7, p. 126, pi. 40,
fig. 4, 1835, p. 311, 1845.— CusiiMAN, Proc. Hawaiian Knt. Soc, vol. 12, p. 39,
1944, (Genotype: (Ophion merdarius Stephens, not Gravenhorst) = Op/iion
comhunluH Gravenhorst.] Monohasic.

HcniconpiluH Agassiz, Xomenciatoria zoolof^ici index univci-salis, p. 178, 1846.
Emendation of Enicospilus Steplieiis. [Autotype: Ophion undulatus Graven-
horst.)

Allocampius Foer.ster, Vcrh. naturh. V^er. jjreuss. Rheinlande, vol. 25, j). 150,
1868. — Thomson, Opuscula cntomologiea, fasc. 12, p. 1186, 1888. [Geno-
type: Ophion comhuHtua Gravenhorst.) Present inclusion and designation.

Allocampius Tiiomso.n, Opu.'<cula enlomologica, fasc. 12, p. 1189, 1888, not
Foerster. (Genotype; Ophion undulatus Gravenhorst.) Monobasic.

466 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Dispilus Kriechbaumer, Berlin. Ent. Zeitschr., vol. 39, p. 309, 1894; Zeitschr.

Hym. Dip., vol. 22, p. 154, 1901. [Genotype: Ophion {Dispilus) natalensis

Kriechbaumer.] Monobasic.
Eremotylus (Foerster) Ashmead, Trans. Amer. Ent. Soc, vol. 23, p. 192, 1896, not

(Foerster) Thomson. [Genotype: {Ere?7iotyhis arctiae Ashmead) =Enicospi-

lus glabratus (Say) .] New combination. Monobasic.
Pleuroneur ophion Ashmead, Proc. U. S. Nat. Mus., vol. 23, p. 86, 1900. —

CusHMAN, Proc. Hawaiian Ent. Soc, vol. 12, p. 43, 1944. [Genotype:

Pleuroneur ophion hawaiiensis Ashmead.] Monobasic.
Cymatoneura Kriechbaumer, Zeitschr. Hym. Dip., vol. 1, pp. 22, 74, 1901.

[Genotype: Ophion undulatus Gravenhorst. By designation of Viereck, Proc.

U. S. Nat. Mus., vol. 42, p. 635, 1912.]
Pterospilus Kriechbaumer, Zeitschr. Hym. Dip., vol. 1, p. 156, 1901. [Genotype:

Ophion (Enicospilus) dubius Tosquinet. By designation of Viereck, U. S.

Nat. Mus. Bull. 83, p. 126, 1914.]
Irispilus Kriechbaumer, Zeitschr. Hym. Dip., vol. 1, p. 156, 1901, [Genotype;

Ophion (Enicospilus) trimaculatus Tosquinet.] Monobasic.
Eremotylus (Foerster) Kriechbaumer, Zeitschr. Hym. Dip., vol. 1, p. 152, 1901,

not (Foerster) Thomson. [Genotype: Ophion undulatus Gravenhorst. By

present designation.]
Leptophion Cameron, Proc. Zool. Soc. London, 1901, p. 227. [Genotype: Lepto-

phion longiventris Cameron.] Monobasic.
Dicaynptus Szepligeti, in Wytsman, Genera insectorum, fasc. 34, p. 28, 1905.

[Genotype: Dicamptus giganteus Szepligeti.] Monobasic.
Metophion Szepligeti, in Wytsman, Genera insectorum, fasc. 34, p. 28, 1905.

[Genotype: Metophion bicolor Sz6pligeti. By designation of Viereck, U, S.

Nat. Mus. Bull. 83, p. 94, 1914.]
Ceratospilus Szepligeti, in Wytsman, Genera insectorum, fasc. 34, p. 28, 1905.

[Genotype: Ceratospilus biroi Szepligeti.] Monobasic.
Atoponeura Szepligeti, in Wytsman, Genera insectorum, fasc. 34, p. 34, 1905.

[Genotype : (Atoponeura concolor Szepligeti, preoccupied in Enicospilus by E.

concolor [Cresson]) = Enicospilus atoponeurus, new name.] Monobasic.
Ophiomorpha Sz^ipligeti, in Wytsman, Genera insectorum, fasc. 34, p. 34, 1906.

[Genotype: (Ophion curvinervis Cameron, preoccupied by 0. curvinervis

Kriechbaumer) =0?) /won earner onii Dalla Torre. By designation of Hooker,

Trans. Amer. Ent. Soc, vol. 38, p. 134, 1912.]
Crypiocamptus BrEthes, Anal. Mus. Nac Buenos Aires, vol. 19, p. 230, 1909

(new name for Allocamptus Thomson, not Foerster). [Autotype: Ophion
undulatus Gravenhorst.]
Eremotyloides Perkins, Trans. Ent. Soc. London, 1914, pp. 530, 532. — Cushman,

Proc. Hawaiian Ent. Soc, vol. 12, p. 45, 1944. [Genotype: Eremotylus or-

bitalis Ashmead.] Monobasic.
Amesospilus Enderlein, in Michaelsen, Beitr. Kentn. Land- und Siisswasser-

fauna Deut. Sudw. Afiikas, vol. 1, p. 222, 1914-1916. [Genotype: Ophion

unicallosus Vollenhoven.] Monobasic and designated.
Schizospilus Seyrig, Mus. Nat. Hist. Nat., Mission Scientifique de I'Omo, vol.

3, Zool. fasc. 18, Hym. 2, Ichn., p. 79, 1935. [Genotj^pe: Schizospilus divisus

Seyrig. Original designation.]

As in the case of Ophion complete citations in the above synonymy,
especially under Enicospilus and Henicospilus, would require space
out of proportion to its value. The most comprehensive works are

GENEKIC REVISION OF THE OPHIONINI — CUSHMAN 467

tho same as those cited under Ophion, but as with that p'uus, the
major ])ortion of the Uterature is scattered through many periodicals
and the majority of the species are not to be found in pubhshed keys.

The name of this genus was originally spelled Enicospilu^, but
Agassiz (1S46) emended it to Henicospilus, presumably supposing
that Stephens had derived the fii*st part of the name from the Greek
word (i>ik6$. he being the ])roper transliteration of e. Several authors,
notably Dalla Torre, have adopted the emended spelling, widle othei^s,
notably Schniiedeknecht, have used the original spelling. To obtain
an unbiased opinion as to the proper form of the name I recently sub-
mitted the question to a committee on nomenclature set up in the
division of insect identification, U. S. Bureau of Entomology and Plant
Quarantine. The unanimous opinion of the committee was that, since
Stephens did not uidicate the derivation of the name, the original
spelling should be preserved. This decision was based on Opinion 34
of the International Commission on Zoological Nomenclature, which
rules that mdess e\'idence of the derivation of a name is clearly indi-
cated in the original pubUcation the original spelling should be pre-
served.

It will be noted that I have cited as the genotype of Enicos'pilus,
Ophion combvsiits Gravenhorst instead of Ophion merdarius Graven-
horst as given by Viereck, 1914. This change is based on Stephens's
identification of his own figure, originally identified as "Enicospilus
merdariv-'i" Asithout indication of the author of the species, as
comhufttus Gravenhorst; in other words, {Enicospilus merdarius
Stephens, 1835, not [Ophion] Enicospilus merdarius [Gravenhorst!) =
Enicospilus combustus (Gravenhorst). In this interpretation the com-
mittee mentioned abov(> also concurred unanimously.

Head (pi. 50, figs. 9, 22): Occipital carina present; temple usually
naiTOw and receding, rarely buccate and reaching outside tangent of
eye; eyes and ocelli large, eyes deeply emarginate; mandible usually
abruptly narrowed between base and middle and strongly twisted,
rarely gradually narrowed and only slightly twisted; ma.xillae and
labium of normal length.

Thorax: Tronotal sinus (pi. 55, fig. 74) narrow, spiracular sclerite
concealed; scutellum margined, usually to apex; speculum not defined;
prepectoral and pos([)<'cl,orHl carinae (pi. 52, fig. 40) strong, tlu^ latter
rarely interrupted medially; basal constriction of propodeum (pi. 52,
fig. 44) not divided. Wings (pi. 53, fig. 52; pi. 54, figs. 01, 62; pi. 55,
fig. 70) with stigma, narrow, cMiilling radius near base, usually sub-
parallel-sided distad of radius and rather abruptly tapering apically,
rarely very slender and merging imperceptibly with metacarpus; basal
abscissa of radius thickened and sinuate^ or undulant, very rarely
straight and unthickencd; apical abscissa strongly curved; fenestra

725504—47 5

468 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

present, with or without scleromes, rarely small and poorly defined;
abscissula straight; nervellus broken below (rarely at) middle, upper
abscissa usually inclivous. Legs with middle and hind trochanters not
toothed apically; pectination of claws normal (pi. 55, fig. 80).

Abdomen (pi. 56, figs. 100, 101): Tergite 2 without an umbo, its
spiracles usually at or near apical third, epipleura usually completely
separated and infolded, very rarely partially or entirely unseparated.

In some of the characters herein treated as making up a combination
that identifies an Enicospilus, especially in those of the wings, there
is considerable variation. Most of the synonyms listed above are
based on variations of wing characters.

The stigma, typically almost parallel-sided with a rather abrupt
apical taper, occasionally is like that of Thyreodon, extremely long
and slender and merging imperceptibly with the metacarpus. The
basal abscissa of the radius exhibits variation from marked curvature
and undulation even exceeding that of the genotype of Dicamptus to
the straightness and slenderness of that typical of Ophion. The
fenestra varies from a very small and poorly defined one, as in Ophio-
morpha concolor Szepligeti, to one occupying most of the apical part
of the discocubital cell and with all of the structures distinct; some
species lack scleromes entirely, others have from one to six. If only
one sclerome is present it may be either the proximal or the distal one,
or the quadra may be faintly sclerotized over most of its surface.
Other marked differences are exhibited in the course or form of the
discocubitus, the shape of the second discoidal cell, the proportional
lengths of the veins of the transverse brace, the strength of the post-
pectoral carina (rarely interrupted medially), and the form and
sculpture of the propodeum. In a very large majority of the species
the spiracles of tergite 2 are at or near the apical third, but in a few
unusually stout species they are much closer to, though still distinctly
beyond, the middle. Such species also tend to a lack of definite
separation between the second tergite and its epipleura, whereas
typically the epipleura are completely separated and inflexed; other
species exhibit partial separation, the basal half or more of the epi-
pleuron being separated and inflexed and the apical portion not
clearly separated and not mflexed. Typically the mandible is abruptly
narrowed before the middle and so twisted that the two teeth are in a
plane nearly vertical to the longitudinal body axis, but the variation
in its form extends all the way from nearly the condition found in
Stauropoctonus, with the torsion so great that the upper tooth appears
to be on the inner margin, to nearly that in Ophion, with less torsion
and gradual taper from base to apex as exhibited by Enicospilus
flavoplagiatus Cushman.

GENERIC REVISION OF THE OPHIONINI — CUSHMAN 4G9

Few species show marked departure from the normal in more than
one of the characters.

The use of the difTercnt number of scleromcs as a basis for the
segregation of genera or siii>gcnera, as in the case of Dispilw^ Kriech-
baumer, IVi^pilu.s Krieclibaumer, and Schizospilu^ Seyrig cannot be
defended, for it separates closely related species and groups together
unrelated species. The same is true of so-called genera based on the
j)osition of the scleromes, such as Amesospiluti P^nderlein, or on the
form of the basal abscissa of radius, such as Dicamptus Sz6pligeti, or
of the discocubitus, such as Aioponeura Sz^pligeti. For example,
from the figures alone in a paper by Seyrig (Mission Scientifique de
rOmo, vol. 3, fasc. 18, Ichneumonidae, pt. 2, 1935, Mus. Nat. Hist.
iSat.), in which he argues for the division of Enicospilus into three
genera entirely on the number and position of the alar scleromes, it
seems quite obvious that Enicospilus mediv-s Seyrig, E. mollis Seyrig,
Amesospilus Justus Seyrig, and A. rupeus Se3^rig are more closely
related to one another than any is to Amefiospllus fortis Seyrig or to
Enicospilus rubens Tosquinet; and the other characters mentioned in
the descriptions seem to bear this out.

To argue that a very laige genus is ipso fa-cto unwieldy and should
therefore be broken up into smaller genera by the most convenient
characters is unscientific. Many of the same characters can be used
in a more truthful manner within a genus, in association with other
characters, to segregate really related species. It is entirely possible
that by the use of combinations of characters the huge genus Enicos-
pilus can be divided into natural groups of somewhat less than generic
status, and it may be found possible to use some of the many names
already proposed in the subgeneric sense.

For additional examples of fenestra and wing venation in Enicos-
pilus beyond those in the present paper and in the paper by Seyrig
cited above, see the figures in the following publications: Cushman,
Arb. morph.-tax. Ent. Beiheftc aus Berlin-Dahlem, vol. 4, p. 296,
figs. 1-14, 1937; Cushman, Proc. Hawaiian Ent. Soc, vol. 12. p. 55,
1944; Cameron, Biologia Centrali-Amcricana, Ins., llym., vol. 1,
pi. 12, 188G; Brues, Bull. Mus. Conip. Zool., vol. 02, pi. 1, 1918.

In erecting his genera Allocamptus and Eremotylus, Foerster was
segregating frf)m the old genus Opiiion (hose Euroi)ean species in
whicii the discocubitus is not angularly i)rok('n, Allocamptus to include
those with scleromes in the discocubital cell and Eremvtylus those
without scleromes. In his unpublished manuscript he placed Opiiion
repeiitinus Holmgren, rarnululus (iravcnhorsl , combustus (Jravenhorst,
and merdarius Gravenhorst in Allocamptus, with tlie bust indicated as
his choice for genotype. To Erc.motylus he assigned 0. hombycivorus
Gravenhorst and undulatus Gravenhorst, without indicating the

470 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

genotype. To Eremotylus in the sense of Thomson have been referred
such American species as (Thyreodon) Enicospilus texanus (Ashmead)
(new combination) (pi. 55, fig. 64), {Eremotylus) Enicospilus rufoniger
(Hooker) (new combination) (pi. 55, fig. 66), and Enicospilus ameri-
canus (Christ) (pi. 55, fig. 69), the last under the name Eremotylus
macrurus (Linnaeus). Thomson's statement that Allocamptus
Foerster is coextensive with Enicospilus appears to have been the
first (imphed) reference of species to Allocamptus, and the name in
this restricted sense appears never to have been used in combination
with a specific name, most authors having followed Thomson's lead
in applying to it Ophion undulatus Gravenhorst. That Thomson
considered Foerster's generic names invahd because of lack of asso-
ciated species is shown by his use of the names Allocamptus and
Eremotylus. He applied Allocamptus to what Foerster thought of as
Eremotylus and Eremotylus to something different from either, that is,
Ophion marginatus (Gravenhorst) Jurine. 0. undulatus should not
be accepted as the genotype of Allocamptus Foerster, as was indicated
by Viereck (1914), since it obviously does not agree with the original
description, because of the lack of scleromes; and Allocamptus Thom-
son is a different concept, even though both are here considered
synonymous with Enicospilus. The case of Eremotylus is somewhat
different, for Ophion marginatus, the designated genotype, apparently
agrees with the original description, and that genus should be ac-
credited to Foerster, even though it appears that Thomson considered
that he was pubhshing it in a valid manner for the first time.

Dispilus Kriechbaumer first appeared as a subgeneric name under
Enicospilus without description other than that of its genotype. In
his second reference to the name Kriechbaumer proposed Pterospilus
as a new genus to include the subgenera Henicospilus (to be restricted
to species ^vith one alar sclerome), Dispilus (with two scleromes), and
Trispilus, a new subgenus (with three scleromes).

Both the angulate discocubitus and the small fenestra exhibited by
the genotype of Pleuroneurophion Ashmead (pi. 55, fig. 68) occur
elsewhere in Enicospilus, usually not in combination. The only
really anomalous feature of Pleuroneurophion is the exserted ovipositor
(pi. 56, fig. 100). None of the species referred to Pleuroneurophion by
Cameron, Szepligeti, and Uchida agrees in this character. In the
paper cited above I have already pointed out these facts and treated
Pleuroneurophion as a subgenus of Enicospilus.

The original inclusion of Ophion undulatus Gravenhorst in Cyma-
toneura and its subsequent designation as genotype by Viereck makes
that genus isogenotypic with Allocamptus Thomson. Those who
recognize this as a genus distinct from Enicospilus should use Cyma-
toneura as the generic name, since Allocamptus Foerster has priority

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 471

ovor Allocamptu.'^ Thomson. In cK-scribinc; Cymaioneura, Kricrh-
baunicr mentioned only two chnracters, the hick of scleronies in the
wintj and the course of the basal abscissa of the radius, "an oder
nahe der Basis verdickt, geschlanjrelt odor wellenforinig." As I
have indicated elsewhere, both of these structures are subject to great
variation within Enicospilus. The really most characteristic feature
of the genotype of Cymaiomura is the form of the head with the
temple strongly buccate behind the upper part of the eye and rapidly
diminishing in breadth and convexity below. This does not occur
in any of the other s])ecies referred to Cymatoneiira by Kriechbaumer
or in others that I know, though many have the temples strongly
convex but more uniformly so. Some of these, including the geno-
type of Evicospilvs, have distinct scleronies and the radius very
different. Others, witli the wing characters of Cymatoneura, have the
temples very narrow. Another somewhat peculiar feature of the
genotype is the slight concavity of the clvpeus, best seen in profile.
This is approached in such species as {Thyreodon) Enicospilus texanus
(Aslimead)," in which the clypeus is straight in profde with no inflec-
tion or reflexed margin apically. Thomson was quite wrong in
stating that undulatus lacks the postpectoral carina.

The synonymy of Leptophion Cameron is on the authority of J. F.
Perkins, who states that the genotype has alar scleromes and that the
claws are very sharply bent apically.

There is nothing in the description of Dicamptus Sz6pligeti to dis-
tinguish it from Enicospilus. Though I have been unable to find a
specimen of the genotype in the material available, I have identified
(Dicamptus) Enicospilus grammospilus (Enderlein) (new combination)
(pi. 55, fig. 70), which appears to bo properly referred to Dicamptus,
and on the strength of this synonymizc Dicamptus. Morley treats
Dicamptus as a subgenus of Allocamptus Thomson, thereliy imputing
ff> it "vertical" mandibles. Uchida, on the other hand, states that
the mandibles are "horizontal." He synonymizes "Allocamptus
Morley (not Thomson)" with Dicamptus, overlooking the fact that
Morley ineludcd in Allocamptus the genotype, undulatus
'^fravenhorst).

Most of the few characters given in the original description of
Metophion Sz6{)ligeti (the presence of two scleronies in the wing, th(»
strongly curved apical al^seissa of the radius, and the margined scu-
tolhmi) indicate affinity with hnir,ospilv^<i; in fact, the only discc^rdant
f'liararter is found in the expression "randmal nieht ausgebildet."
One would hardly say that the stigma is not developed in Enicospilus
or in any genus of Ophionini that I have seen. Before me arc speci-
mens of a specifically unidentified species of Enicospilus from the

" New combinatlun.

472 PROCEEDINGS OF THE NATIONAL MUSEUM vol.96

Oriental fauna that agree with the description of Metophion in all
but this character and in having the nervulus antefurcal rather than
postfurcal. They also agree with the description of the genotype
in the form of the alar scleromes, "einem mondformigen und einem
gewohnhchen Chitinfleck," that is, a somewhat triangular proximal
sclerome and a crescentic one on the anterodistal margin of the
quadra. Another character worthy of special mention in which the
specimens agree with the generic description is the unusually narrow,
sometimes alm^ost pointed, base of the second discoidal cell. On the
strength of these facts I synonymize Metophion with Enicospilus.

I have not seen the genotype of Ceratospilufi Sz6pligeti, but certain
Philippine specimens of Enicospilus agree in all but the postfurcal
nervulus, and this is too subject to variation for recognition as a generic
character.

Neither the medially thickened discocubitus nor the lack of scle-
romes is sufficient to segregate Atoponeura Sz^pligeti as a genus dis-
tinct from Enicospilus. An unidentified species of Enicospilus from
the Oriental Region exhibits these two characters (pi. 55, fig. 67).

It seems doubtful that Sz^pligeti knew the designated genotype of
Ophiomorpha, although he included it originally. In describing
Ophion curvinervis, Cameron stated, and his figure shows, that the
basal abscissa of the radius is distinctly curved, which would preclude
its tracing to Ophiomorpha in Sz6pligeti's key and its agreement with
the original description of the genus. I have examined two specimens
of Ophiomorpha concolor Szepligeti,^^ the most logical choice for
genotype, identified by Seyrig, and find that it has the basal abscissa
straight, only slightly thickened and very gradually tapering from
base to apex; also, the fenestra is obsolescent, being represented only
by a small hairless area in the normal position of the fenestra. Un-
fortunately, in describing curvinervis Cameron refers to the fenestra
only with the observation that the wings lack scleromes.

Cryptocamptus Brethes was proposed as a substitute for Allo-
camptus Thomson, not Foerster, its author not realizing that Cyma-
toneura was already available.

In describing Eremotyloides in a key to genera, Perkins did not
indicate any type except inferentially. On page 529 of the paper in
which the genus was described (p. 530) he stated, "I have not seen
the typical species of Eremotylus, Forst., and I think that Ashmead is
wrong in attributing the one variable Hawaiian species to it." On
page 532 in a discussion of the variation of Hawaiian Ophionini he
employs the combination "Eremotyloides orbitalis" without giving the
author of the species, but it seems entirely clear that Eremotylus

" {Ophiomorpha concolor Sz§pllgeti, preoccupied in Enicospilus by E. concolor Cresson) =EniscospUut
parvifenestratus, new name.

GENERIC REVISION OF THE OPIIIONINI — CUSHMAN 473

orbitalifi Ashmcad is intended as the genotype of Eremotyl aides, and
Viereck so interpreted it. In the reference cited above I have treated
Eremntyloides as a subgenus of Enicospilus showing that the fenestra
may be large or small and with or without sclcromes. A part of the
wing of EnkospHns (Eremotyloides) fullawayi Cushman is illustrated
herewith (pi. 55, fig. 65).

Amesospilns Enderlein is another segregate from Enicospilus based
on a wing character, the lack of central scleromcs, which it shares
with many otherwise very divei"se species.

Schisospilus Seyiig is a frankly artificial genus distinguished from
Enicospilus solely by the possession of two or more scleromes on the
quadra.

GENERA OF OPHIONINI NOT INCLUDED IN THIS REVISION
Genus BARYTATOCEPHALUS Schuiz

Banjcephalus Brauxs, Term^sz. Fiizet., vol. 18, p. 43, 1895. — Sz6pligeti, in
Wyt-sman, Genera inscctorum, fasc. 34, p. 24, 1905. — Schmiedeknecht,
Opiiscula ichncumoiiologica, fasc. 18, p. 1426, 1908; suppl., fasc. 24, p. 15,
1935. — Shestakov, Konowia, vol. 5, p. 257, 1926 (preoccupied by Bary-
cephalus Giinther, 1860). (Genotype: Darycephalns mocsaryi Brauns. By
designation of Viereck, U. S. Nat. Mus. Bull. 83, p. 19, 1914.]

Barytatocephalus Schulz, ZooI. Ann., vol. 4, p. 23, (1909) 1911 (substitute name
for Barycephalmt Brauns, preoccupied). [Genotype: Barytalocephalus moc-
saryi Brauns.] Autobasic.

The possession of the postpectoral carina, the weakly broken but
strongly rcclivous nervellus, and the large head are suggestive of
Clvitorapha Cushman, but the very small eyes and ocelli, the short
antennae, the unseparated clypeus, the lack of notaulices, the e.x-
carinate and polished propodeum, and the strongly contrasting color
pattern of Barytatoccphalus ai)pear to distinguish it.

The genus is still known only from the two original Hungarian
species. Brauns suggested that his two species might be the se.xes
of the same species, but Shestakov, in describing the male of sominiger
Brauns, showed them to be distinct.

Genus DICTYONOTUS Kriechbaumer

Dir.tyonotus KniEciiBAL