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HARVARD UNIVERSITY

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Museum of Comparative Zoology

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MAR - 5 1958

UNIVERSITY OF KANSAS

I

SCIENCE BULLETIN

>,'•

UNIVERSITY OF KANSAS PUBLICATIONS

University of Kansas Science Bulletin— Vol. XXXVIII— Part II

March 20, 1958
Lawrence, Kansas

ANNOUNCEMENT

The University of Kansas Science Bulletin (continuation of the
Kansas University Quarterly) is issued in parts at irregular inter-
vals. Each volume contains from 1000 to 1800 pages of reading
matter, with necessary illustrations. Exchanges with other institu-
tions and learned societies everywhere are solicited. All exchanges
should be addressed to

The University of Kansas Science Bulletin,
Library of the University of Kansas,

Lawrence, Kan.

PUBLICATION DATES

The actual date of publication (i. e., mailing date) of many of the
volumes of the University of Kansas Science Bulletin differs so
markedly from the dates bourne on the covers of the publication or
on the covers of the separata that it seems wise to offer a corrected
list showing the mailing date. The editor has been unable to verify
mailing dates earlier than 1932. Separata were issued at the same
time as the whole volume.

Vol.

XX— October 1, 1932.
XXI— November 27, 1934.
XXII— November 15, 1935.
XXIII— August 15, 1936.
XXIV— February 16, 1938.
XXV— July 10, 1939.
XXVI— November 27, 1940.
XXVII, Pt. I— Dec. 30, 1941.
I— May 15, 1942.
Nov. 12, 1942.
July 15, 1943.
Pt. II— Oct. 15, 1943.
XXX, Pt. I— June 12, 1944.
Pt. II— June 15, 1945.

XXVIII, Pt.

Pt. II
XXIX, Pt. I

Vol.

XXXI, Pt. I— May 1, 1946.

Pt. II— Nov. 1, 1947.
XXXII— Nov. 25, 1948.

XXXIII, Pt. I— April 20, 1949.
Pt. II— March 20, 1950.

XXXIV, Pt. I— Oct. 1, 1951.
Pt. II— Feb. 15, 1952.

XXXV, Pt. I— July 1, 1952.

Pt. II— Sept. 10, 1953.
Pt. Ill— Nov. 20, 1953.

XXXVI, Pt. I— June 1, 1954.

Pt. II— July 15, 1954.

XXXVII, Pt. I— October 15, 1955.

Pt. II— June 29, 1956.

XXXVIII, Pt. I— Jan. 6, 1957.

Pt. II— March 20, 1958.

Editor Edward H. Taylor

Editorial Board.

Edward H. Taylor, Chairman
Charles Michener
Paul Roofe
E. L. Treece

WOBTHIE H. HORR

Parke H. Woodard, Secretary

UNIVERSITY OF KANSAS

SCIENCE BULLETIN

DEVOTED TO

THE PUBLICATION OF THE RESULTS OF

RESEARCH BY MEMBERS OF THE

UNIVERSITY OF KANSAS

Volume XXXVIII, Part II

University of Kansas Publications

Lawrence, March 20, 1958

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA, KANSAS

1958

26-8050

L!ir«tiM

MA^ 5 1958

mm

Contents of Volume XXXVIII, Part II

Number page

13. Contribution to the Herpetology of Thailand.

Edward H. Taylor and Robert E. Elbel, 1033

14. Report on a Collection of Amphibians and Reptiles

from Harbel, Republic of Liberia.

Edward H. Taylor and Dora Weyer, 1191

15. The Tetranychidae of Mexico ( Acarina ) .

Robert E. Beer and D. S. Lang, 1231

16. A Revision of the Bees of the Subgenus Epinomia in the

New World ( Hymenoptera : Halictidae ) .

Earle A. Cross, 1261

17. The Status of the Mexican Lizards of the Genus

Mabuya Robert G. Webb, 1303

18. The Integumental Anatomy of the Monarch Butterfly

Danaus plexippus L. (Lepidoptera: Danaiidae).

Paul R. Ehrlich, 1315

19. A Revision of the Subgenus Melanonemotelus of Amer-

ica North of Mexico (Diptera: Stratiomyidae ) .

Wilford /. Hanson, 1351

20. Two New Fossil Plants from the Pliocene of North-

western Texas A. Byron Leonard, 1393

21. Anatomical Findings at Autopsy Immediately Follow-

ing Hanging Paid G. Roofe, Howard A. Matzke,

Irwin L. Baird and Edward C. Galbreath, 1405

22. A Statistical Method for Evaluating Systematic Rela-

tionships Robert R. Sokal and Charles D. Michener, 1409

23. The Role of Tissue Spaces and Nerve Fibers in the

Spread of Tetanus Toxin in the Rat.

Alexander A. Fedinec and Howard A. Matzke, 1439

24. The Family Succineidae (Gastropoda: Pulmonata) in

Kansas Charles D. Miles, 1499

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 13

Contribution to the Herpetology of Thailand

BY

Edward H. Taylor l and Robert E. Elbel 2

Abstract. A collection of Amphibia and Reptiles from Thailand is reviewed.
The following forms are regarded as new: Micrixalus magnapustulosus,

Mabuya macularia postnasalis, Mabuija macularia quadrifasciata, Mabuya mac-
ularia malcolmi, Leiolopisma siamensis, Sibynophis collaris triangularis and
Parahelicops boonsongi.

The known herpetological fauna of Thailand is listed as follows: Salaman-
ders 1, caecilians 2, frogs and toads 76, turtles 24, crocodiles 2, lizards 86, and
snakes 130, with a grand total of 321 species and subspecies.

CONTENTS

PAGE

Introduction 1034

Acknowledgments 1035

Methods 1035

Historical review 1036

List of Thai Provinces 1038

A list of the Amphibia known from Thailand 1039

A list of the Reptilia known from Thailand 1040

Taxonomic treatment 1047

Class Amphibia 1047

Order Gymnophiona 1047

Family Caeciliidae 1047

Order Salientia 1048

Family Ranidae 1048

Family Rhacophoridae 1068

Family Microhylidae 1070

Family Bufonidae 1075

Family Pelobatidae 1077

Class Reptilia 1079

Order Testudines 1079

Family Platysternidae 1079

Order Crocodylia 1081

Family Crocodylidae 1081

Order Squamata ' 1081

1. Department of Zoology, University of Kansas, Lawrence, Kansas.

2. Department of Zoology, University of Oklahoma, Norman, Oklahoma.

(1033)

1034 The University Science Bulletin

PAGE

Suborder Sauria 1081

Family Gekkonidae 1081

Family Agamidae 1083

Family Varanidae 1101

Family Scincidae 1101

Suborder Serpentes 1125

Family Typhlopidae 1 1 25

Family Dipsadidae 1126

Family Colubridae 1 130

Subfamily Sibynophinae 1 130

Subfamily Aeroehordinae 1 134

Subfamily Colubrinae 1134

Subfamily Boiginae 1142

Subfamily Natricinae 1149

Subfamily Homalopsinae 1159

Family Elapidae \16\

Family Hydrophiidae 1163

Family Crotalidae 1168

Literature Cited 1 174

Index " H80

INTRODUCTION

In the years 1953 and 1954 the senior author identified the species
in certain collections of reptiles and amphibia sent to him from
Thailand through the U. S. State Department. With the exception
of a few duplicate specimens the collections were returned to
Thailand, the specimens retained being placed in the Museum of
the University of Kansas. These collections, brought together
largely through the efforts of Dr. Boonsong Lekagul and the junior
author, will eventually become the basis of a national Thai collec-
tion. In 1955 the junior author, who had spent the preceding four
years in Thailand with the United States Operations Mission to
Thailand, returned to the United States with a number of specimens
which are now incorporated in the EHT-HMS Collection, property
of the senior author.

This latter collection also contains considerable studied material
from Thailand. One part of this was collected by Baron de
Schauensee and was reported by the senior author, in 1934.*
Another part was obtained by exchange with Dr. Malcolm Smith,
long resident of Thailand and physician to his Majesty the King of
Siam (now Thailand). Some of these are cotypes, paratypes or
topotypes, all identified by Dr. Smith. f These have been of especial
help in the study of our collections.

* Proceedings of the Academy of Natural Sciences, Philadelphia, vol. 86, 1934, pp.
281-310, pis. 17, text figs., 1-4.

1 In describing new forms Dr. Smith frequently designated a small series of numbers
as the "type series" although he may have had numerous other specimens before him.

Contribution to Herpetology of Thailand 1035

A third collection, from French Indo-China, obtained by ex-
change through the kindness of Dr. Rene Bourret of the Institut
Oceanographique de lindochine, has been indeed valuable for com-
parative study.

The country of Thailand is a strategic area in the study of the
south Asian faunae since its southern peninsula represents the high-
way along which emigrants from the continent reached the terri-
tories of the East Indies and the Philippines; and at the same time
species from the latter areas reached the peninsula and the main-
land. While it is not always certain in which direction the move-
ment took place, on the basis of the known distribution of a species,
it is possible to postulate its place of origin and later movement.
Thus it becomes evident that tracing the distribution of species
throughout Thailand can yield information of broad interest.
Moreover the mainland part of Thailand, especially the northern
part, is in a measure the pathway of exchange between Burma and
the Chinese — Indo-Chinese territories and thus the distributional
data lends much to our understanding of the faunae of these terri-
tories.

ACKNOWLEDGMENTS

We wish to express our thanks to Dr. Boonsong Lekagul of
Thailand who was instrumental in collecting or acquiring much of
the material from Thailand; to Mr. John Legler, curator in charge of
the herpetological collections of the University of Kansas, who has
been most helpful in making available materials and specimens in
his charge; to Mr. H. G. Deignan, U. S. National Museum for help
in clarifying certain locality designations. We especially acknowl-
edge use of the published works of Dr. Malcolm Smith treating of
the faunae of Thailand and Southeastern Asia.

METHODS

In this paper we are listing Thai species in the University collec-
tions (designated by the letters KUMNH) and certain ones in the
private collection of the senior author (designated EHT-HMS).
No attempt has been made to give all literature references and
synonyms, nor have we attempted to give all Thai localities in which
specimens have been taken. The names Siam and Thailand are
interchangeable. Siam appears in the older literature, Thailand in
the more recent. However, on postage stamps of the country one
or the other name appears at this time ( 1956 ) .

Concerning the field numbers, the early RE and Y numbers were
collected by Robert E. Elbel except for RE 1300 to RE 1556 and

1036 The University Science Bulletin

RE 2000 to RE 2500 which were collected by Robert E. Elbel and
H. G. Deignan; the numbers RE 3000 to RE 3975 were collected
by Robert E. Elbel and Dr. Boonsong Lekagul; the numbers above
RE 3975 were collected by Robert E. Elbel; all other field numbers
were collected by Dr. Boonsong Lekagul.

As for the spelling of Thai names, we are using the scheme of
transliteration from the Thai alphabet adopted by the Thai Gov-
ernment nearly two decades ago, and likewise used in the U. S.
Army Gazetteer of 1944. This is often at variance with spellings
encountered in European or older American maps, or in other re-
ports. We feel that an effort should be made to stabilize the system
of transliteration.

HISTORICAL REVIEW

A considerable amount of collecting has already been done in
Thailand aud numerous reports have appeared as evidenced by
the bibliography. No effort has been made to make the list com-
plete. Many of the older works, in which references to specimens
from Siam are made, are omitted.

Among the most important works on the herpetological fauna
of Thailand are those of Dr. Malcolm Smith, especially his volumes
on the Reptilia and Amphibia in "The Fauna of British India" and
the "Reptilia and Amphibia of the Malay peninsula from the Isthmus
of Kra to Singapore." Numerous shorter papers of his, dealing with
the Thai reptiles and amphibians, have appeared in various journals.

The snake, lizard and batrachian catalogues of the British Mu-
seum by George A. Boulenger are indispensable in a study such
as this.

Two papers by Major Stanley Flower are "Notes on a collection of
Reptiles and Batrachians made in the Malay Peninsula in 1895-96",
and "Notes on a second collection of reptiles made in the Malay
Peninsula and Siam." These two with a similar paper on the Ba-
trachians constitute a major contribution.

In the report at hand the locality records of many specimens
collected by the junior author are pin-pointed by recording the
name of the subvillage, village, district, and province, and where
a particular mountain is involved this is given with the approximate
elevation. Since most maps of Thailand fail to show the 71
provinces, a chart is presented showing the location of each. Often
a village, a district, and province may have the same name and in
older records it is not always possible to determine which is in-
tended.

Fig. 1. Map of Thailand. ( The numbers refer to names of provinces
and in themselves have no significance.)

1038

The University Science Bulletin

List of Thai Provinces

Corrected spellings as used by the U. S. Army Gazetteer, 1944.

1.

Chainat

37.

Lamphun

2.

Sing Buri

38.

Lampang

3.

Lop Buri

39.

Phrae

4.

Sara Buri

40.

Nan

5.

Ang Thong

41.

Uttaradit

6.

Ayutthaya

42.

Tak

7.

Nonthaburi

43.

Sukhothai

8.

Pathum Thani

44.

Phitsanulok

9.

Thon Buri

45.

Kamphaeng Phet

10.

Phra Nakhon (Bangkok)

46.

Phichit

11.

Nakhon Nayok

47.

Phetchabun

12.

Praehin Buri

48.

Nakhon Sawan

13.

Sarriut Prakan

49.

Uthai Thani

14.

Chachoengsao

50.

Kanchanaburi

15.

Chon Buri

51.

Suphan Buri

16.

Bayong

52.

Bat Buri

17.

Chanthaburi

53.

Nakhon Pathom

18.

Trat

54.

Samut Songkhram

19.

Chaiyaphum

55.

Samut Sakhon

20.

Nakhon Batchasima (Khorat)

56.

Phet Buri

21.

Buri ram

57.

Prachuap Khiri Khan

22.

Surin

58.

Chumphon

23.

Khu Khan (Sisaket)

59.

Banong

24.

Ubon

60.

Phangnga

25.

Nong Khai

61.

Surat Thani

26.

Loei

62.

Nakhon Si Thammarat

27.

Udon Thani

63.

Phuket

28.

Sakon Nakhon

64.

Krabi

29.

Nakhon Phanom

65.

Trang

30.

Khon Kaen

66.

Phatthalung

31.

Maha Sarakham

67.

Satun

32.

Kalasin

68.

Songkhla

33.

Boi Et

69.

Pattani

34.

Mae Hong Son

70.

Yala

35.

Chiang Mai

71.

Narathiwat

36.

Chiang Bai

A LIST OF THE AMPHIBIANS KNOWN FROM THAILAND *

Caudata
Salamandridae

Tylototriton verrucosus Anderson

Gymnophiona
Caeciliidae

*Ichthyophis glutinosus ( Linnaeus )
Ichthyophis monochrous ( Bleeker )

* The asterisk in front of a name indicates that the species is treated herein.

Contribution to Herpetology of Thailand 1039

Sahentia
Pelobatidae

Megophrys nasuta (Schlegel)
Megophrys longipcs (Boulenger)
*Megophrys aceras ( Boulenger )
Megophrys feae (Boulenger)
Megophrys monticola (Giinther)
Megophrys pelodytoides (Boulenger)
*Megophrys hasselti hasselti ( Tschudi )

BUFONIDAE

*Bufo asper Gravenhorst

Bufo borbonica ( Kuhl and van Hasselt)

Bufo macrotis Boulenger
*Bufo melanostictus Schneider

Bufo parvus Boulenger

Bufo penangensis Stoliczka

Nectophryne hosii Boulenger
Hylidae

Hyla anneetens (Jerdon)
Ranidae

*Ooeidozyga lima Kuhl and van Hasselt
*Phrynoglossus laevis martensi (Peters)

Elachyglossa gyldenstolpi Andersson

Rana cyanophlyetis Schneider
*Rana tigerina pantherina Fitzinger [ = Rana rugulosa Wiegmann]

Rana cancrivora cancrivora Gravenhorst

Rana cancrivora raja M. Smith
*Rana limnocharis limnocharis Boie in Wiegmann

Rana macrodon Kuhl

Rana kochangae M. Smith

Rana doriae Boulenger

Rana pileata Boulenger
*Rana macrognatha macrognatha Boulenger

Rana hascheana ( Stoliczka ) [ = Rana limborgi Sclater fide M.
Smith]

Rana aenea M. Smith
*Rana kuhlii Schlegel in Dumeril and Bibron

Rana laticeps Boulenger

Rana mortenseni Boulenger
*Rana cubitalis M. Smith
*Rana nigrovittata (Blyth)

Rana humeralis Boulenger

Rana miopus Boulenger

Rana macrodactyla (Giinther)

Rana nicobariensis ( Stoliczka )

Rana alticola Boulenger
*Rana erythraea (Schlegel)

Rana scutigera Andersson

Rana signata ( Giinther )

Rana tasanae M. Smith

1040 The University Science Bulletin

Rana glandulosa Boulenger

Rana luctuosa (Peters)

Rana haasei M. Smith
*Rana chalconota (Schlegel)
*Micrixalus magnapustulosus sp. nov.

Micrixalus tenasserimensis (Sclater)
Rhacophoridae

*Rhacophorus leucomystax leucomystax ( Gravenhorst )

Rhaeophorus nigropalmatus Boulenger

Rhacophorus prominanus M. Smith

Rhaeophorus colletti Boulenger

Rhaeophorus bimaculatus Boulenger

Philautus asper ( Boulenger )

Philautus horridus (Boulenger)

Philautus vittatus ( Boulenger )

Philautus petersi (Boulenger)

Philautus pictus (Boulenger)

Philautus bimaculatus (Peters)

Chirixalus doriae Boulenger

MlCHOHYLIDAE

Microhyla pulchra ( Hallowell )

Microhyla inornata Boulenger

Microhyla ornata ( Dumeril and Bibron )

Michohyla annamensis M. Smith

Microhyla butleri Boulenger

Microhyla heymonsi Vogt

Microhyla annectens Boulenger
*Microhyla berdmorei (Blyth)

Kalophrynus pleurostigma (Miiller)

Kaloula mediolineata M. Smith
*Kaloula pulchra pulchra Gray
*Glyphoglossus molossus Giinther

Calluella guttulata (Blyth)

A LIST OF THE REPTILIA KNOWN FROM THAILAND

Testudines
Dermochelidae

Dermochelys coriacea (Linnaeus)
Cheloniidae

Eretmoehelys imbricata (Linnaeus)
Chelonia mydas ( Linnaeus )
Lepidochelys olivacea (Eschscholtz)
Platysternidae

*Platysternon megacephalum Gray
Emydidae

PCyclemys mouhotii Gray
Cyclemys dentata (Gray)
Cuora amboinensis (Daudin)
?Geoemyda spengleri (Gmelin)
Geoemyda spinosa ( Bell in Gray )

Contribution to Herpetology of Thailand 1041

Geoemyda trijuga edeniana (Theobald)

Geoemyda grandis Gray

Damonia ** subtrijuga (Schlegel and S. Miiller)

Hieremys annandalii (Boulenger)

Notochelys platynota (Gray)

Siebenrockiella crassicollis (Gray)

Batagur baska ( Gray )
Testudinidae

Testudo elongata Blyth

Testudo emys Schlegel and S. Miiller

Testndo impressa (Giinther)
Trionychidae

Pelochelys bibroni (Owen)

Chitra indica (Gray)

Dogania subplana ( Geoffroy-St. Hilaire)

Amyda eartilaginea ( Boddaert )

Crocodylia
Crocodylidae

*Crocodylus siamensis Schneider
Crocodylus porosus Schneider

A LIST OF THE REPTILES KNOWN FROM THAILAND

Sauria
Gekkonidae

*Cyrtodactylus intermedins (Smith)

Cyrtodactylus consobrinoides (Annandale)

Cyrtodactylus oldhami (Theobald)

Cyrtodactylus peguensis (Boulenger)

Cyrtodactylus angularis (Smith)

Cyrtodactylus brevipalmatus (Smith)

Cnemaspis siamensis ( Smith )

Cnemaspis affinis ( Stoliczka )

Aleuroscalabotes felinus (Giinther)

Phyllodactylus siamensis Boulenger
*Hemidactylus frenatus Schlegel (in Dumeril and Bibron)

Hemidactylus garnoti Dumeril and Bibron
*Cosymbotus platyurus (Schneider)

Cosymbotus craspedotus (Mocquard)?
*Peropus mutilatus Wiegmann

Hemiphyllodactylus typus typus Bleeker
*Gekko gecko (Linnaeus)

Gekko smithii Gray

Ptychozoon lionatum Annandale

Ptychozoon kuhli Stejneger
Agamidae

* Draco whiteheadi Boulenger
*Draco maculatus (Gray)
*Draco haasei Boettger

** Mertens and Wermuth (Zool. Jahrb., Band 85, Heft 5, 1955) use the name Malay-
emys Lindholm, 1931.

1042 The University Science Bulletin

*Draco blanfordi Boulenger

Draco divergens Taylor

Draco volans Linnaeus

Draco fimbriatus Kuhl

Draco punctatus Boulenger

Draco cyanolaemus Boulenger

Draco taeniopterus Gunther

Draco fonnosus Boulenger
*Draco melanopogon Boulenger

Draco microlepis Boulenger

Draco quinquefasciatus Gray
*Goniocephalus armatus annatus ( Gray )
*Goniocephalus armatus crucigerus (Boulenger)

Goniocephalus lepidogaster (Cuvier)

Goniocephalus doriae Peters

Goniocephalus abbotti Cochran

Calotes cristatellus ( Kuhl )
*Calotes floweri Boulenger
*Calotes versicolor- ( Daudin )
*Calotes emma Gray

* Calotes mystaceus Dumeril and Bibron

* Liolepis belliana belliana (Gray)
*Physignathus cocincinus Cuvier

Varanidae

* Varanus bengalensis nebulosus (Gray)
Varanus rudicollis (Gray)

Varanus s. salvator (Laurenti)
Varanus d. dumerilii (S. Miiller in Schlegel)
Lacertidae

Takydromus sexlineatus Daudin
Scincidae

*Mabuya macularia postnasalis subsp. nov.
*Mabuya macularia quadrifasciata subsp. nov.

* Mabuya macularia malcolmi subsp. nov.
*Mabuya m. multifaseiata (Kuhl)
*Mabuya longicaudata (Hallowell)

Mabuya praesigne (Boulenger)

Mabuya rugifera (Stoliczka)
*Eumeces quadrilineatus (Blyth)
*Dasia olivacea Gray
*Sphenomorphus indicus indicus (Gray)

Sphenomorphus tersus (Smith)
*Sphenomorphus maculatus (Blyth)

Lygosoma quadrupes (Linnaeus)

Siaphos quadrivittatum (Peters)
*Leiolopisma rupicola (Smith)

* Leiolopisma eunice Cochran
Leiolopisma reevesii reevesii (Gray)
Leiolopisma reevesi melanostictum (Boulenger)
Leiolopisma tavesae Smith

Contribution to Herpetology of Thailand 1043

Leiolopisma kohtaoensis Cochran
PLeiolopisma doriae (Boulenger)

Leiolopisma vittigerum vittigernm (Boulenger)
*Leiolopisma siamensis sp. nov.

Riopa koratense ( Smith )

Riopa bowringii (Giinther)

Riopa isodactyla (Giinther)

Riopa herberti (Smith)

Tropidophorus berdmorei ( Blyth )

Tropidophorus robinsoni Smith

Tropidophorus thai Smith

Tropidophorus microlepis Giinther

Ophioscincus anguinoides (Boulenger)

Ophioscincus roulei (Angel)

Ophioscincus gyldenstolpei (Lonnberg)

DlBAMIDAE

Dibamus novae-guineae Dumeril and Bibron

Serpentes
Typhlopidae

Typhlops braminus (Daudin)
Typhlops floweri Boulenger
Typhlops lineatus Boie
Typhlops albiceps Boulenger
*Typhlops diardi muelleri Schlegel
Aniliidae

Cylindrophis rufus rufus (Laurenti)

XENOPELTIDAE

Xenopeltis unicolor Reinwardt in Boie
Boidae

Python reticulatus (Schneider)

Python molurus bivittatus Schlegel
Dipsadidae

Pareas laevis (Boie)

Pareas malaccanus (Peters)

Pareas margaritophorus (Jan)
*Pareas carinatus (Boie)

Pareas hamptoni (Boulenger)
*Pareas macularius Theobald

Haplopeltura boa (Boie)

COLUBRIDAE

Xenoderminae

Xenodermus javanicus Reinhardt
Sibynophiinae

Sibynophis geminatus (Boie)
*Sibynophis collaris triangularis subsp. nov.
Acrochordinae

Acrochordus javanicus Hornstedt
*Acrochordus granulatus (Schneider)

1044 The University Science Bulletin

Colubrinae

Gonyosoma oxycephalum (Boie)

Elaphe flavolineata (Schlegel)

Elaphe taeniura (Cope)
*Elaphe radiata (Schlegel)

Elaphe porphyracea porphyracea (Cantor)
*Ptyas mucosus (Linnaeus)
*Ptyas korros ( Schlegel )

Zaocys carinatus ( Giinther )
*Lycodon capucinus Boie

Lycodon laoensis Giinther

Lycodon fasciatus (Anderson)

Lycodon subcinctus Boie
*Ohgodon cyclurus (Cantor)

Oligodon purpurascens (Schlegel)

Oligodon taeniatus (Giinther)
*Oligodon cinereus (Giinther)
*OHgodon quadrilineatus (Jan)

Oligodon barroni ( Smith )

Oligodon joynsoni ( Smith )

Liopeltis scriptus (Theobald)

Dryocalamus davidsonii (Blandford)

Calamaria vermiformis Dumeril, Bibron, and Dumeril

Calamaria pavimentata Dumeril, Bibron, and Dumeril

Plagiopholis nuchalis (Boulenger)

Pseudorhabdion longiceps (Cantor)

Ahaetulla formosa Boie

Ahaetulla caudolineata Gray

Ahaetulla subocularis (Boulenger)

Ahaetulla cyanochloris (Wall)
*Ahaetulla ahaetulla ahaetulla ( Linnaeus )
Boiginae

*Boiga multimaculata (Boie)

*Boiga cyanea (Dumeril, Bibron, and Dumeril)

*Boiga cynodon (Boie)

Boiga dendrophila dendrophila (Boie)
*Dryophiops rubescens (Gray)
*Psammodynastes pulverulentus (Boie)

Psammophis condanarus indochinensis Smith

Chrysopelea paradisi Boie
*Chrysopelea ornata (Shaw)
*Dryophis prasinus Boie

Dryophis mycterizans ( Linnaeus )
*Dryophis nasutus (Lacepede)
Natricinae

*Pseudoxenodon macrops macrops ( Blyth )

Natrix piscator piscator (Schneider)
*Natrix piscator flavipunctata (Hallowell)

Natrix trianguligera (Boie)

Contribution to Herpetology of Thailand 1045

Natrix percarinata (Boulcnger)

Natrix inas ( Boulenger )

Natrix groundwater! Smith

Natrix desehauenseei Taylor

Natrix modesta (Giinther)

Rhabdophis s. subminiata (Schlegel)
*Rhabdophis subminiata helleri (Schmidt)
*Rhabdophis stolata (Linnaeus)

* Rhabdophis nigrocincta (Blyth)

* Rhabdophis chrysarga (Boie)
Macropisthodon rhodomelas (Boie)
Macropisthodon flaviceps ( Dumeril and Bibron )

*Parahelicops boonsongi sp. nov.

Opisthotropis spenceri Smith
Homalopsinae

*Enhydris plumbea ( Boie )

Enhydris jagorii (Peters)

Enhydris smithi (Boulenger)
*Enhydris enhydris (Schneider)

Enhydris bocourti (Jan)

[Enhydris chinensis (Gray)]
*Homalopsis buccata (Linnaeus)

Cerberus rhynchops (Schneider)

Bitia hydroides Gray

Herpeton tentaculatum Lacepede
Elapidae

Bungarus flaviceps Reinhardt

Bungarus fasciatus (Schneider)
*Bungarus candidus (Linnaeus)

Callophis maculiceps (Giinther)

Callophis gracilis hughi Cochran

Maticora bivirgata (Boie)

Maticora intestinalis (Laurenti)
*Naja naja kaouthia Lesson in Ferussac

Hamadryas hannah Cantor
Hydrophiidae

Laticauda colubrina (Schneider)

Laticauda laticaudata (Linnaeus)

Aepyurus eydouxi (Gray)

Kerilia jerdoni siamensis Smith
*Enhydrina schistosa (Daudin)
*Praescutata viperina ( Schmidt )

Thalassophis anomalus Schmidt

Kolpophis annandalei (Laidlaw)
*Lapemis hardwickii Gray

Microcephalophis g. gracilis (Shaw)

Pelamis platurus (Linnaeus)

Thalassophina viperina (Schmidt)
*Hydrophis mamillaris (Daudin)
*Hydrophis caerulescens (Shaw)

1046 The University Science Bulletin

Hydrophis klossi Boulenger

Hydrophis cyanocinctus Daudin

Hydrophis torquatus diadema Gunther

Hydrophis torquatus agaardi Smith

Hydrophis ornatus ornatus (Gray)

Hydrophis brookei Gunther

Hydrophis fasciatus atriceps Gunther
Viperidae

Vipera russellii siamensis Smith
Crotalidae

*Agkistrodon rhodostoma (Boie)

Trimeresurus monticola Gunther

Trimeresurus puniceus ( Boie )
*Trimeresurus albolabris Gray
*Trimeresurus popeorum Smith

Trimeresurus kanburiensis Smith

Trimeresurus purpureomaculatus andersoni Theobald

Trimeresurus wagleri wagleri (Boie)

Contribution to Herpetology of Thailand 1047

TAXONOMIC TREATMENT

CLASS AMPHIBIA

Family Caeciliidae

Ichthyophis glutinosus (Linnaeus)

Caecilia glutinosa Linnaeus, Systema Naturae, vol. 1, 1758, p. 393 (type locality
is here designated by us as Colombo, Ceylon. Linnaeus first described the
species in Mus. Adolph Frid., p. 19, tab. 4, fig. 1. The figure probably
represents the type).

Two adults, EHT-HMS No. 31729 (RE 4248), 31730 (RE 4263)
and seven young, 31731-31737 (RE 4291 young) from Ban Nong
Wai (subvillage), Na Phung (village) Dan Sai (district), Loei
(province) Thailand on Phu Nam Lang (mt.) elev. approx. 1780 m.,
Nov. 14-17, 1954, R. E. Elbel coll.

Published data on Ichthyophis glutinosus (as well as 7. mono-
chrus) show a very great range of variation in the number of pri-
mary and secondary grooves about the body. We very strongly
suspect that more than one form is involved in each species.
Boulenger states that the range in number of folds or grooves in
glutinosus is from 240 to 400, a variation of 160 folds, while in
monochrus the range is 235 to 360, a variation of 125.

The geographic range is large and an examination of series from
various localities will probably demonstrate that the variation men-
tioned above is geographical. A study of the dentition of individuals
of equal age will doubtless prove of much importance in delineating
other forms.

The type locality for 7. monochrus is Sinkawang, Western Borneo.
It was collected by Dr. Bleeker and later sent to the British Museum.
The type locality for I. glutinosus is uncertain. Presumably the
origin of the specimen described by Linnaeus is unknown.

One of the earliest descriptions of this species is that of Albertus
Seba, Thesaurus, vol. II, 1734, p. 26, pi. 25, fig. 2. The type locality
given is Ceylon. The description follows:

Num. 2. Serpens, Caecilia, Ceilonica. Squamis caret: at harum
loco tenuibus tegitur annulis, arete sibi mutuo conjunctis, spadiceo
colore tinclis. Alba quasi fascia per ventrem protenditur. Caput
peculiare nihil sibi vendicat, praeterquam quod late tumens, bi-
nisque amplis naribus, in antica parte, prope os, sit pervium.
Linguam in eo reperire non datur, at trachae tamen hiatum. Non

1048 The University Science Bulletin

admodum flexilibus est articulis; sed in se ipsam coacta riget. Vide
prolegomena.

Since it would appear that no conclusion as to the type locality
can now be drawn, we propose to designate Colombo, Ceylon as the
type locality. Thus, a study of comparative geographic variation
can be made.

The Siamese specimens have the folds numbering 431-438, the
primaries and secondaries being of nearly the same length.

The specimens, Nos. 31730 and 31729 yield the following data re-
spectively: Grooves, 431, 438; complete grooves, 64, 66; maxillary
teeth, 18-20, 19-?; vomerine teeth, 17-20, 15-?; outer mandibular
teeth, 19-20, 19-?; splenial teeth (inner mandibular), 14-14, 15-13;
premaxillaries, 5, 5.

It will be noted that the groove-counts of these specimens are
considerably higher than the "400" mentioned by Boulenger.

Order SALIENTIA
Family Ranidae

Ooeidozyga lima Kuhl and van Hasselt

Ooeidozyga lima Kuhl and van Hasselt, Isis, 1822, p. 475 (type locality, Java).
Oxyglossis * lima Smith, Journ. Nat. Hist. Soc. Siam, vol. 2, no. 2, Dec. 1916,

p. 164 (Klong Wang Hip, Nakon, Sritamarat ) ; ibid., pp. 172-175, pi.

(tadpole); Flower, Proc. Zool. Soc. London, Nov. 14, 1899, p. 886.
Ooeidozyga lima Smith, Proc. Zool. Soc. London, 1927, pp. 202-203; Bull.

Raffles Mus., no. 5, 1931, pp. 3-32.

This diminutive aquatic frog is rarely seen out of water. Its habit
of resting near the top of the water with only the eyes and nostrils
exposed, allows it to become aware of the approach of danger and
disappear below the surface with dispatch.

The nostrils are dorsal, nearer the eye than the tip of the snout
and surrounded by a slight ridge which develops a papilla on its
outer edge. The tympanum, relatively very large, is completely
covered by skin so that its outline is not visible without dissection.
The inner edge of mandible has a slight shelf which unites anteriorly
with one from the opposite side, forming a "second palate" anteriorly
which, seen directly from below, entirely or partially conceals the
choanae. No vomerine teeth are present. The tongue is pointed
posteriorly and free for much of its length.

Externally the body is covered with rough tubercles which on
the sides and ventral surface tend to form rows, some bearing a

* Long known under the genus Oxyglossus Tschudi (1838) (non Swainson); this was
replaced by Oxydozyga Tschudi (1838) by Stejneger, Proc. U. S. N. M., vol. 66, 1925,
p 33 Later, Malcolm Smith (1927) replaced the latter by the older Ooeidozyga Kuhl
and van Hasselt, 1822.

Contribution to Herpetology of Thailand 1049

series of pores perhaps derived from the neuromast system of the
tadpole.

The fingers are pointed, lacking all but a remnant of web, the
second longer than the first. The toes are almost completely
webbed, the tips of the digits slightly less pointed than the fingers.
A prominent tubercle that occurs below the tibiotarsal articulation,
that is itself covered with tiny tubercles, will identify the species.
The broad median dorsal cream stripe is usually present, as well as
the elongate brown stripes on the back of the thighs.

The younger specimens have the upper eyelids heavily tubercu-
late. The venter is slightly dark with longitudinal rows of silvery-
white spots marking the site of low tubercles. As they grow older
the dark pigment forms indefinite lines on the chin and on the
underside of the thighs, while the dorsum becomes variously
lineated. There are however usually two lateral stripes and a pair
of dorsal stripes. The latter may be broken leaving short lines and
spots. The tuberculation is intensified and practically no area lacks
the closely set spiny or rounded tubercles.

The largest specimen, 31720, measures 31 mm. snout to vent.
EHT-HMS Nos. 31720-23 (RE 3954) are from Ban Na Phua (sub-
village), Kan Luang (village), Na Kae (district), Nakhon Phanom
(province), Thailand, approx. 200 m. elev., July 28, 1954, Robert
E. Elbel and Dr. Boonsong Lekagul, colls. Three specimens, EHT-
HMS Nos. 31751-53, That Phanon (city), That Phanon (district)
Nakhon Phanom (province), Nov. 20, 1954, Robert E. Elbel, coll.

Phrynoglossus laevis martensii (Peters)

Phrynoglossus martensii Peters, Monatsb. Akad. Wiss. Berlin, 1867, p. 29 (type
locality, Bangkok, Siam).

Oxyglossus laevis martensi Smith, Jour. Nat. Hist. Soc. Siam, vol. 2, no. 2,
pp. 172-175, pi.; idem, vol. 2, no. 2, p. 227 (commonly and widely dis-
tributed); idem, ibid., pp. 885-916, pis. 59-60.

Phrynoglossus laevis martensi Smith, Bull. Raffles Museum, no. 5, 1931, pp. 3-32.

A very young example of P. I. martensii ( KUMNH No. 40195 ) is
from Pakchong (village), Sikiu (district), Khorat (province), Thai-
land, taken Aug. 23, 1952 and a second, older specimen (EHT-HMS
No. 31837) is from Udon Thani (city), Udon Thani (province),
Thailand, both specimens obtained by the junior author.

In No. 31837 the following characters may be discerned. The
snout is very flat, bluntly rounded when seen from above, without
trace of a canthus rostralis. The nostrils are widely separated, the
distance between them slightly greater than the interorbital distance.
The nostril is nearer the eye than the median point on the upper
lip. The tympanum is large, covered with skin but its outline can

1050 The University Science Bulletin

be seen, its diameter a little more than half of the length of the
eye. A fold of skin from the eye runs over and behind the tym-
panum and on to the side of the neck.

The tongue is elongate oval, rounded behind, free on the sides,
not or but slightly free behind. The vocal sac is present, the two
slits being at the level of the back of the tongue. The two palatal
shelves meet anteriorly forming a platform which does not conceal
the choanae when seen from below. Eustachian tube openings
larger than choanae. There are no vomerine teeth, but lying back
of choanae, and between their inner levels, there are two heavily
pigmented areas. The first finger is longer and larger than the
second, the inner digits with a lateral fringe or ridge extending to
the tips, no web or only a remnant of a web. The toes are three-
fourths to four-fifths webbed, the tips feebly dilated. An enlarged
compressed inner metatarsal tubercle about three-fifths as long as
the first toe. There is no tarsal fold or tubercle. The metatarsals
are united.

The skin on the head and dorsum is smooth but on the sides there
is a series of five or more pustules, some other indefinite granules,
and often small granules about the vent. The chin is smooth but
the venter is traversed by numerous fine well-marked wrinkles. The
thigh is smooth on the underside.

The color is brown lavender with a median cream stripe. Four
very indefinite darker stripes can be discerned when the specimen
is submerged in a clear liquid. The tibia and tarsus have one or
more indistinct darker bands. The chin and throat are brown,
heavily stippled with cream.

Measurements in mm.: Snout to vent, 21; width of head, 8; length
of head, 7.7; arm, 11.2; leg, 31.7.

We follow Malcolm Smith in regarding the genus Phrynoglossus
valid, and not a synonym of Ooeidozyga.

Rana tigerina* pantherina (Fitzinger)

Hydrostentor pantherinus Fitzinger, Ausb. osterr. Naturf., Sitz. kais. Akad. Band

42, p. 414 (type locality, Hong Kong).
Rana tigrina pantherina Steindachner, Reise der Osterreichischen Fregatte,

Novara, urn die Erde im 1857, 1858, 1859. Amphibien, Zool. Theil, Bd. 1,

1867, p. 17, pi. 1, figs. 14-17; Boulenger, Rec. Ind. Mus., vol. 20, June 1920,

p. 17.

Three specimens of Rana tigerina pantherina are in the collection.
These are: EHT-HMS Nos. 31820 (RE 3674), Phu Phan (mt.),

* Daudin spells the specific name of this species tigerina (Histoire naturelle des rain-
ettes des grenouilles des crapauds. An. XI (1803), p. 64, pi. 20). This can scarcely be
regarded as a typographical error since it is so spelled by Daudin in Histoire naturelles
generate et particuliere des reptiles, tome 8, An. XT (1803), p. 125-126.

Contribution to Herpetology of Thailand 1051

550 m., Sakon Nakhon (district), Sakon Nakhon (province), June
12, 1954, Dr. Boonsong Lekagul and Robert E. Elbel colls.; No.
31692 (RE 4018), Ban Na Muang (subvillage), Na Haeo (village),
Dan Sai (district), Loei (province), approx. 1780m. "same range
as Phu Nam Lang ( mt. ) but north." September 29, 1954, Robert
E. Elbel coll.; and No. 31821 (RE 4255), Ban Nong Wai (sub-
village), Na Phung (village), Dan Sai (district), Loei (province),
approx. 1780m. "range of Phu Nam Lang (mt.)", Nov. 15, 1954,
Robert E. Elbel coll.

The largest specimen, 31692, measures 104 mm. from snout to
vent. The femora have been broken by the collector so that accurate
measurements of the legs cannot be made. The toes are about four-
fifths webbed, the tips rounded into tiny discs. There is a free skin-
flap on the outer edge of the fifth toe, and a slight ridge or fringe on
the inner side of the first toe. The skin-fold above the tympanum
curves down closely behind the tympanum. The dorsal folds are
discontinuous on each side forming five to seven irregular lines of
short glandular ridges, which reach to the groin or onto the rump.
There is a pointed median denticulation on the lower jaw with two
shorter ones on each side.

On the body there are numerous scattered black or dark brown
spots, many associated with glandular ridges. The back part of the
thigh is dark brown reticulated with cream, as is the front side of
the femur, tibia and the lower part of the sides. The chin and
throat are cream with a series of more or less symmetrical lines and
spots. The upper lip bears cream spots surrounded by brown. The
top of the head is nearly uniform brown and the venter nearly uni-
foim cream.

The other two specimens are marked in much the same manner.
The ridges on No. 31820 are somewhat more elongate. The tibiotar-
sal joint reaches the nostril. In No. 31821 the joint reaches the eye.
In all, the first finger is longer than the second. Its tip may be some-
what better developed into a rounded disc than those on the other
fingers. The inner side of the second and third fingers have a dis-
tinct lateral fringe.

Rana limnocharis limnocharis Gravenhorst

(Fig. 2)

Fana limnocharis Gravenhorst, Deliciae Musei Zoologici Vratislaviensis con-
tinens Chelonias et Batrachia, Fasc. 1, Lipsiae, 1829, p. 42 (type locality,
Java ) .

Specimens of Rana limnocharis in the collection are from the fol-
lowing localities: EHT-HMS No. 31778 (RE 3674), Phu Phan

1052

The University Science Bulletin

(mt. ), 550m., Sakon Nakhon (district), Sakon Nakhon (province),
June 12, 1954, "large gravid female 59 mm. snout to vent"; Dr. B.
Lekagul and B. E. Elbel colls. No. 31750 (BE 3954) (lot of two,
one very young) from Ban Na Phua (subvillage), Kan Luang
(village), Na Kae (district), Nakhon Phanom (province), Thailand,

Fig. 2. Rana I limnocharis Gravenhorst. EHT-HMS No. 31778 9 Phu Phan
(mt. ), Sakon Nakhon (district and province), Thailand. Actual snout-vent
length, 41.

"on the edge of the range of Phu Phan (mt.), 200m. elevation";
Sept. 29, 1954. Nos. 31776-77 (BE 4018), from Ban Na Muang
(subvillage), Na Phung (village), Dan Sai (district), Loei (prov-
ince), Thailand; Sept. 29, 1954, B. E. Elbel coll. KUMNH Nos.
40186, 40188 (both BE 3259), Phu Kading (mt.), 1045 m., Si Than
(village), Wang Saphung (district), Loei (province), Thailand,

Contribution to Herpetology of Thailand 1053

Jan. 4, 1954; Dr. B. Lekagul and R. E. Elbel colls. Nos. 40008-
40009, Thailand, Feb. 1955 (no locality data); 40012, Phathalung,
Thailand, Feb. 1955; 40013, Mae Hong Son, Thailand, Feb. 1955,
Dr. Boonsong Lekagul coll.

One peculiar character * evident in these specimens is the pres-
ence of a ventral area outlined by a slight groove or fold that
crosses the breast, passes to a point on side behind the axilla, then
runs back to the median point between thighs. This line represents
the line of attachment between the skin and the muscles below.
In males at least the entire area is glandular and at certain times pro-
duces a covering of minute granulations most conspicuous on the
area following the breast. There is also a slight fold running from
the breast to the angle of the mouth. These are always evident but
are very conspicuous in the gravid female.

The male usually may be distinguished by its smaller size and
the deep black coloring of the vocal-sac area on the chin and throat.
A median cream line may be present or absent in either sex. This
species is common everywhere in Thailand except in high moun-
tains.

Rana macrodon Kuhl

Rana macrodon f Kuhl, in Tschudi, Classification der Batraehier, 1838, and in
Mem. Soc. Sci. Nat. Neuchatel, vol. 2, 1839, p. 80 (attributes the name to
Kuhl), (type locality, Java).

Rana macrodon var blythi * Boulenger, Rec. Ind. Mus., vol. 20, pp. 40-45,
part. Nakon Sritamarat (=Nakon Si Thammarat) type locality.

The following specimens are in the collection: KUMNH No.
40014 5 Nakhon Si Thammarat, "Thailand"; 40015 yg. "Thailand";
40016 "Thailand" collected by Dr. Boonsong Lekagul.

The first two specimens have been eviscerated. The third (No.
40016) is a male approximately 120 mm. snout- vent length; head
width, 46 mm.; length of head, 52 mm. In this specimen the diam-
eter of the tympanum (8 mm.) is less than the length of eye (11
mm.). The elongate toothlike projections from the anterior part
of the lower jaws reach an elevation of almost four millimeters.
The vomerine teeth are in two compressed elevated ridges, diag-
onally placed, reaching from a point a little distance in front of
choanal level to a point considerably behind posterior choanal level;
palatal glands open in an irregular curved series of thirteen pores
nearly midway between the anterior choanal level and the anterior

* This character is present in certain other species of Rana but is never conspicuous.

f Malcolm Smith ( 1930 ) regards blythi as a form not easily differentiated from macro-
don, and Boulenger later states that the name in "a racial sense cannot be used." The
association of Rana magna and Rana acanthi with macrodon as subspecies, as recently
proposed, we believe should be disregarded.

1054 The University Science Bulletin

part of palate; the jaw denticulations fit into deep depressions in
the upper jaw.

In No. 40014 5 the tympanum is a little less than 5 mm. in diameter
and its shape is oval rather than round. The snout-to- vent length
is 102 mm.

Rana macrognatha macro gnatha Boulenger

(Fig. 3)

Rana macrognatha Boulenger, Ann. Mag. Nat. Hist., ser. 8, vol. 20, p. 414;

Rec. Ind. Mus., vol. 20, p. 51 (type locality, Karin Hills, Burma).
Rana macrognatha macrognatha M. Smith, Jour. Nat. Hist., Soc. Siam, vol. IV,

no. 4, July 25, 1923, pp. 218-219, pi. 9, fig. 2 (adult male head; Nakon

Sritamarat Mts., Siam).

Two specimens, EHT-HMS Nos. 31724?, 31725 <? (Lot RE
3820), were taken at Phu Phan (int.), Sakon Nakhon (district),
Sakon Nakhon (province), Thailand, at an elevation of 550m.,
June 29, 1954, by Dr. Boonsong Lekagul and Robert E. Elbel.

Diagnosis: Head of male broad (25.5 mm.) equally as broad
as long, more than half the length of head and body; leg short,
the tibiotarsal articulation reaching to anterior part of eye; first
finger slightly longer than second; a free-edged flap or swelling
beginning between orbits extends some distance behind them; a
slight swelling on sides of occiput behind eyes; no dorsolateral
glandular fold. Diameter of tympanum equals length of eye in
male; no glands below tympanum or on humerus; skin covered with
minute granules intermixed with larger, often pearly-tipped pustules.

Description of No. 31725 J1 : A median pointed denticulation
on lower jaw flanked on each side by a broad-based, sharply pointed
spine, which fits into a slight depression of the upper jaw; choanae
partly concealed by shelf on maxillary when seen from below;
vomerine teeth (three to five in number) on two converging
diagonal ridges, largely behind posterior level of choanae, separated
widely from choanae and from each other; openings to vocal sac
are behind tongue, the openings small, puckered; posterior horns
of tongue prominent, the tongue narrowly free behind (in female
more than a third free). Palatal glands open through two canals
each with a small opening that is closer to level of choanae than
to front end of palate.

Distance between nostrils equal to their distance from eye and
from the median point on upper lip; canthus rostralis not evident;
width of an upper eyelid less than interorbital width; between
orbits a tonguelike raised area extending from front of eyes back
onto occiput, the posterior edge free, rounded; occipital area swollen

Fig. 3. Rana macrognatha macrognatha Boulenger. Upper figure
EHT-HMS No. 31724 9 . Actual snout-vent length 49 mm.; lower EHT-
HMS No. 31725 6 . Actual snout-vent length, 50 mm. Both from Pliu
Phan (mt. ), Sakon Nakhon (district and province), Thailand.

1056 The University Science Bulletin

on each side posterior to eye; tympanum distinct, its diameter equals
eye length, covered with closely adherent skin, the surface with
minute granules; no glands present at mouth angle or below tym-
panum and none on humerus; an inverted V-shaped fold back of
occiput; two more or less distinct rows of elongated pustules be-
hind tympanum extending to groin; dorsum and limbs with smaller
pustules, those on limbs usually arranged in indefinite longitudinal
rows, the pustules with pearly tips; chin with a few indistinct longi-
tudinal wrinkles; venter with very numerous, moderately distinct
transverse folds or wrinkles; a fold across breast reaches to near
mouth angle; median and posterior ventral part of thigh with large
granules. Arms short, the first finger slightly longer than the sec-
ond; a narrow fringe along part of inner side of the second and third
fingers; subarticular tubercles well developed; three palmar (meta-
carpal) tubercles, innermost elevated, outermost elongated.

Legs short, when folded at right angles the heels barely touch;
the tibiotarsal articulation reaches front edge of eye. An elongate
inner metatarsal tubercle in line with a distinct tarsal fold and a fold
or fringe that extends along outer edge of inner toe; no outer meta-
tarsal tubercle; a fringe along outer edge of fifth toe; outer metatar-
sals separated by a web; distinct discs on toes, with a groove on
outer edge; toes about three-fourths webbed.

Color: In preservative the frontal and interorbital areas light
brown; occiput darker brown, the color running forward on each
side onto eyelids; sides a little darker than the light brown back;
legs barred with dark brown; soles of feet brown. Venter and under-
side of limbs cream; brown spots on upper and lower jaws; tym-
panic area light tan.

Measurements in mm.: ( $ and § ): Snout to vent, 50, 49; head
length, 26, 21; axilla to groin, 19, 20; arm, 26, 27; leg, 67, 72; tibia,
24, 22; foot, 31.5, 31.

Remarks: The specimen from the same locality, identified as the
female of the species, lacks the specialization of the head of the
male. There is a strongly defined cream stripe on the head and body
from lip to near vent; the fold across the breast and shoulders is
more distinct. Toothlike projections on the jaws are only slightly
indicated; and the upper eyelid is as wide as the interorbital dis-
tance. The diameter of the tympanum is about three-fourths of the
length of the eye. Variation in the width of the head is indicated
in the measurements.

Contribution to Herpetology of Thailand

1057

Rana kuhli Schlegel

(Fig. 4)

Rana kuhli (Schlegel [Mus. Ludwig Batavia]) Dumeril and Bibron, Erpetologie
Generate. . . ., vol. 8, 1841, pp. 384-386 (type locality, Java); Smith,
Jour. Nat. Hist. Soc. Siam, vol. 2, May 1917, p. 227 (peninsular, western,
and northern Siam: common on many of the hills above 300 m. ); idem, vol.
2, no. 4, Dec. 1917, pp. 262-263, pi. fig. 1 (tadpole), (Doi Nga Chang,
March, 700-1000 m. elev.); Giinther, Beptiles of British India, 1864, pp. 404-
405, pi. XXVI, fig. A (but not fig. B) (part).

The following specimens are in the collection: KUMNH No.
40185 (RE 3156), Khao Sawan (mt), approx. 600m. elev., Sieo
(village), Loei (district and province), Thailand, Nov. 23, 1953,
Dr. Boonsong Lekagul and Robert E. Elbel, collectors. Nos. 40189-
40190 (RE 3236), Ban Muang Khai (subvillage), Tha Li (village),
Tha Li (district), Loei (province), Thailand, Dec. 10, 1953 (ap-
prox. 600m. elev.) Dr. Boonsong Lekagul and Robert E. Elbel,

Fig. 4. Rana kuhli Schlegel. EHT-HMS No. 31738 Ban Na Muang, Na Haeo
(village), Dan Sai (district), Loei (province), Thailand. Actual snout-vent
length, 72 mm.

1058 The University Science Bulletin

collectors. EHT-HMS Nos. 31738-31739 (RE 4018), Ban Na Muang
(subvillage), Na Haeo (village), Dan Sai (district), Loei (prov-
ince), Thailand (approx. 1780 m. elev. and in the same range as
Phu Nam Lang (mt.) but farther north), Sept. 29, 1954, Robert E.
Elbel, collector.

This common form is characterized by a wide, rather depressed
head crossed by a white or cream bar. Taylor ( 1934 ) has inadvert-
ently recorded this frog from "Chiang Mai," Thailand, as Rana
corrugata.

Rana cubitalis Smith

(Fig. 5)

Rami cubitalis M. Smith, Jour. Nat. Hist. Soc. Siam, vol. 2, 1917, p. 277 (type
locality, Doi Nga Chang, Thailand); Rec. Ind. Mus., vol. 26, 1925, p. 138
(tadpole); Boulenger, Rec. Ind. Mus., vol. 20, June, 1920, p. 138, 139
( redescription of $ type from "Doi Nga Chang, Thailand, 1600 ft. elev.");
M. Smith, Bull. Raffles Museum, no. 3, Apr. 1930, pp. 103-104, fig. 5
( [arm, showing gland] mountains of Nakon Sritamarat, Thailand; Karen
Hills, Burma).

Three specimens of this handsome frog (EHT-HMS Nos. 31747-49
[RE 3986]) are in the collections. They are from Ban Na Muang
(subvillage), Na Haeo (village), Dan Sai (district), Loei (prov-
ince), Thailand, approx. 1780 m. elevation. Sept. 25, 1954, Robert
E. Elbel, collector. One is a male, two are females.

The male seems to differ from the females not only in the
presence in the glandular development of the first finger and the
large gland on the forearm, but also in having the fine granula-
tions of the back interspersed with numerous granules and larger
pustules. The sides are covered with very numerous pustules each
in turn covered with numerous fine spinous granules (ten to forty).
The granules and pustules on the femora tend to form longitudinal
rows while on the tibia they tend to unite into elongate glandular
folds. There is no distinct tarsal fold but a slight rounding elevated
ridge follows the inner edge of the tarsus and there is a very slight
outer line of minute granules continued onto the edge of the fifth
toe; two strong metatarsal tubercles are present. Four or five
vomerine teeth are present on ridges lying between the choanae.
The ridges are separated from each other by a distance equal to
the length of one ridge, and from the choanae by an equal distance.
The vocal openings are small and situated in the floor of the mouth
behind level of tongue.

The color of the male is very light brownish-gray with a brown
blotch indicated in the tympanic area and a brown stripe present

Contribution to Herpetology of Thailand 1059

below the canthus rostralis. The anterior part of the narrow dorso-
lateral glandular ridge has a blackish brown border. The limbs
have only a suggestion of transverse markings. The venter is

Fig. 5. Rana cubitalis M. Smith. EHT-HMS No. 31748 9 Ban Na Muang,
Na Haeo (village), Dan Sai (district), Loei (province), Thailand. Actual
snout-vent length, 76 mm.

smooth or with only fine transverse wrinkles. On the posterior
face of the thighs there are granules or areolae.

The females, much larger and gravid, have the dorsum fawn
to buff-brown with a brown line strongly defined in the loreal region
and on the tympanum. A brown line is present on the under-

1060 The University Science Bulletin

side of the arm near its insertion. A few flecks of brown are on
the lower jaw and a few scattered brown spots are present on the
sides and along the dorsolateral glandular fold. They are pro-
nounced on the anterior face of the thigh, tibia and tarsus. The
foot and underside of tarsus are dark lavender-brown. The venter
and the underside of the thighs are uniform light (cream?). The
chin and throat have scattered dark pigment in one female, while
in the other the entire ventral surfaces of the body and limbs are
immaculate.

We do not find the openings of the palatal glands on the palate
in front of the vomerine teeth in either sex.

Annandale (Mem. Asiat. Soc. Bengal, vol. 6, 1917, pp. 141-142,
fig. 5 [head] ) gives a brief description of frog specimens from Pegu,
lower Burma, under the name of Rana leptoglossa Cope. From the
figure given we would guess that it is actually a member of this
species which was described in 1917.

Rana nigrovittata (Blyth)
(Figs. 6 and 7)

Lymnodytes nigrovittata Blyth, Jour. Asiat. Soc. Bengal, vol. 24, 1855, p. 718
(type locality, Mergui, Tenasserim).

The following specimens, collected by Robert E. Elbel and Boon-
song Lekagul, are referred to Rana nigrovittata: KUMNH Nos.
40182-40184 (RE 3420), Ban Sang Kho, (subvillage), Khok Phu
(village), Sakon Nakhon (district and province), Thailand, Feb. 9,
1954, elev. approx. 500 m.; No. 40188 (RE 3259), Phu Kading (mt.),
elev. 1045m., Si Than (village), Wang Saphung (district), Loei
(province), Thailand, Jan. 4, 1954, EHT-HMS No. 31772 (RE
3820), Phu Phan (int.), elev. 500m., Sakon Nakhon (district),
Sakon Nakhon (province), Thailand, June 29, 1954.

This last specimen, a male, differs from the descriptions of this
species in having large moundlike areas, resembling paratoids,
between the tympani, narrowly separated from the orbits, and
caused seemingly by hypertrophy of muscles below the skin. We
give it herewith a detailed description. However, it seems to agree
reasonably well with R. nigrovittata Blyth.

Diagnosis: A pair of well-defined dorsolateral folds; interorbital
space wider than upper eyelid; an internal vocal sac, the openings
small, puckered, very far back in mouth; a well-defined gland on
humerus near the insertion of the arm; skin minutely and uni-
formly granular except on posterior part of rump, where part of

Contribution to Herpetology of Thailand

1061

Fig. 6. Rana nigrovittata (Blyth). KUMNH No. 40188, Phu
Kading (mt. ), 1045 m. elev., Si Than, Wang Saphung (district), Loei
(province), Thailand. Actual snout-vent length, 57 mm.

the granules are larger; toes three-fourths to four-fifths webbed,
the toes with moderately large dises.

Description of species: Head obtusely pointed the width of the
head distinctly greater than its length; canthus rostralis indicated,
rounded; loreal region concave; nostril about equidistant between
eye and the median point on upper lip; interorbital area wider
than the upper eyelid; tympanum large, distinct, superficial, not
covered by skin, its greatest diameter (6.2 mm.) much less than

1062

The University Science Bulletin

length of eye (9.1 mm.), separated from the eye by a distance of
four millimeters; no distinct gland below tympanum (however,
the skin on upper jaw is thickened somewhat and now softened).

Fig. 7. Rana nigrovittata (Blyth). EHT-HMS No. 31772,
Phu Phan (mt. ), 500 m. elev., Sakon Nakhon (district and
province), Thailand. Actual snout-vent length, 70 mm.

Tongue elongate, notched, with two rounded processes behind,
free for about one fourth of its length; choanae lateral; two bony
ridges at anterior level of the choanae extending back of their
posterior level, separated from choanae by a distance equal to
their distance from each other, bearing two vomerine teeth on
the posterior end of each ridge. A median denticulation on lower

Contribution to Herpetology of Thailand 1063

jaws; no lateral denticulations but a thickening of the anterior part
of the jaw is evident. First finger longer than second; subarticular
tubercles large. Three large metacarpal tubercles, the two outer
partially fused posteriorly; four supernumerary tubercles; leg
elongate, the tibiotarsal articulation reaches the tip of the snout;
no dorsal fold; two metatarsal tubercles the inner twice as long
as wide, the outer distinct, rounded; no fringe on outer side of
fifth and inner side of first digits; disks well-developed, edged by
a groove; outer metatarsals separated by a web; toes from three-
fourths (on inner) to four-fifths webbed on outer fingers; sub-
articular tubercles large.

Skin above on head and body with very small uniform granules
but becoming a little larger on rump; a few larger tubercles on
back part of upper eyelid; no distinguishable fold behind tympanum;
sides indistinctly granular, the granules of variable size; chin and
venter smooth; area on posterior face of thighs granular; most of
ventral surface of thighs glassy smooth; upper surface of tibia and
tarsus with fine spiny tubercles, often pearl-tipped. A pair of
dorsolateral glandular folds.

Color, in preservative: Above uniform brown, the swollen areas
on occiput distinctly lighter; sides of snout dark brown; a lateral
brown stripe, lighter back of eye, darkest along outer edge of the
dorsolateral fold; arms dark brown above with some brown spots
with lighter background on posterior ventral surface; leg with irreg-
ular dark bars above composed of coalescing spots; back surface
of femur brown reticulated with light brown; chin and throat,
venter and underside of limbs with fine brown pigment; the spot-
ting on the legs extends onto the undersides; palms light, soles
blackish brown; upper jaw with a brownish-gray stripe extending
to below tympanum.

Measurements in mm.: No. 31772: snout to vent, 70; width of
head, 31; length of head, 27; arm, 42; leg from vent, 113; tibia, 38;
foot, 51.

Remarks: The three additional specimens listed are females and
lack the humeral gland. Below the tympanum are two distinct
glandular areas. In these specimens, a groove, midway between the
anterior level of the choanae and front of the palate, marks the
outlets of the palatal glands. We cannot find this in the male
specimen, and it may represent another significant difference in
populations.

2—8050

1064 The University Science Bulletin

Rana erythraea (Schlegel)

(Fig. 8)

Hijla erythraea Schlegel, Abbildungen neuer oder unvollstandig bekannter
Amphibien . . ., 1837, p. 27, pi. 9, fig. 3.

Eleven Rana erythraea (EHT-HMS Nos. 31681-31691 [Lot RE
4292] were collected at Ban Nong Wai ( subvillage ) , Na Phung
(village), Dan Sai (district), Loei (province), Nov. 17, 1954, by
R. E. Elbel. All are of medium size, 35 to 42 mm. in snout-vent
length. The green dorsal color has disappeared but the dorsolateral
cream stripes covering the broad dorsolateral glandular fold are
strongly in evidence.

The species attains a length of 78 mm. for females; 42 for males.
It is widespread in the lowlands in southeastern Asia and Indonesia.
The senior author has reported it on certain of the Philippine Islands.

Rana chalconota (Schlegel)

Hi/la chalconota Schlegel, Abbildungen neuer oder unvollstandig bekannter
' Amphibien . . ., 1837, p. 24, pi. 9,* fig. 1.

A single poorly preserved specimen of Rana chalconota from Ban
Chawang (village), Chawang Sieo (district), Nakhon Si Thammarat
(province), Thailand is in the collection. It bears the KUMNH
number 40007 (BL 20094). It was taken Feb. 1954 by Dr. Boon-
song Lekagul.

Rana sp.

Nine tadpoles of an unidentified Anuran, presumed to be a Rana
were taken on Khao Sawan (mt. ), approx. 600 m. elev., Sieo
(village), Loei (district and province), Thailand, by Dr. Boonsong
Lekagul and R. E. Elbel, Nov. 29, 1953. The lot is numbered
EHT-HMS No. 31841 (RE 3173). While these have been some-
what discolored by preservatives a median lighter muscular area be-
ginning in the occipital region is evident. Near the beginning of
the tail are two somewhat diagonal, rather large discrete brown spots
closely approximated. A transverse bar or spot is present somewhat
in advance of these and a median dark spot is present in the occipi-
tal area. The frontal region may have a few darker flecks or spots.
A few small brown spots occur on the caudal fin and tail. The
spiracle is sinistral; the anus dextral.

The mouth is surrounded by a papillate fringe. At the anterior
median point is a short curved tooth series. The upper beak is of
blackish horn as is the lower. Both are denticulate. On each side

* Van Kampen (1923) regards fig. 3 of pi. 50, the illustration of this species (not
fig. 1, pi. 9).

Fig. 8. Bona erythraea (Schlegel). Upper EHT-HMS No. 31684.
Actual snout-vent length, 41 mm.; lower EHT-HMS No. 31689, actual
snout-vent length, 40 mm. Both from Ban Nong Wai, Na Phung,
Dan Sai, Loei (province), Thailand.

1066

The University Science Bulletin

of the upper beak are five series of denticulations, diminishing in
length posteriorly. The beak on the lower jaw is narrower than
upper. On each side and extending somewhat posterior to it are
five rows of denticulations growing longer toward the lower edge of
the frill; on each side are about 25 very short fragmentary rows of
denticulations. At most posterior part of area a median flat pad-
like region edged anteriorly with denticulations.

Micrixalus magnapustulosus sp. nov.

(Fig. 9)

Type: EHT-HMS No. 31838 $ , Ban Na Phua ( subvillage ) , Kan
Luang (village), Na Kae (district), Nakhon Phanom (province),
Thailand, elevation approx. 200 m., collected July 28, 1954, by R. E.
Elbel and Dr. B. Lekagul.

Diagnosis: A diminutive frog with unnotched tongue a little
longer than wide, free laterally and free posteriorly for about one

Fig. 9. Micrixalus magnapustulosus sp. nov. Tvpe
EHT-HMS No. 31838-5, Ban Na Phua, Kan Luang,
Na Kae (district), Nakhon Phanom (province), Thai-
land. Actual snout-vent length, 16 mm.

Contribution to Herpetology of Thailand 1067

third of its length; no trace of vomerine teeth; lower jaw symphysis
with a suggestion of a median denticulation; eye large, longer than
snout (measured mesially); a sinuous fold from corner of eye be-
hind and partly across tympanum to above arm insertion; tympanum
covered with skin but its outline partly discernible; a curving fold
runs from corner of mouth; dorsum and sides with numerous large
pearl-tipped pustules, the skin between them with smaller pustules
or roughened ridges; toes four-fifths webbed, the web reaching at
least the small, slightly rounded tip of the fourth toe.

Description of the type: Tip of the snout broadly oval, the nostrils
slightly elevated, situated about an equal distance from the eye
and the median point on edge of upper lip, separated from each
other by a distance equal to or very slightly less than median length
of snout; width of an eyelid equal to or slightly less than the inter-
orbital distance; tympanum covered with skin but moderately
distinct, its diameter ( 1.5 mm. ) much less than length of eye-open-
ing (2.2 mm.); a sinuous fold from eye runs downward and back-
ward behind tympanum and terminates on shoulder above the in-
sertion of arm; a small curving fold runs back for a short distance
behind mouth-angle; no canthus rostralis; the loreal area slightly
convex rather than excavated; snout extends beyond mouth for
about half a millimeter; eye large, strongly elevated, its length
greater than median length of snout, but equal to a line from eye
to the middle point on upper lip.

Tongue a little longer than wide (3x2.7 mm.), free posteriorly
for more than a fourth of its length, and free laterally; no tongue
papilla, but the surface minutely granular; choanae small, nearly
lateral, not concealed by maxillary shelf when seen from below;
no trace of vomerine teeth; openings of the Eustachian tubes smaller
than choanae; no evidence of vocal slits. The symphysis of the
lower jaws has no denticulate elevation.

Arm short, only the toes reaching beyond tip of snout; first finger
very little longer than second; only a mere vestige of a web, the
edges of the digits not or but slightly ridged on sides. Three small
palmar tubercles the inner and outer a little more distinct than
middle one, the outer elongated and completely separated from the
middle one; four distinct subarticular tubercles.

Leg short, the tibiotarsal articulation reaches the middle of the
eye; toes with only a slight development of the terminal discs, the
web reaching the discs on one or more toes; a well-defined, some-
what compressed, elongate inner metatarsal tubercle; a very tiny

1068 The University Science Bulletin

outer tubercle situated at the terminus of a slight pustular ridge
along outer toe; well-developed subarticular tubercles; a short
diagonal tarsal fold extending less than half length of tarsus; when
legs are folded at right angles to body the heels fail to touch.

Skin on snout and interorbital area with a very indistinct eleva-
tion across head between anterior edges of eyes (the "eyespot" in-
cluded ) ; body with large craterlike pustules, the sides of which are
excavated by numerous "valleys"; pustules pearl-tipped, between
larger pustules are tiny pustules or ridges; sides with less distinct
pustules; legs with numerous smaller pearl-tipped pustules tending
to form longitudinal rows; chin nearly smooth; venter without dis-
tinct granules; underside of thighs smooth except on the posterior
part; posterior face of thighs have a very few scattered granules.

Color in preservative: Generally brown above with two vague
lighter areas on middle of back preceded and followed by slightly
darker areas or lines; upper side of upper arm fawn brown, the fore-
arm banded with brown. Leg, including tarsus, and foot, with nar-
row bands; chin and throat brownish with some lighter flecks; venter
and part of underside of thighs nearly immaculate; soles dark
lavender.

Measurements in mm.: Snout to vent, 16; width of head, 7; length
of head, 6; arm, 8; leg, 22.5; tibia, 7; foot, 7.7.

Remarks. We are unable to determine the age of the specimen
but believe it to be nearly adult, nor can we state certainly the
character of the pupil, but believe it to be horizontal.

Family Rhacophoridae

Rhaeophorus leucomystax leucomystax (Gravenhorst)

(Fig. 10)

llyla leucomystax Gravenhorst, Deliciae Musei Zoologici Vratislaviensis con-
tinens Chelonias et Batrachia, fasc. I, Lipsiae, 1829, p. 26.

Bhacophorus leucomystax Smith, Jour. Nat. Hist. Soc. Siam, vol. 2, no. 3,
p. 229. Flower, Proc. Zool. Soe. London, Nov. 14, 1899, pp. 898-899, pi.
59, figs. 3, 3a (tadpoles).

Polypedates leucomi/.stax leucvmi/.stax Taylor, Proc. Acad. Nat. Sci. Phila-
delphia, vol. 86, 1934, p. 283 (Chieng Mai).

Two specimens of this arboreal species (KUMNH Nos. 40193-94
[RE 3034]) are from Khon Kaen (city and province), Thailand,
Nov. 13-14, 1953, Robert E. Elbel, coll., and one, No. 40017 is from
Thailand without definite locality. Two of them (40193-94) are
marked on dorsum with dark gray or brownish spots on a gray back-
ground; the third, No. 40017, is nearly uniform gray with some very
indefinite mottling. The limbs of all specimens are barred dimly

Contribution to Herpetology of Thailand 1069

with darker color. On the sides there may be a row of dark spots,
as if a dark stripe beginning behind eye and passing through
tympanum and beyond had been broken up. A very narrow cream
line borders the lip in all of the specimens.

The posterior face of the thigh is brown, this color enclosing
cream spots or flecks, while a similar coloration is dimly indicated

Fig. 10. Rhacophorus leucomystax leucomystax ( Gravenhorst ) . KUMNH
No. 40194 Khon Kaen (city and province), Thailand. Actual snout-vent length,
60 mm.

on the front of the thigh and groin areas. A small postcranial fold
is indicated.

The palatal glands open in a transverse line of eight pores about
midway between the anterior point of the palate and the anterior
level of the choanae. The vomerine teeth are 8-10 on two con-
verging ridges beginning on the anterior level of the choanae and
terminating near their posterior level. The subtriangular openings

1070 The University Science Bulletin

of the Eustachian tubes are of smaller area than the choanal open-
ings.

The webbing on the hand is reduced to a vestige between the
first three fingers; no web is evident between the two outer fingers.
The webbing of the toes becomes progressively greater. The two
inner toes are less than half webbed, while the other toes are from
one-half to three-fourths webbed.

The pads have a deep groove around their outer edges and each
is traversed by a groove somewhat posterior to the center on the
underside.

No. 40017 has a more extensive web, the web reaching the base
of the pads on some of the toes.

Family Microhylidae

Microhyla berdmorei (Blyth)
(Fig. 11)

Engystoma (?) berdmorei Blyth, Jour. Asiat. Soc. Bengal, vol. 24, 1856, p.
720 (type locality, Pegu, Burma).

Microhyla berdmorei Parker, A monograph of the frogs of the family Micro-
hylidae. 1934, pp. 127-28 (synonymy and distribution).

Microhyla foivleri Taylor, Proc. Acad. Nat. Sci. Philadelphia, vol. 86, 1934,
pp. 284-86, fig. 1, pi. 17, fig. 2 (type locality, Chieng Mai, Siam).

Microhyla malcolmi Cochran, Proc. Biol. Soc. Washington, vol. 40, 1927,
p. 182 (type locality, Pak Jong, Siam).

A specimen of Microhyla berdmorei (KUMNH No. 40019) is in
the collection from Phatthalung, Thailand, Dr. Boonsong Lekagul,
coll. It is a female with the abdomen distended with eggs.

The following structural characters obtain in this specimen:

Diagnosis: A median groove present on the upper surface of the
widened digital discs producing two separate elevations. Snout
longer than eye; a rounded canthus rostralis; three palmar tuber-
cles, the two outer closely approximated; two small rounded meta-
tarsal tubercles; tibiotarsal joint extends about six millimeters be-
yond the tip of snout; venter and underside of limbs glassy smooth.

Description of species: Tip of snout forming a narrow oval,
projecting beyond mouth 1.8 mm.; nostril equidistant between eye
and the median point on upper lip; loreal region not concave;
tympanum concealed by skin and partly covered by muscles; tongue
narrowed anteriorly, widened posteriorly, free for half its length;
shelf on the inner side of the maxillary joins with its fellow firmly
to form a small anterior platform that does not conceal the choanae
when seen from below; latter small, transversely oval; a rather
broad transverse ridge across palate in front of oesophageal opening.

Contribution to Herpetology of Thailand

1071

Skin smooth above and below ( perhaps more so than normal be-
cause of the distension of the skin caused by the eggs within ) ; the
fingers reach beyond tip of snout when arm is laid forward; first
finger much shorter than second, which in turn is shorter than the
fourth; edges of fingers with slight lateral ridges but web is lacking
except for the merest remnant; subarticular tubercles prominent;

Fig. 11. Microhyla berdmorei (Blyth). KUMNH No. 40187, Phu Kading
(int.), 1045 m., Si Than, Wang Saphung (district), Loei (province), Thailand!
Actual snout- vent length, 39 mm.

palmar tubercles three, the two outer almost united, partially sepa-
rated by a narrow groove.

Terminal pads scarcely wider than fingers, divided by a groove
above, forming two small swellings on dorsal part of pad. Toes
completely webbed, the tips of the toes widened and divided above
by a groove forming two "scalelike" swellings; outer metatarsals
united. Legs long, the tibiotarsal joint extending several milli-

1072 The University Science Bulletin

meters beyond tip of snout; heels overlap ten millimeters when legs
are folded at right angles.

Color: Purplish above becoming lavender-brown on sides. There
is trace of a bar between eyes which begins a darker area that nar-
rows on occiput, widens again on shoulders and then becomes lost
on back. Some black rounded spots above and behind arm in-
sertion; black spots on front of thigh and on the posteroventral
region on tibia; underside of chin heavily mottled with brown; un-
derside of tarsus and foot purplish brown; ventral surface of body
and limbs largely cream. A triangular black mark in anal region.

Measurements in mm.: Snout to vent, 41; width of head at tym-
panum, 12.3; length of head, 16; arm, 20; leg, 77; tibia, 27; foot, 34.

A second specimen KUMNH No. 40187 (RE 3254), also a gravid
female, agrees in most details with the preceding. It is from Phu
Kading (int.), 1045 m. elev., Si Than (village), Wang Saphung (dis-
trict), Loei (province), Thailand, Jan. 1, 1954, R. E. Elbel and
Dr. B. Lekagul, colls.

Kalotila pulchra pulchra Gray

Kaloula pulchra Gray, The zoological miscellany, 1831, p. 38 (type locality,
"China"); Taylor, Proc. Acad. Nat. Sci. Philadelphia, vol. 86, 1934, p. 284,
pi. 17, fig. 1 (Chieng Mai, Siam).

Callula pulchra Gunther, Reptiles of British India, 1864, p. 437 (Siam, etc.);
Boulenger, Catalogue of the Batrachia Salientia and Ecaudata in the collec-
tion of the British Museum, 2nd ed., 1882, pp. 170-171, figs. (Siam, etc.);
Nieden, Das Tierreich, Anura II. Lief, 49, 1926, pp. 22-23, fig. 18; Jour.
Nat. Hist. Soc. Siam, vol. 2, no. 1, June, 1916, pp. 40-41; idem, vol. 2, No. 3,
May, 1917, 226-231 (common and widely distributed in Siam).

Kaloula pulchra pulchra Parker, A monograph of the frogs of the family Micro-
hylidae, London, 1934, pp. 84-86 (Bangkok, Chantaboon; literature list and
synonymy ) .

A large specimen of Kaloula p. pulchra (KUMNH No. 31516)
from Rat Buri, Thailand was collected by Dr. Boonsong Lekagul.
The typical coloration is present. It measures 67 mm. in length.

The choanae, due to the extremely short snout, are at the anterior
end of the palate. They are transversely widened and followed by
strong ridges that are almost contiguous on the median line. There
are no maxillary or vomerine teeth.

Ghj))hoglossus molossus Gunther

(Fig. 12)

Glyphoglossus molossus Gunther, Proc. Zool. Soc. London, 1868, p. 483, pi. 38,
fig. 1 (type locality, Pegu).

The strongly truncate snout of this curious burrowing frog differ-
entiates it from most of the Microhylidae, which are characterized,
for the most part by narrow, somewhat pointed snouts. Their eyes

Contribution to Herpltology of Thailand

1073

are diminutive, their length less than the distance between the eye
and the tip of the snout. However, the upper eyelid is relatively
heavy, overhanging and overlapping the lower. The head itself
is strongly convex above, the occipital region highest, sloping gradu-
ally to the tip of the snout, the median line actually higher than the

Fig. 12. Glyphoglossus molossus Gunther. KUMNH No. 40018 "Thailand."

Actual snout-vent length, 73 mm.

1074 The University Science Bulletin

eyes. Both upper and lower jaws are truncate, giving the head
a very blunt appearance. A rather dim fold crosses the head just
behind the eyes and reaches down to the mouth angle. The
choanae are diagonally elongate, partly divided by a fleshy process
from the anterior edge. While vomerine teeth are absent there are
two small bony bolitoid processes behind, but slightly internal to
the choanae, resembling vomerine teeth. A rounded median papule
lies immediately in front of, and contiguous with the broad trans-
verse fold lying across the palate some distance in front of the oeso-
phageal opening. The tongue is peculiar. It is somewhat rounded
behind with a suggestion of an anterior notch, and on each side
a thickening leaving a median depression. The thickenings narrow
posteriorly, leaving two diverging fingerlike ridges that fail to reach
the back edge of the tongue. The posterior part of the tongue is
free for a little less than a fourth of its length. Openings of the
Eustachian tubes are large and circular. The vocal sac in the males
is evidenced by two slitlike openings, one on each side of the tongue
near the anterior part of the mouth. Externally, a transverse fold
marks the position of the median vocal sac, which has two extensions
running back above arm to shoulder.

The digits on the hand have lateral ridges (or fringes) along
their inner edges, while between the second and third there is a
tiny web remnant. There are four subarticular tubercles followed
by three small tubercles on the metacarpals, and two large palmar
tubercles, the outer elongate extending to the back border of the
palm, the inner shorter and more distinct but larger than inner.

The toes, fully webbed, have their tips somewhat enlarged and
thickened into small terminal pads. There are two or three sub-
articular tubercles on toes, except outer. A large sharp free-edged
shovel represents the inner metatarsal tubercle; the small outer
tubercle is approximated to first. When the limb is adpressed the
anterior edge of the shovel reaches the tip of the snout.

The skin of face, snout, eyelids, anterior part of lower jaw, and
lips covered with tiny firm rounded, sometimes subspinose tubercles.
The body skin is wrinkled sometimes showing indistinct granula-
tion. The rather smooth venter is neither granular nor areolate.

The entire body is discolored by rust and the preserving fluid.

Measurements in mm.: Snout to vent, 60; between nostrils, 5.3;
arm, 31; leg, 81; foot, 36; shovel, 9; tibia, 25.

Remarks: The data have been drawn from KUMNH No.
33519 ^ from Rat Buri, Thailand. A second specimen a female

Contribution to Herpetology of Thailand

1075

containing ovarian eggs is without specific locality other than
"Thailand." This specimen is somewhat larger and the sides and
limbs are flecked and spotted with cream on a black-brown ground
color. Both were collected by Dr. Boonsong Lekagul.

Family Bufonidae
Bufo asper Gravenhorst

(Fig. 13)

Bufo asper Gravenhorst, Deliciae Musei Zoologici Vratislaviensis continens
Chelonias et Batrachia, Fasc. 1. Lipsiae 1929, p. 58.

Two specimens of this large toad are present in the collection.
These are KUMNH Nos. 40005 (BL 20125) and 40006 (BL 20039)

Fig. 13. Bufo asper Gravenhorst. EHT-HMS No. 30304 Km. 16 on
Pahang Road, NW Kuala Lumpur, Selangor, Fed. Malav States. Actual
snout-vent length, 80 mm.

1076 The University Science Bulletin

both from Nakhon Si Thammarat, peninsular Thailand, Sept. 2,

1953, Dr. Boonsong Lekagul, coll.

The specimens are characterized as follows: a broad supraorbital
ridge, often discernible with difficulty, and a short thick supratym-
panic ridge; snout short, bluntly oval, obliquely truncate; loreal
region vertical, not or scarcely concave; interorbital region wider
than an eyelid; tympanum small, distinct; fingers free, toes com-
pletely webbed, the digital tips widened; a strong inner tarsal
ridge and two metatarsal tubercles, the inner largest. Tibiotarsal
articulation reaches to eye; upper surfaces of body with rough,
pyramidlike pustules and tubercles; parotid glands small but
prominent.

Nearly uniform blackish brown; some light flecks on upper lip
and under thighs.

The specimen figured is from Kuala Lumpur somewhat south of
the Thai border. The species reaches a length of 215 mm.

Bufo melanostictus Schneider

Bufo melanostictus Schneider, Historiae amphibiorum naturalis et literariae
. . . fasc. primus, 1799, p. 216 (type locality, "ex India orientali").

This common and widespread toad is represented by numerous
specimens in the collection. The following localities are repre-
sented: EHT-HMS Nos. 31740-41 (BE 3946), Phu Kho (mt.), 522
m. elev., Kan Luang (village), Na Kae (district), Nakhon Phanon
(province), B. E. Elbel and Dr. B. Lekagul colls., July 28, 1954;
Nos. 31742-43 (BE 4261), Ban Nong Wai ( subvillage ) , Na Phung
(village), Dan Sai (district), Loei (province), Thailand, Nov. 15,

1954, approximately 1780 m. elev., (same range as Phu Nam Lang
(mt.) B. E. Elbel coll.; No. 31744 (BE 3833), Phu Phan (mt.),
Sakon Nakhon (district), Sakon Nakhon (province), Thailand, July
2, 1954, elev. 550 m., Dr. B. Lekagul and B. E. Elbel, colls.; Nos.
31745-46 (BE 4030), Ban Na Muang (subvillage), Na Haeo (vil-
lage), Dan Sai (district), Loei (province), Thailand, elev. circa
1780 m. (in same range of mountains as Phu Nam Lang (mt. ) but
farther north,) Sept. 30, 1954, B. E. Elbel, coll.

Two other specimens from Thailand are without locality data,
the tags having been lost. One measures 102 mm. snout to vent.
The specimen is a female, the ovaries filled with small ovarian eggs
and a stomach packed with wingless termites.

Contribution to Herpetology of Thailand 1077

Family Pelobatidae

Eight species representative of this family have been reported
from Thailand as follows: Megophrys nasuta (Schlegel), M.
longipes (Bonlenger), M. aceras (Bonlenger), M. pclodytoides
(Bonlenger), M. carinense (Boulenger), M. feae (Bonlenger), M.
hasselti hasselti (Tschndi), and M. monticola Giinther.

Megophrys hasselti hasselti (Miiller)
(Fig. 14)

Leptobrachium hasselti Miiller in Tschudi, Classification der Batrachier,
. . . 1838, p. 81, (attributed to Miiller MS) (type locality, Java).

Two specimens, a male, EHT-HMS No. 31762 (BE 4078), and
a dried female, 31763 (BE 4078), from Ban Na Muang ( subvillage ) ,
Na Haeo (village), Dan Sai (district), Loei (province), Thailand,
Oct. 4, 1954, B. E. Elbel, coll.

Diagnosis: Head as wide or wider than body; canthus rostralis
sharp; arms and legs weak, slender, the tibiotarsal articulation not
reaching eye; no palpebral appendages; no vomerine teeth; two
moderate palmar tubercles; one metatarsal tubercle, the outer tu-
bercle missing vertebrae procoelous.

Description: Head as wide as long, the canthus sharp, termi-
nating at nostril; nostril a little nearer eye than median point on lip;
loreal region sloping, somewhat concave; snout in front of nostrils
sloping obliquely; interorbital distance once to once and a half the
width of an upper eyelid, and a third greater than distance between
nostrils; tympanum rather small, its diameter less than half the
length of eye-opening, the upper part covered by a sinuous fold
from eye to angle of jaws. Eyes prominent.

Posterior two thirds of the tongue free, notched behind; two
large slitlike openings to the vocal sac far back near mouth angle;
choanae large; no vomerine teeth; openings of the palatal glands
two, lying between inner level of choanae, but slightly in advance
of the choanae.

Skin on dorsum minutely granular or pustular, the pustules larger
and more prominent on sides; chin with small granules; entire
venter with larger granules or areolations; underside of thighs
smooth except for a few proximal scattered granules; a few granules
about vent; a more or less distinct fold on the side, anteriorly touch-
ing, but not continuous with the supratympanic fold.

Arm small, slender; first finger a little longer than second, latter
a little longer than fourth; small subterminal pads but no sub-

1078

The University Science Bulletin

articular tubercles; undersurface of digits with some broken elon-
gate ridges or callous areas that may incorporate the tubercles;
inner fingers and inner edge of outer fingers with a narrow fringe or
ridge, but no web; two prominent palmar tubercles; toes short, one-

Fig. 14. Megophrys hasselti hasselti (Miiller). EHT-HMS No. 31762
Ban Na Muang, Na Haeo (village), Dan Sai (district), Loei (province), Thai-
land. Actual snout-vent length, 46 mm.

third to one-half webbed on inner toes but continued as a fringe or
ridge to the terminal pad. A well-defined inner metacarpal tuber-
cle; no outer tubercle; no tarsal fold or ridge; when limbs are
folded at right angles to body the heels fail to meet; an elongate
ridge on underside of the third and fourth toes, indicated also on
second toe.

Contribution to Herpetology of Thailand 1079

Color in preservative: Above lavender with darker lavender or
purplish; darker irregular spots on back and sides; a dark longi-
tudinal interorbital line; loreal region and lip with three or four
spots, a dark line from nostril to eye, and from eye along the supra-
tympanic fold, widening on the tympanum; arms and legs banded
dark lavender above. Belly and chin lighter (yellowish) with
clouding of lavender, or numerous flecks of brown or lavender.

Measurements in mm.: Snout to vent, 46; head width, 21.5; head
length, 21.5; arm, 29; leg, 49; tibia, 16.5; foot and tarsus, 22.

Remarks: A second specimen is badly dried. It measures 60 mm.
snout to vent.

Megophrys aceras ( Boulenger )

Megalophnjs mantana aceras Boulenger, Fascic. Malay. Zool., I, 1903, p. 131
(type locality, Malay Peninsula ) .

One specimen examined — No. 20113 without a definite Thai local-
ity. Very probably the specimen is from Nakhon Si Thammarat,
collected by Dr. Boonsong Lekagul.

CLASS BEPTILIA

Order Testudines

Family Platysternidae

Platysternon megacephahim Gray

(Fig. 15)

Platysternon megaceplwlum Gray, Proc. Zool. Soc. London, 1831, p. 107; and
Illustrations of Indian Zoology, vol. 2, 1834, p. 62 (type locality, south
China).

A single young specimen from Thailand has lost its tag and is of
uncertain provenance. However, it is practically certain that it came
from Dan Sai (district), Loei (province). The species may be
diagnosed by the following characters: a blunt median dorsal keel
not or but lightly notched; costal plates each with a tubercular keel.
Nuchal scales very small; five vertebrals wider than costals; five
costals, the three posterior smaller than largest marginals; twelve
marginals on each side; serrate behind. Head large; tail elongate,
covered with shields as follows : 28 paired subcaudals -f- 3 single
scales; a dorsal series of 28 widened scales each with a median
dorsal keel except three or four at base; the terminal scute is com-
pressed, equally as long as the five preceding scales; the dorsal series
of scutes separated from the subcaudals by an intercalated series on
the base of the tail extending to the eighth pair of subcaudals; two
or three intercalated scales on sides of elongate dorsal terminal scute.

1080

The University Science Bulletin

Plastron small with six paired plastral elements and a very small
median scale (abnormal?) between the gulars and numerals; bridge
of plastron narrow, separated from marginals by three or four in-
framarginals.

Length of carapace, 50; width of carapace, 44; tail, 52.

Olive brown above with some black spots on the vertebral keels

Fig. 15. Platysternon megacephalum Gray. KUMNH No. 40084, Thailand.
Actual width of carapace, 44 mm.; greatest length, 50 mm.

Contribution to Herpetology of Thailand 1081

and costal keels; blackish spots on anterior costal keels (see figure).
This species is an inhabitant of mountain streams and is said to be
a tree climber!

Order CROCODYLIA

Family Crocodylidae

Crocodyhis siamensis Schneider

Crocodilus siamensis Schneider, Historiae Amphiborium naturalis et literariae,
Fasc. II, 1801, p. 157 (type locality, Siam, based on drawings, fide M.
Smith).

One typical young specimen (KUMNH 40085 [RE 784] ) is in the
collection from Rung Roraphet (lake), Nakhon Sawan (district and
province), Thailand, collected by Robert E. Elbel Feb. 6, 1953.

This species may be distinguished rather readily from Crocodylus
palustris, by having only four maxillary teeth instead of five (all
visible from the outside of the closed mouth); from Crocodylus
porosus by a shorter snout and in having the rows of bony dorsal
plates in direct contact instead of having the bony parts separated
by soft skin. In siamensis there are usually four ( rarely two ) postoc-
cipital scutes, that are normally absent in porosus.

Order SQUAMATA

Suborder SAURIA

Family Gekkonidae

Cyrtodactylus intermedins* (Smith)

Gymnodactylus intermedins Smith, Jour. Nat. Hist. Soc. Siam, 1917, p. 221
( type locality, Khao Sebab, SE Siam ) ; The fauna of British India including
Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, 1935, pp. 44-45,
pl. 1, fig- 1.

A specimen of this species from Phatthalung, Thailand, was ex-
amined in the collection of Dr. Roonsong Lekagul. It is a female
and while agreeing in color markings it lacks the preanal pores of
the male. There is, however, a long series of enlarged femoral
scales extending most of the length of the femur. The essential
characters of the form are: male with a wide-angled series of
preanal pores, a group of enlarged preanal scales and a series of six
to ten enlarged femoral scales; eleven to twelve supralabials and
nine to ten infralabials; a lateral fold of slightly enlarged scales;
belly with 40 to 50 scales between folds. Rody covered with large

* This species formerly was associated with Gymnodactylus. Garth Underwood, Proc.

Zool. Soc. London, vol. 124, pt. 3, 1954, p. 475, revives the generic name Cyrtodactylus

Gray with the type species C. pulchellus, restricting Gymnodactylus, senso strictu to South
America.

1082 The University Science Bulletin

trihedral tubercles separated by small granules; tail with flat scales
and whorls of larger tubercles, while there are transversely enlarged
plates on underside. Grayish or brownish with blackish-brown
cream-edged bands across back, the one on the neck reaching to
eyes; tail banded.

Hemidactylus frenatus Schlegel

Hemidactylus frenatus Schlegel in Dumeril and Bibron, Erpetologie Generale
. . ., vol. 3, 1836, p. 366 (type locality, Java).

The following specimens obtained by Robert E. Elbel are in the
collections: EHT-HMS Nos. 31830-31834 (RE 4971), Phu Phak
Khi Nak (mt. ), near Ban Nam Yen ( subvillage ) , Kok Sathon (vil-
lage), Dan Sai (district), Loei (province), approximately 1300 m.
elev., between the ranges of Phu Nam Lang and Phu Lorn Lo (mts. ),
Mar. 18, 1955; and No. 31835 (RE 3985), Ban Na Muang (sub-
village), Na Haeo (village), Dan Sai (district), Loei (province),
Thailand, approx. 1780 m., in same range as Phu Nam Lang
(mt), Sept. 25, 1954.

KUMNH No. 31428 (RE 640), Banpong (city and district), Rat
Buri (province), Thailand, Apr. 12, 1952; No. 31411, Banpong
(city and district), Rat Buri (province), Feb. 29, 1952; No. 40086
(RE 914), Ban Lat (subvillage), Ban Kaeng (village), Phukhieo
(district), Chaiyaphum (province), Thailand, Dec. 16, 1952, all
collected by Robert E. Elbel.

Cosymbotus platyurus (Schneider)

Stellio platyurus Schneider, Amphib. Physiol., vol. 2, 1792, p. 30, (type locality,
unknown).

A series of specimens of Cosymbotus platyurus KUMNH Nos.
31413-31427 (RE 613, 619, 746) are all from Banpong (city),
Banpong (district), Rat Buri (province), collected by Robert E.
Elbel, Apr. 5, Apr. 8, and June 18, 1952, respectively.

Peropus mutilatus Wiegmann

Peropus mutilatus Wiegmann, Nova Acta Acad. Leopold-Carol., vol. 17, 1835,
p. 238 (type locality, Manila, P. I.).

Four specimens of this species collected by Robert E. Elbel are
in the collection. These are EHT-HMS No. 31829 (RE 4575) from
Ban Muang Khai (subvillage), Tha Li (village), Tha Li (district),
Loei (province), Thailand; No. 31409 (RE 1085), Khorat (city
and province), Thailand, Sept. 10, 1952; No. 31410, Banpong (city
and district), Rat Buri (province), Thailand, Feb. 29, 1952.

Contribution to Herpetology of Thailand 1083

Gekko gecko (Linnaeus)

Lacerta gecko Linnaeus, Systema Naturae, 10th ed., 1758, p. 205 (type locality,
"habitat in Indies").

The following specimens are in the collections: EHT-HMS Nos.
31822-31823 (RE 4079), Ban Na Muang ( subvillage ) , Na Haeo
(village), Dan Sai (district), Loei (province), Thailand, Oct. 4,

1954, R. E. Elbel, coll. and EHT-HMS Nos. 31824, 31825 ( RE 5297),
Phu Namlang (mt.), 1780 m., Ban Khok (subvillage), Na Phung
(village), Dan Sai (district), Loei (province), Thailand, May 23,

1955, Robert E. Elbel, coll.; KUMNH No. 33518 (BL 20024), Khao
Khansong, Sriracha, Chon Buri, Thailand, July 15, 1953, Dr. Boon-
song Lekagul, coll.; and No. 40020 (BL 20024), Ubon, Thailand,
Nov. 9, 1953, Dr. Boonsong Lekagul, collector.

This very widely distributed species is easily recognized by its
bluish (or ultramarine) to lavender color with numerous rusty-red
spots on both upper and ventral surfaces. The first specimen, No.
31822, is the largest we have seen and doubtless is near the maxi-
mum size.

The two largest specimens, Nos. 31822, 31823, have the following
measurements (in mm.) respectively: snout to vent, 185, 176; tail,
179, 150; total length, 364, 326; head width, 47, 42; head length,
54, 52; snout to arm insertion, 62, 60. The femoral pores are, 10-11,
12-11.

Family Agamidae

Genus Draco Linnaeus

Within the territory of Thailand there are more species of these
unusual gliding lizards of the genus Draco, than in any other area
of equal size. In the extreme southern part of peninsular Thailand,
south of the Isthmus of Kra, Thailand shares with the Federated
Malay States many species which have a distribution extending
into Indonesia, some even into the western Philippines. There
are certain other forms that seem to have developed in continental
Asia that occur in the mainland area of Thailand and extend vary-
ing distances down into the peninsula. Thus from the mainland
we have Draco macidatus Gray, D. haasei Boettger, D. whiteheadi
Boulenger, D. divergens Taylor, and D. taeniopteriis Giinther.
Species presumably confined to the southern peninsular area and
the Indonesian-Philippine region are Draco quinqaefasciatus Gray,
D. volans Linnaeus, D. punctatus Boulenger, D. cyanolaemus Bou-
lenger, D. hlanfordi Boulenger, D. formosus Boulenger, D. micro-

1084 The University Science Bulletin

lepis Boulenger, D. melemopogon Boulenger. Another species with
an Indonesian distribution, D. fimbriatus Kuhl, has been reported
by Taylor (1934) from Chiang Mai, Thailand. One might suspect
that still other species from the south have already found their way
farther into northern continental territory. Malcolm Smith (1930)
has suggested the need of more distributional data on these forms
in Thailand. We would further suggest the need for larger series
from everywhere so as to ascertain the amount of variation that
may be expected.

The group, consisting of maculotus, type locality unknown; di-
vergens, type locality, Chiang Mai, northern Thailand; haasei,
Chantaboon, southeastern Thailand; whiteheadi, Hainan Island —
may represent only subspecies of a single species. Until their
distribution and variation is better known, we propose to maintain
them as separate species. Giinther's (1864) figure of maculotus
is of a specimen from the coastal area of Thailand. Malcolm Smith
suggests "Penang" as the type locality. If the type characteristics
of the type specimens of maculotus can be found in a population
from Thailand or Malaya, the type locality should be fixed so that
the species or subspecies may be placed properly and referred to
under the correct name. If this cannot be done a locality must be
fixed arbitrarily.

The following key should be helpful in ascertaining identities of
these interesting lizards.

Key to Thailand species of the genus Draco

1. Tympanum scaly 2

Tympanum naked, lacking a covering of scales 6

2. Larger (to 105 mm. snout to vent); nostril directed upward; no com-

pressed spine on the supraciliary edge; wing membranes red with
a series of five curving blackish bands, each as wide as the red
interspaces; black band across the neck behind the dewlap,

quinquefasciatus
Smaller (to 85 mm. snout-vent length); nostril lateral directed out-
ward; a small compressed spine on supraciliary edge 3

3. A blue spot on each side of base of gular appendage ( dewlap ) 4

No blue spot on each side of base of dewlap 5

4. No distinct row of lateral, conical, or trihedral nuchal scales; a row of

trihedral scales, widely spaced on the base of the wing membrane;
lateral nuchal expansion ("wattles") covered above with scales
larger than largest dorsals, which are twice size of ventrals; base
of dewlap orange without blue marks. Coppery red above with
metallic reflections and black spots; on wings, orange with rounded
dots on proximal half; head scales large, flat, with no or with few
indistinct keels; usually a low nuchal crest of eight scales ... haasei

Contribution to Herpetology of Thailand 1085

Nostril pointing directly outward; dewlap tapering gradually to a fine
point, blackish on anterior part, blue at the end, red behind the
base; no lateral blue marks on each side; upper head scales strongly
keeled; 8-9 supralabials; on each side of back a series of en-
larged keeled dorsal scales; hind limb reaches halfway between
elbow and axilla; wing reddish brown above with dark bars and
small black spots; immaculate beneath whiteheadi

5. Nostril pointing directly outward; upper head scales strongly keeled;

no series of trihedral scales on side of neck, but a series along the
base of the wing membranes; scales on lateral nuchal expansion
about as large as largest dorsal scales; gular appendage often twice
as long as head; dorsal scales but little larger than ventrals; wing
membranes orange with scattered black spots rather uniform in
size; wing membrane yellow below with one or two black spots;

dewlap yellowish, with blue spots at base maculatus

Nostril directed outward and somewhat upward; head and neck
prominently marked with black paired spots or transverse bars;
spotting of wing membrane largely confined to proximal part, not
arranged in rows; underside of yellow wing membrane much
spotted with black; a nuchal crest involving 25 scales; dewlap
yellow with paired blue spots at base, the length more than one
and a half times that of head; loreal region and labials forming a
broad shelf; snout constricted behind nostrils; a row of trihedral
scales from neck along sides of body at base of wing; no distinctly
Y-shaped group of scales, but a median row of three connecting
with a transverse curving row of larger scales on snout; low tail
crest present divergens

6. Nasal scale lateral, the nostril directed outward 7

Nasal scale dorsal, the nasal directed upward; dewlap longer than

head 10

7. No caudal crest evident 8

A low caudal crest present 9

8. Supralabials, 7-10, smooth, unkeeled; a Y-shaped series of scales on

snout; small nuchal crest; dewlap much longer than head; snout

usually equal to or shorter than diameter of orbit volans

Snout equals diameter of orbit; nostril directed outward and slightly
upward; 8-10 strongly keeled labials; dewlap translucent, covered
with large scales; no caudal crest; wing membranes edged with
brick red and with four or more black bands spotted with white;
throat bluish; dewlap yellow cyanolaemus

9. Supralabials, 10-13; small nuchal crest; nostril directed outward and

slightly upward; upper head scales very small; snout longer than
diameter of orbit; wing membranes dark brown with lighter lines;
below dirty gray, immaculate, or with a few scattered spots; chin
and throat bluish with a network enclosing white spots; dewlap

red or salmon pink fimbriatus

Snout equals diameter of orbit; Y-shaped series of scales on snout;
two conical, triangular or spinelike tubercles on end of supraciliary
(orbital) edge; 10-11 upper labials feebly keeled; dewlap of male
about as long as head or shorter; tail crest strong; back and head

1086 The University Science Bulletin

with large black dots; four blotches form a cross between shoul-
ders; wings black streaked with whitish; uniform whitish below;
dewlap and nuchal appendages bright chrome yellow . punctatus

10. Dewlap coal black; a transverse light orange or whitish band across

neck connecting with color of underside of nuchals expansions;
snout shorter than diameter of orbit; 11-15 supralabials; breast and

belly usually with brown spots melanopogon

Not so marked ]1

11. Smaller; snout-vent length less than 85 mm.; snout equal to or shorter

than orbit 12

Larger; 100-125 snout-vent length; snout longer than diameter of
orbit 13

12. Snout shorter than orbit; dewlap not or but little longer than head;

11-14 supralabials; arm extended, the hand reaches beyond tip of
snout; wing membranes with fine black ill-defined transverse bands;
base of dewlap and underside of nuchal expansion purplish red,
with a black spot at base of dewlap in males; head and body

80 mm. microlepis

Snout equals diameter of orbit; 7-9 supralabials; dewlap with very
large scales slightly longer than head, arm reaches well beyond
snout; hind limb to axilla or shoulder wing membranes with five
arched transverse black bands sometimes forked at base; head and
body 75 mm taeniopterus

13. Supralabials, 9-10 keeled; dewlap much longer than head, translucent,

covered with large scales; slight nuchal fold; arm reaches much
beyond tip of snout; wing membranes marbled with dark brown
and with lighter spots and thin white lines; immaculate beneath;

scarlet on underside of nuchal expansions blanfordi

Snout as long as diameter of orbit; 9-11 keeled supralabials; dewlap
a little longer or equal to head length; thin, translucent, covered
with large scales; ann reaches much beyond snout; wing mem-
branes olive, edged with maroon or crimson; five transverse bands
of blackish mottling, often indistinct; throat maroon or crimson in
male, dark green in female formosus

Draco whiteheadi Boulenger

Draco whiteheadi Boulenger, Proc. Zool. Soc. London, Nov. 1899, pp. 956-957,
pi. 66, fig. 1 (type locality, Hainan Island); Smith, The fauna of British
India including Ceylon and Burma; Reptilia and Amphibia, vol. II, Sauria
Feb. 7, 1935, p. 140 (northern Siam).

Diagnosis: Nostril directed laterally, tympanum scaled; head
small, the snout considerably longer than the diameter of the orbit;
8-9 supralabials; dewlap one and one-half times as long as head;
dorsal scales a little larger than ventrals, obtusely keeled; arm
reaches tip of snout; wing membranes brick red above with round
black spots, immaculate below; dewlap blue at end, blackish in
front, red at base; related to D. maculatus but with longer snout,
etc.

Contribution to Herpetology of Thailand 1087

Description of species: Head short, the snout slightly shorter
than the diameter of the orbit; rostral less than twice as long as
wide, with eight glandular pits on front surface, bordered by two
labials and five other scales; nasals craterlike, the nostril directed
outward, separated from the rostral and labials by one row ot
scales; five or six scalerows separate the nasals; snout constricted
behind nasals; a Y-shaped series of scales on snout, the basal part
consisting of two large keeled scales, and the arms of two or three
enlarged keeled scales reaching outward and backward; supra-
orbital areas outlined by semicircular series of enlarged irregularly
keeled scales that are separated mesially by three longitudinal series;
outer row of supraoculars very large, keeled; one (or two) large
keeled supraciliary scales extending half the length of the supra-
ciliary border; on middle of the latter half of the border, a com-
pressed triangular scale, and at the posterior border a large com-
pressed tubercular one, flanked by one or two similar scales; six
enlarged suboculars more or less continuous with a short series
of three pyramidal scales, the last two largest; two or three irregular
canthals; nine loreal series; supralabials ten, all with two to four
pits; an enlarged triangular tubercle between scaled tympanum
and angle of mouth; twin tubercles some distance back of tym-
panum, and another pair above and slightly posterior to tympanum;
"occipital" scale narrow, elongate, flanked by groups of five en-
larged irregular scales on each side; distal to each of the two groups
and slightly posterior is a somewhat elevated group of about six
irregular enlarged scales, separated from each other by four small
or two enlarged scales; on nape at normal point for the beginning
of a nuchal crest are two conical scales surrounded by eight scales,
altogether forming a rosette. Infralabials 8-8, also bearing pits;
mental unpitted, its border about equal to rostral border, touched
behind by two labials and five scales; gular scales minute; male
gular appendage (21 mm.) longer than head (14 mm.), covered
on its distal half by large regular scales; lateral nuchal expansions
relatively small, with larger scales above.

The nuchal crest is not or barely indicated; a series of large
somewhat compressed pyramidal or trihedral scales begins on neck
and curves up above arm, then follows along the base of the wing
membrane to level of thighs, about 16 on neck anterior to wing,
ten or eleven along wing base; a short indefinite row of similar
scales higher up on shoulders; scales of the dorsal rows along
back much larger than the small ventral keeled scales, keeled or
smooth, often with their edges raised suggesting lateral keels. A

1088 The University Science Bulletin

distinct caudal crest; a series of mucronate scales begin on the
dorsal part of the tail separated at first by three scalerows from trie
caudal crest and more posteriorly comes to border the crest scales;
another serrate series that begins on outer edge of the base of the
tail comes later to border the two median ventral series, which
have high keels. A fringe of enlarged scales along the posterior
edge of thigh; front dorsal part of thigh and tibia with enlarged
keeled scales; 27 lamellae under fourth toe; scales on breast, on
dorsal part of upper arm, and on dorsal part of forearm, enlarged,
keeled; posterior edge of forearm with a fringe of larger scales.

Color in preservative: Above brownish, reticulated or dotted with
some lighter marks, and with deep black marks on head and neck;
wing membranes light tan with black marbling or flecking, not
forming bands or rows of spots; wing membrane below immaculate
gray save for a narrow elongate curved black mark near outer
border. Chin darker, speckled with lighter; breast flecked with
brownish spots; tail gray, banded with darker to tip; dewlap
blackish on anterior margins, remainder whitish, probably orange-
red in life; along side of the base of the dewlap are black marks
separating the light color of the underside of the lateral nuchal
expansions from that of the dewlap area.

Measurements in mm.: Snout to vent, 64; tail, 102; length of head,
14; width of head, 10.5; height of head, 8.5; snout to orbit, 5; snout
to tympanum, 12; axilla to groin, 33; arm, 27; leg, 33; length of
gular appendage, 21; width of alar membrane, 22.

Remarks: Malcolm Smith (loc. cit.) has referred D. whiteheadi
to the synonymy of Draco maculatus (Gray), but mentions the fact
that specimens of this form occur in northern Thailand, Hainan and
Tongking. The type is 86 mm. snout to vent, the tail, 148 mm.

Draco haasei Boettger

Draco haasei Boettger, Zool. Anz., no. 433, 1893, pp. 424-425 (type locality,
"Pratchedi Kan Sabab Chantaboon," Siam).

Diagnosis: Belated to D. maculatus Gray but the upper side of
the lateral nuchal expansion covered with scales which are larger
than the largest scales on the back, which in turn are twice the size
of the belly scales; the base of the gular appendage is deep orange,
without blue flecks.

Description of species: Head small. Snout a little longer than the
diameter of the orbit. Nostril lateral, directed outwards. Tym-
panum scaled. Upper head scales large, weakly keeled. A pair of
scales on the supraocular region distinctly enlarged, much larger

Contribution to Herpetology of Thailand 1089

than the supralabials; a distinct compressed or conical scale in the
beginning of the last third of the snpraciliary arch and a second
broader and shorter one at the end of the supraciliary region; eight
supralabials; gular appendage of the male long, nearly twice head
length, covered with scales which are somewhat larger than the
belly scales. Upper side of lateral nuchal expansions covered with
very large, strongly keeled scales, larger than supralabials and the
largest back scales. A very short and low nuchal crest consisting of
eight scales. Scales of the back irregular, of very dissimilar form
and size, the largest reaching twice the size of the sharply keeled
belly scales, and with obsolescent keels or lacking keels altogether.
On each side of the back and especially distinct in the posterior part,
is a row of large trihedral keeled scales separated rather widely.
The arm when laid forward reaches well beyond the tip of the snout;
the hind leg reaches the axilla.

Color: Above coppery red with metallic reflections and marked
with blackish clots, those on the neck arranged symmetrically; a
triangular black interorbital spot; wing membrane delicate orange
color with whitish longitudinal lines and on the proximal half richly
strewn with small round black spots. Underside unicolor and only
on the anterior point of wing membrane one or two black spots.
Underside of the head with brownish reticulation; lateral nuchal
expansion below, deep orange lacking dark spots.

Total length, 178; head length, 14; head width, 11; rump length,
51; arm, 28; leg, 38; tail, 113; dewlap, 23.

The type locality is Chantaboon [Chanthaburi], Siam. It was
taken on a tree trunk near the village of Pratchedi, Kan Sabab,
[Khao Sabap (mt. )] together with Draco taeniopterus Giinther.
The species was collected by Dr. Eric Haase, Director of the Royal
Siamese Museum in Bangkok, who at the same time collected the
following specimens at Bangkok: Lygosoma chalcides Linnaeus,
Dryocalamus davidsoni Blanford, Rana tigerina Daudin, Microhyla
inornata.

[The above is a free translation of the type description.]

Draco macidatus (Gray)

Dracunculus maculatus Gray, Catalogue of the specimens of lizards in the col-
lection of the British Museum, London, 1845, p. 236 (type locality, un-
known ).

Draco maculatus Giinther, Reptiles of British India, 1864, p. 125, pi. 13, fig. C.
( full drawing of a Siamese specimen. Also reports specimens from "Pinang"
and Tenasserim); Smith, Bull. Raffles Mus., Singapore, Straits Settlements,
no. 3, Apr. 1930, p. 21; Fauna of British India including Cevlon and Burma;
Reptilia and Amphibia, vol. II Sauria, Feb. 7, 1935, pp. 138-140, fig. 42.

1090 The University Science Bulletin

Diagnosis: Tympanum scaled; head scales large, partly keeled;
nasal lateral, the nostril directed outward; gular appendage uniform
orange, very long, with a blue spot on each side of the base. A
series of rather flat enlarged scales forming a Y-shaped group on the
snout.

Description of species: (from KUMNH No. 40043 [BL 20008],
Nakhon Si Thammarat, peninsular Thailand), Dr. Boonsong Leka-
gul coll. Head moderately large, the snout as long as the diameter
of the orbit; nasal scales lateral, craterlike, the nostril directed out-
ward; rostral about two and one-half times as wide as high, bordered
by two labials and seven postrostrals; nasal separated from the
rostral by one scale, from its fellow by five scalerows; a median row
of three large somewhat elevated scales on snout; on each side of
the most posterior is a row of three still larger scales which run out-
ward and backward to the upper edge of the orbital rim; three
canthal scales, the last forming part of rim of orbit, very large, dimly
keeled, followed by two supraciliaries; a compressed scale on supra-
ciliary border some distance behind the two; a knoblike scale at
back of the supraciliary (orbital) edge; a rather indefinite row of
scales form semicircles about upper supraocular areas, separated
mesially by two or three scalerows; a partial row of four large
supraoculars and another somewhat smaller row; occipital area
somewhat inflated with two converging heavy keellike ridges on
each side of the median occipital region; scales in occipital region
large, irregular, the "occipital" usually one of the smaller ones; the
keel crossing two large scales on each side; a slight nuchal crest in-
volving 10-12 scales; supralabials, 8-10; four or five slightly enlarged
suboculars separated from the labials by two scalerows; three or four
larger postoculars continuous with a series of three large scales run-
ning back from eye on the temporal region, the last somewhat
pyramidal; a larger moundlike scale on anterior border of scaled
tympanum; a few flat enlarged scales in upper temporal area,
slightly craterlike; a compressed tubercular scale some distance
above and a group of three a little behind the tympanum; mental
bordered by two labials and five postmental scales; one or two en-
larged scales along side of neck; smooth or slightly keeled dorsal
scales nearly equal to the ventrals which are strongly keeled; a row
of enlarged keeled trihedral scales along base of wing membranes;
scales on top of nuchal expansion keeled, larger than dorsals; on
under surface a patch of scales with edges curiously crenulated;
dewlap elongate, the distal part covered with large imbricating
smooth scales; an indistinct or low dorsal crest; scales on breast, on

Contribution to Herpetology of Thailand 1091

dorsum of upper arm and forearm, on anterior dorsal part of thigh
and lower leg, enlarged, keeled; a distinct fringe of large scales bor-
dering posterior side of leg and outer edge of forearm. The keeling
on the tail, owing to its preservation, is difficult to describe (scales
soft and flattened); wing membrane ample; leg reaches a little more
than halfway from elbow to axilla; arm brought forward the fingers
extend beyond tip of snout.

Coloration: Gray to ultramarine with indefinite darker head
marking; the sides of neck and back flecked or reticulated dimly
with darker gray; wing membrane flesh color ( reddish or orange in
life) with very numerous brown or blackish spots or short bars
forming indistinct irregular transverse rows, and longitudinal rows,
the latter indistinctly connected by fine whitish lines; wing mem-
branes gray, immaculate below; tail banded with darker; hind leg
with one or two lighter bands.

Measurements in mm.: Length, snout to vent, 82; tail, 126; head
length, 17; head width, 11; snout to tympanum, 14.5; snout to arm
insertion, 31; axilla to groin, 45; dewlap, 27; arm, 33; leg, 40.

Remarks: Two females, KUMNH Nos. 40044 (BL 20106) and
40045 (BL 20109), Dr. Boonsong Lekagul collector, are from Thai-
land. The tags have disintegrated but they are most probably from
Nakhon Si Thammarat. They cannot be associated certainly with
any particular species having a scaled tympanum. The wing mem-
branes are somewhat reddish or pinkish, the black coloration form-
ing five or six transverse lines, the spots strongly confluent. The
gular areas are probably uniform light orange in life; the dewlap is
short but distinct. While the dorsal nuchal crest is absent, one
specimen has a row of three larger median scales just back of the
occipital region. The caudal crest is wanting but the small occipital
ridges or keels are present. These may represent a subspecific form
as yet unrecognized.

Two specimens, EHT-HMS Nos. 31789 (BE 4971) and 31790
(BE 4971) are seemingly typical. They were taken from Phu Phak
Khi Nak (int.), approx. 1300 m. elev., between Phu Nam Lang and
Phu Lorn Lo (mts.) near Ban Nam Yen (subvillage), Kok Sathon
(village), Dan Sai (district), Loei (province), Thailand, Mar. 18,
1955, B. E. Elbel, collector.

Draco blanfordi Boulenger

Draco major (nee Laurenti) Blanford, Jour. Asiat. Soc. Bengal, vol. 47, 1878,
p. 125 (type locality, forest east of "Tavoy, India").

Draco blanfordi Boulenger, Catalogue of the lizards in the British Museum,
vol. 1, 1885, p. 267, pi. 20 (head); Smith, The fauna of British India in-
cluding Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, Feb. 7,
1935, pp. 141-142, fig. 41, B.

1092 The University Science Bulletin

Two specimens are in the collections, KUMNH Nos. 40042, 40097.
The first has the following characters:

Rostral bordered by seven scales; nasal large, separated from
rostral by a single scale; snout about as long as the diameter of the
orbit; nostril in craterlike nasal, directed upward; tympanum not
covered with scales; scales on head rugose, keeled or ridged, espe-
cially on frontal and snout region; a longitudinal ridge on anterior
frontal region consisting of three or four large scales; on each side of
this the scalerows from the snout curve outward; at posterior end of
the ridge the keels of the scales are transverse or curve outward and
backward; a semicircular scalerow of larger scales outlines the orbit
above, anteriorly bearing high keels directed outward; these two
series separated mesially by three scalerows; supraoculars variable
in size but eight or ten are larger than the rest, all bearing low
keels; "pineal" eye in a tiny scale between a pair of larger scales;
supralabials, 9-11; infralabials, 10-11; edges of lips with very nu-
merous small well-defined pits, which represent gland openings;
three or four irregular canthal scales; a large tubercle at posterior
corner of eye; a row of three scales behind eye, the last largest;
two rows of enlarged subgular scales behind mental separated from
the labials; lateral nuchal expansion ample, the outer scales both
above and below enlarged; gular appendage ( dewlap ) longer than
head, translucent, the tip rounded, covered with scales larger than
ventral scales. Dorsal scales somewhat irregular, smooth or slightly
keeled; a faint trace of a nuchal crest; ventrals larger than dorsals,
with strong keels; an irregular row of enlarged scales or groups of
scales along side near base of wing membranes. Tail strongly com-
pressed with a slight caudal crest and a ventrolateral row of mucro-
nate scales; undersurface of the tail with two rows of very heavily
keeled scales. Arm brought forward, the hand extends beyond the
snout; leg brought forward, the longest toe reaches to about halfway
between the elbow and arm insertion. The wing membranes are
ample, supported by four elongate ribs, and two short ribs an-
teriorly; a slight webbing on back of leg, fringed with enlarged
serrate scales; 28 lamellae under longest toe.

Color in preservative: Generally brown or gray brown; head
grayish, the upper orbital area outlined in dark brown; a median
dark spot between eyes; a pair of occipital dark spots; upper nuchal
region with several small dark flecks; when wing membranes are
stretched under water very dim longitudinal discontinuous lines of
brown can be discerned on the dorsal surface; below uniform gray

Contribution to Herpetology of Thailand 1093

somewhat darker near anterior border (probably yellowish in life);
dewlap whitish, possibly yellow in life; blackish areas on sides of
neck, and a cream area under the nuchal expansion; back nearly
uniform gray-brown without distinct banding or mottling; tail not
banded, lighter than body.

Measurements in nun.: Length, snout to vent, 127; tail, regener-
ated, 117; snout to eye, 10; snout to auricular opening, 23; snout to
arm insertion, 38.5; axilla to groin, 67; arm, 52; leg, 65.

Remarks: The single described specimen is No. 40042 from Nak-
hon Si Thammarat, Thailand, collected by Dr. Boonsong Lekagul.
The species is regarded as rare but it probably ranges over most of
the country. It has been obtained chiefly in peninsular Thailand.

The second specimen of this species in the collection is No. 40097
(RE 1433), from Hinlaem (village), Trakhanun (district), Kanach-
anaburi (province), Thailand, Nov. 6, 1952. Robert E. Elbel and
H. G. Deignan, collectors. This preserved specimen shows the
color as bluish gray-green with some metallic reflection. The wing
membranes are light brown, lined with narrow whitish streaks (in
No. 40042 the white has disappeared leaving the brown color ap-
pearing as dim narrow stripes; when held to the light the membrane
is translucent and the white streaks above are visible through the
wing). Venter light, lighter than the underside of the wing mem-
brane. There is some evidence of mottling under the chin. The
dewlap is creamy in color and semitransparent; there are large
cream-white areas on underside of the nuchal expansions.

Somewhat above each tympanum is an enlarged pointed tuber-
cular scale. Most of the characters listed for No. 40042 likewise
apply to this second specimen.

Draco melanopogon Boulenger

Draco melanopogon Boulenger, Catalogue of the Lizards in the British Mu-
seum, vol. 3, 1887, p. 492 (type locality, Malacca); Laidlaw, Proc. Zool.
Soc. 1901, pt. 1, p. 307; Boulenger, Fascic. Malay Zool., vol. 1, 1903, p.
152; and A vertebrate fauna of the Malay Peninsula from the Isthmus of
Kra, to Singapore including the adjacent islands; Reptilia and Batrachia,
1912, pp. 62-63; Smith, Bull. Raffles Mus., No. 3, April 1930.

Draco nigriappcndiculata Bartlett, Crocodiles and Lizards of Borneo, 1895,
p. 82.

Diagnosis: This species is characterized by a small head, the
snout shorter than the diameter of the orbit; tympanum distinct,
not covered with scales; nostrils in craterlike scales, the openings
directed upward and slightly backward; scales on latter two thirds
of the supraciliary border not enlarged or specialized; arm as long
as leg or nearly so; no Y-shaped grouping of enlarged scales on

1094 The University Science Bulletin

top of snout; no trace of a nuchal crest; a narrow groove between
the elevated orbits on midline of head. The gular appendage
(dewlap) is a deep black color.

Description of species: (KUMNH No. 40047). Head relatively
small; the snout slightly shorter than diameter of orbit, the frontal
region declivous, the area between nostrils flattened; five scales
between nasals; rostral about four times as wide as high, bordered
by two labials and nine postrostrals; nasal high, craterlike, separated
from rostral by one scale, nostril directed upward and somewhat
backward; scales on snout small, subequal; the scales on the front
edge of the bony orbit including canthals and anterior supra-
ciliaries are elongate, heavily keeled, the scales standing nearly
vertical. Supraorbital region strongly elevated, the outer supra-
oculars only slightly larger than other scales, the semicircular scale
series not differentiated clearly; a deep groove between orbits;
occipital region somewhat inflated, the scales larger than the supra-
oculars but very unequal in size; "occipital" with a distinct "eye
spot," as large as any surrounding scales; supralabials, 12-13, each
with a median longitudinal ridge, keel, or series of tubercles, the
lower edges slightly elevated; infralabials, 12-13, each with two
longitudinal ridges; mental subtriangular, as wide as rostral,
bordered by two labials and four or five scales, the outer largest;
no distinguishable rows of enlarged gulars; an indistinct rounded
tubercle at extreme posterior part of the supraciliary edge on the
orbital rim; no trace of a nuchal crest; wing membrane ample;
a row of about seven large distinct keeled scales along basal border
of membrane; scales on the nuchal expansions enlarged above and
below near outer edge; gular appendage of the male elongate,
narrowed towards tip. Arm brought forward, the anterior fourth
of forearm and hand reach beyond snout; leg laid forward, the toes
reach the axilla; a narrow web, or fringe, bordering the posterior
border of the hind leg, is edged with a series of broadened scales;
tail long, without crest, but at base on each side is a row of larger
scales forming a flaring serrate edge; dorsal body scales small,
subequal, feebly keeled or smooth, smaller than scales on ventral
surfaces; very numerous glandular pits in scales or membranes
within mouth near edge of lip.

Color: Generally brownish above on body; wing membranes dark
blackish brown with rows of small white (yellow in life) spots
that form both indefinite longitudinal and transverse rows; no
markings on underside of wing membranes; breast and ventral
surfaces with small brown spots. Chin, underside of neck, and

Contribution to Herpetology of Thailand 1095

arms with some brownish flecks. Head rather light brown. Dorsum
of neck and body with lighter yellowish or whitish flecks or mark-
ings. Underside of hind legs and under tail whitish (yellowish).

Measurements in mm.: KUMNH No. 40047 Phatthalung, Thai-
land, Sept. 15, 1954; No. 40048 Ban Chawang (village), Chawang
(district), Nakhon Si Thammarat (province), Thailand, Mar. 1954;
No. 40049 (exact locality uncertain), 1953, respectively: length,
snout to vent, 80, 82, 73; tail, 91 (R), 153, 146; snout to orbit, 4,
4.5, 3.8; snout to ear, 10, 10, 10,3; snout to arm insertion, 28.5, 28,
25; axilla to groin, 48, 48, 41; height of head, 8, 8, 7; arm, 42, 43, 37;
leg, 41, 45, 42.

Remarks: These three male specimens are the only ones in the
collection; they agree very well in markings and general character-
istics. However, there appears to be some variation in the length
of the snout, the gular area of females is gray, and the arm is as
long as or longer than the leg.

Goniocephalus* armatus armatus** Gray

Again a armata Gray, Zool. Jour., vol. 3, 1827, p. 216 (type locality, Singapore).
Acanthosaura armata Boulenger, Catalogue of the lizards in the British Museum,

vol. 1, 1885, p. 301, pi. 22, fig. 1.
Goniocephalus armatus arma.us M. Smith, Fauna of British India including

Ceylon and Burma; Reptilia and Amphibia, vol. II, Sauria, Feb. 7, 1935,

pp. 157-160.

Although certain adult characters are not now evident, two juve-
nile specimens, from Phatthalung (district and province), largely
on the basis of geographic probability, are assigned to this sub-
species. They are KUMNH Nos. 40053, 40054. They were col-
lected by Dr. Boonsong Lekagul, Sept. 15 and Nov. 1, 1954.

Goniocephalus armatus crucigerus ( Boulenger )

Acanthosaura crucigera Boulenger, Catalogue of the lizards of the British

Museum, vol. 1, 1885, p. 302, pi. 22, fig. 2 (type locality, Tavoy, Tenas-

serim ) .
Gonioccplwlus armatus crucigerus M. Smith, Fauna of British India, including

Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, Feb. 7, 1935,

pp. 160-161, fig. 49.

The following specimens are in the collections: EHT-HMS No.
31645 (RE 5240), Ban Bo ( subvillage ) , Na Haeo (village), Dan
Sai (district), Loei (province), Thailand, May 15, 1955, R. E. Elbel,
coll., ("in the range of Phu Nam Lang (mt. ) but farther north,

* Kaup's original spelling was Gonocephahis (1825), but was corrected to Goniocephalus
by him in 1827.

* Acanthosaura Gray. Malcolm Smith has synonymized that genus with Goniocephalus
since he regards the postorbital spine as not being of generic importance. Until further
studies arf made we will follow this disposition of the species formerly in Acanthosaura.

3—8050

1096 The University Science Bulletin

elev. approx. 1780 m."); EHT-HMS Nos. 31714-16 (RE 3984), Ban
Na Muang (subvillage), Na Haeo (village), Dan Sai (district),
Loei (province), Thailand, approx. 1780 m. elev. ("range of Phu
Nam Lang mt."), Sept. 25, 1954, R. E. Elbel, coll.; EHT-HMS Nos.
31708-31710, same data; EHT-HMS Nos. 31712-31713 (RE 5490)
from Phu Nam Lang (mt. ) 1780 m., Ban Khok (subvillage), Na
Phung (village), Dan Sai (district), Loei (province), Thailand,
June 2, 1955, Robert E. Elbel, coll.; KUMNH Nos. 40099 $ ,
40010 5 (RE 1432-1434), Hin Laem (village), Tha Khanum (dis-
trict), Kanchanaburi (province), Thailand, Nov. 3 and 6 (respec-
tively), H. G. Deignan and Robert E. Elbel, collectors.

The two specimens from Kanchanaburi province are from a point
about 135 km. east of the type locality but from a higher elevation.
In these the nuchal crest is much higher than the dorsal crest (8
mm. -3 mm.). The postorbital and occipital spines measure four
millimeters, while the length of the orbit is approximately nine milli-
meters. There is a faint suggestion of a gular pouch but since the
specimens are soft the extent of the pouch cannot be accurately
determined. The largest of these is 115 mm. snout to vent.

Specimens from Loei (province) in the collection have a rather
typical color pattern. Most of them are subadult. Several show
the tympanum scaled over more or less.

All the specimens have the nuchal crest separated from the dorsal
crest and the scales of the series have broader bases than are indi-
cated in Smith's figure ( loc. cit. ) .

Two or three of the specimens have ovarian or oviductal eggs.
Smith ( p. 158 loc. cit. ) states that ten or twelve eggs are laid,
12 x 20 mm. in size. The eggs are not spindle-shaped.

Genus Calotes

Five species of the genus Calotes are well known in Thailand.
One other, Calotes tnicrolepis Boulenger, may also be present since
it has been taken on both sides of the country in Tenasserim and
Annam. The species may be distinguished by the following key:

Key to the Species of Calotes in Thailand

1. A part of the lateral scales point backwards and downwards; no fold

in front of shoulder on neck 2

Scales on sides all pointing backwards and upwards 4

2. The leg reaches at least to eye; ventral scales larger than dorsals;

generally greenish in life cristatellus

The leg fails to reach the eye 3

3. Scales in 48-56 rows around body at middle; general color brownish

in life; head slender floweri

Scales in 65-72 rows around body at middle; color brownish in
]jfe microlepis

4. No fold or pit in front of shoulder; two separated spines above

tympanum versicolor

An oblique fold or triangular pit in front of shoulder covered with
small granular scales 5

5. A postorbital spine; upper lip usually lacking a light stripe emma
No postorbital spine; usually a light labial stripe present, con-
tinued on neck some distance mystaceus

One curious character of many, if not all, of the species of this
genus is the hairlike structures on the scales. The keels are usually
present and at least over much of the body they are produced into
spines or mucrones which may have a terminal or a lateral crater.
From this craterlike region a very fine hair emerges which may
attain a length of from .2 to .45 mm. in length. In situ the hair
appears to emerge directly from under the mucrone, or if a mucrone
is absent it may emerge from a small pit near the back of the scale.

Calotes floweri Boulenger

Calotes floweri Boulenger, Fauna of the Malay Peninsula . . ., 1912, p. 70
(type locality, Chantabun, SE Siam and Tunong Tahan, Malay Peninsula);
Smith, Jour. Fed. Malay States Mus., vol. 10, 1922, p. 269; Fauna of British
India including Ceylon and Burma; Beptilia and Amphibia, vol. 2, Sauria,
Feb. 7, 1935; pp. 186-187.

Calotes microlepis (nee Boulenger) Boulenger, Jour. Fed. Malay States Mus.,
vol. 3, 1908, p. 66.

We propose to restrict the type locality of Calotes floweri * Bou-
lenger to Chantabun = Chanthaburi, SE Thailand.

A single Thai specimen (EHT-HMS No. 31717) in our collection
belongs to Calotes floweri. The specimen has lost its field tag and
we are uncertain as to its exact provenance.

The head is relatively narrow, its length equal to twice its width.
The frontal region is flat, and the distance from the edge of the
orbit to the nostril is slightly greater than the diameter of the orbit.
The rostral is wide and low, bordered behind by two labials and
five postrostrals. Following these scales on the snout and frontal
area there is a group of keeled scales arranged in a slender Y-shaped
series, the scales forming the branches of the Y continuous with
the two enlarged "semicircular" scale series on the inner borders of
the supraocular region. The two semicircular series are separated
by three scalerows. At the level of the occipital scale there is a

* Smith ( loc. cit. ) calls attention to certain differences between the northern ( Indo-
chinese) specimens and the southern (Malay) specimens.

1098 The University Science Bulletin

transverse row of slightly elevated scales, and a few enlarged scales
on each side of the occipital region. The scales covering the
temporal regions are unequal with one irregular series from the
eye to above the tympanum. The supralabials are twelve, the last
three not clearly differentiated, the infralabials eleven. The mental
is narrow, about as wide as long, followed by two enlarged rows
of six scales each and separated from the labials ( except first ) and
from each other by from one row of scales ( anteriorly ) to five rows
of scales (posteriorly).

The lateral scales are arranged in more or less distinct transverse
series, the scales pointing backward and downward. Dorsal scales
are subequal in size or larger than those on the venter but the
latter are more strongly keeled.

As is typical in Calotes the body is strongly compressed, the hind
limbs slender, the gular pouch small or absent. The tail is definitely
compressed, but swollen and serrate at its base, and the median
dorsal scales somewhat larger than the rest. There are 64 scales
in a transverse row around the body.

Four indefinite darker marks are present on the dorsum, the first
between the shoulders. Radiating streaks from the eye are in
evidence. The venter is light with or without dark marks. The
tail is lighter than the body and barred with bands of brown and
light tan.

Calotes versicolor (Daudin)

Agama versicolor Daudin, Histoire naturelle des Reptiles, vol. 3, 1802, pp. 395-

397, pi. 44 ( type locality not stated by Daudin ) .
Calotes versicolor M. Smith, The fauna of British India including Ceylon and

Burma; Reptilia and Amphibia, vol. 2, Sauria, Feb. 7, 1935 (this is based

on Kuril's report). Malcolm Smith fixes the type locality as Pondicherry,

India ( may therefore be accepted as the "terra typica" ) .

The specimens of Calotes versicolor in the collection are from the
following localities:

KUMNH Nos. 31388 (RE 621), 31389 (RE 615), 31390 (RE 626),
31391 (RE 623), 31392 (RE 722), 31404 (RE 622), Banpong (city
and district), Rat Buri (province), Thailand, Apr. 7-17, 1952; 31393
(RE 593), Latya (city), Latya (district), Kanchanaburi (province),
Mar. 28, 1952; 31394 (RE 612), Ban Tham (village), Tamuang (dis-
trict), Kanchanaburi (province), Apr. 5, 1952; 31395-31396 (RE
962), Khon San (village), Phukhieo (district), Chaiyaphum (prov-
ince), Thailand, Dec. 24, 1952; all collected by R. E. Elbel.

KUMNH Nos. 40027 (BL 20009), 40028 (BL 20019), Nakhon Si
Thammarat Sept. 1953; 40029 (BL 20096), Rayong, Sept. 21, 1954;

Contribution to Herpetology of Thailand 1099

40030-32 (BL 20123) "Thailand"; 40033 (BL 20141), Mae Hong
Son, Feb. 1955; 40034 (BL 20101), 40035 (BL 20102), 40036 (BL
20122), 40037 (BL 20124), 1954-55. Phatthalnng; all collected by
Dr. Boonsong Lekagul.

KUMNH Nos. 40089-90 (BE 3473), 40095 (BE 3551), Phu Lom
Lo (int.), 2100 m. Kok Sathon (village), Dan Sai (district), Loei
(province), Mar. 26, and Mar. 31, 1954; 40092-94 (BE 3175), Khao
Sawan (int.), approx. 600 in. elev., Sieo (village), Loei (district and
province), Thailand, Nov. 29, 1954; all collected by Bobert E. Elbel,
and Dr. Boonsong Lekagul.

Calotes emma Gray

Calotes emma Grav, Catalogue of the specimens of lizards in the collection of
the British Museum, London, 1845, p. 244 (type locality, "Afghanistan");
M. Smith, The fauna of British India including Ceylon and Burma; Reptilia
and Amphibia, vol. II, Sauria, Feb. 7, 1935, pp. 195-197, fig. 55.

This widespread species probably occurs throughout Thailand
wherever suitable forested country exists.

The scales, usually terminating in spines or mucrones, also have
a small fine flexible hairlike projection growing out from a small
cavity just below the terminal spine and extending farther than the
spine. These are usually much finer than human hair and their
length is from .4 to .45 mm. in length.

Specimens of Calotes emma in the collection are from the follow-
ing localities: EHT-HMS Nos. 31667 (BE 5240), Ban Bo (sub-
village ) , "in the range of Phu Nam Lang ( mt. ) but farther north,"
elev. approx. 1780 m., Na Haeo (village), Dan Sai (district), Loei
(province), Thailand, May 15, 1955, Bobert E. Elbel, coll.

Nos. 31674 (BE 3675), Phu Phan (mt.), 550m., Sakon Nakhon
( district and province ) , June 12, 1954.

Nos. 31675 (BE 3918), 31676 (BE 3918), 31677 (BE 3868), 31678
(BE 3868), Phu Kho (mt), 522m., Kan Luang (village), Na Kae
(district), Nakhon Phanom (province), July 17-25, 1954, Dr. Leka-
gul and Bobert E. Elbel colls. No. 31679 (BE 4650), Phu Loin Lo
(elev. 2100 m.), Kok Sathon (village), Dan Sai (district), Loei
(province), Thailand, Feb. 15, 1955, B. E. Elbel.

KUMNH Nos. 40087 (BE 3416), Bang Sang Kho (subvillage),
Khok Phu (village), Sakon Nakhon (district and province), Thai-
land, Feb. 8, 1954; 40088 (BE 3175), Khao Sawan (mt.), 600 m.
Sieo (village), Loei (district), Loei (province), Thailand, collected
by Bobert E. Elbel.

Nos. 40023 (BL 20077), 40024 (BL 20100), 40025 (BL 20125),
Phatthalung (province), Thailand, 1954; 40026 (BL 20088), Ban

1100 The University Science Bulletin

Chawang (village), Chawang (district), Nakhon Si Thammarat
(province), Thailand, Mar. 1954. All collected by Dr. Boonsong
Lekagul.

Calotes mijstaceus Dumeril and Bibron

Calotes mijstaceus Dumeril and Bibron, Erpetologie Generate, vol. 4, 1837, p.
408 (type locality, Burma); M. Smith, The fauna of British India including
Ceylon and Burma; Reptilia and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp.
197-199.

This species is widely distributed in continental Thailand but ap-
pears to be absent in the peninsular part of the country. It reaches
elevations up to and perhaps above 1500 m. elevation.

The specimens of Calotes mijstaceus are from the following locali-
ties: KUMNH Nos. 31399 (RE 719), 31400 (RE 721), 31401 (RE
724), Lam Phaya (village), Nakhon Pathom (district and province),
Thailand, Apr. 17-18, 1952, R. E. Elbel coll. No. 31402 (Y 197),
Boekprai (village), Bangpong (district), Rat Buri (province),
Thailand, May 30, 1952, R. E. Elbel coll. Nos. 31403 (RE 872),
31442 (RE 873), 31443 (RE 893), Non Khun (village), Phukhieo
(district), Chaiyaphum (province), Thailand; Dec. 10, 13, 1952.
R. E. Elbel, collector.

KUMNH 40038 (BL 20026), 40039 (BL 20026), Nakhon Pathom
(province), Thailand, July 18, 1953, Dr. Boonsong Lekagul coll.
No. 40040 (BL 20043), Nakhon Si Thammarat (province), Sept.
1953, Dr. Boonsong Lekagul coll.

Liolepis belliana belliana (Gray)

Uramastyx belliana Gray, Zool. Jour., vol. 3, 1827, p. 220 (type locality, Penang;
based on a drawing by Major-General Thomas Hardwicke [fide Malcolm
Smith]).

Specimens of this large species in the collection are as follows:
EHT-HMS Nos. 31754-31758 (RE 5169), Phu Phak Khi Nak (int.),
elev. approx. 1300 m., near Ban Nam Yen ( subvillage ) , Kok Sathon
(village), Dan Sai (district), Loei (province), "between the ranges
of Phu Nam Lang and Phu Lorn Lo," Mar. 28, 1955, Robert E. Elbel,
collector.

KUMNH Nos. 31380 (RE 556), 31383 (RE 557), Phu Hin Tang
(int.), Latya (district), Kanchanaburi (province), March 25, 1952,
Robert E. Elbel; Nos. 31381 (RE 642), 31384 (RE 643), Wang Pho
(village), Thong Pha Phum (district), Kanchanaburi (province),
Apr. 11, 1952; Robert E. Elbel; Nos. 31385 (Y 11), 31386 (Y 19),
31687 ( Y 13), April 3-6, 1952, Phuchik (village), Pak Tho (district),
Rat Buri (province), Robert E. Elbel; No. 40106 5 , Kantalak (dis-

Contribution to Herpetology of Thailand 1101

trict), Sisaket (province), May 31, 1954, Robert E. Elbel. No.
40041 "Siam," Dr. Boonsong Lekagul, collector.

One large female ( 135 mm. snout to vent, tail, 217 mm. ) has re-
tained the juvenile coloration. There are two broad continuous
lateral stripes; the median stripe is narrower but bifurcates on the
neck, and then the two stripes are broken into elongate spots.
The venter and area under the tail are uniformly light-colored.
The chin and throat are blackish with white flecks except anteriorly.

Physignathus cocincinus Cuvier

Physignathus cocincinus Cuvier, Regne Animal, 2nd ed., vol. 2, 1829, p. 4
( type locality, Cochin China ) .

One young specimen, EHT-HMS No. 31728 (3821) is in the
collection from Phu Phan (int.), 550 m. (104° 05', 16° 55'), Sakon
Nakhon ( district and province ) , Thailand, collected June 29, 1954,
by Dr. Boonsong Lekagul and Robert E. Elbel.

This is the largest of the Siamese agamid lizards. In preserva-
tion the body and limbs have become plumbeous black. Ten dark
brown bands on the lighter ground color of the tail are clearly evi-
dent but the terminal part of the tail is uniform brown, lacking
bands.

The following characters obtain: snout to vent, 144; tail, 370;
femoral pores, 7-6; head length, 42 mm.; width of head, 28 mm.;
supralabials, 13-14; infralabials, 10-10.

Family Varanidae

This family of large lizards is represented in Thailand by five
forms. These are Varanus rudicollis (Gray), Varanus dumerilii
dumerilii (Schlegel), Varanus ftavescens Hardwicke and Gray,
Varanus salvator salvator ( Laurenti ) and Varanus bengalensis nebu-
losus ( Gray ) . The recent revision of the genus by Robert Mertens
(vide infra) shows that each of these is a representative of a dif-
ferent subgenus.

Varanus bengalensis nebulosus (Gray)

Monitor nebulosus Gray, in Griffith's Cuvier 's Animal Kingdom, vol. 9, Synopsis,

p. 27 (type locality, Java).
Varanus nebulosus Smith, Fauna of British India including Ceylon and Burma;

Reptilia and Amphibia, vol. 2, Sauria. Feb. 7, 1935, pp. 403-404, fig. 94

(4).
V[aranus] (Indovaranus) bengalensis nebulosus Mertens, Abh. Senckenb.

Naturf. Ges. Abh., no. 466, 1942, pp. 244-252; pi. 11, fig. 50.

One young specimen, KUMNH 40022 (BL 20002) is in the collec-
tion, from Nakhon Si Thammarat, peninsular Thailand, collected

1102 The University Science Bulletin

Sept., 1953, by Dr. Boonsong Lekagul. In this the nostril is nearer
the eye (5 mm.) than to the tip of the snout (7 mm.); five
of the supraoculars are distinctly widened transversely, four of
them two to three times as wide as long; a distinct longitudinal
groove or depression on the snout; some scales at the tip of snout
and in frontal region larger than the other head scales; dorsal body
scales, as well as ventrals, smooth or only vaguely keeled.

Chin and throat cream, barred with black; some clouding or spots
on side of head and on light supralabial area; body with numerous
irregular transverse series of small cream or white ocelli and some
indistinct dark transverse bands; the ocellated dots become obso-
lete on proximal fourth of the tail; two bands of cream on distal
fourth, only the more distal one complete. Venter with rows of
larger cream dots sometimes more or less connected. The speci-
men measures: snout-vent, 123 mm.; tail, 150 mm.

Family Scincidae

Mabuya macularia (Blyth)

Euprepes macularius Blyth, Jour. Asiat. Soc. Bengal, vol. 22, 1853, p. 652 (type

locality, Rangpur, Bengal ) .
Mabuya macularia M. Smith, the fauna of British India including Ceylon and

Burma; Reptilia and Amphibia, vol. II, Sauria, 1935, pp. 264-266.

Since there is some doubt about the type locality we propose to
fix the type locality at Rangpur, Bengal.

From the variation that obtains in this species it would appear
that several subspecific forms will be recognized eventually. The
last, most drastic treatment of the species is by Malcolm Smith, loc.
cit. It has resulted in placing in it, as synonyms, Euprepes brcvis
Giinther (type locality, Travancore and Anaimalai Hills); Mabuya
madaraszi (Mehely* (type locality, Ceylon); Lygosoma dawsoni
Annandale (type locality, Maddathoray, Travancore) and Mabuya
allapallensis Schmidt, Allapallai Forest, near Chanda, Central Prov-
inces, India.

However, after synonymizing these Dr. Smith recognizes five
forms (numbered from one to five). Four of these are Indian
(India, Pakistan and Ceylon) while one is from southeastern Asia
and the Malay Peninsula.

In the material at hand from Thailand there are three forms that
we are referring to this species. They are distinguished as follows:

* Taylor has revived this name for a Ceylon species. Ceylon lizards of the family
Scincidae. Univ. Kansas Sci. Bull., vol. 33, pt. 2, Mar. 20, 1950, pp. 481-518, text figs,
1-8; A review of the lizards of the Ceylon. Univ. Kansas Sci. Bull., vol. 35, pt. 2, Sept.
10, 1953, pp. 1525-1585.

Contribution to Herpetology of Thailand 1103

Key to the Thailand forms of Mabuya macularia Blyth

1. A postnasal present; no axillary pocket; an area of specialized scales

above ankle, providing a chigger-mite "refuge." A somewhat
darker lateral band, each scale with a small black spot; scales of
the rows below the band each with a similar dark spot, the spots
forming rather straight broken lines. A few tiny blackish spots

with a cream center scattered on rump macularia postnasalis

No typical postnasal present; an axillary pocket present or absent; a
group of specialized scales above ankle providing a chigger-mite
"refuge" 2

2. Four dark dorsal lines on back; an axillary pocket; scales on chin and

throat with brown borders; scales of the dark blackish lateral band
each with a whitish dot; anterior loreal divided leaving a scale
behind supranasal (abnormal?) from 1780 m. elevation.

macularia quadrifasciata
Dorsum olive brown lacking dark lines or spots; no axillary pocket;
no distinct white spots on dark lateral stripe, but some darker
flecks, spots, or marbling, not forming distinct rows.

macularia malcolmi

The following characters are held in common by these forms;
three, five, or seven sharp keels on scales of back and upper parts of
sides ( all three kinds on same specimen ) ; lower lateral scales smooth
or nearly smooth; frontonasal and rostral in contact for a greater
or lesser distance; prefrontals separated narrowly, rarely touching
at one point; first pair of chinshields separated narrowly or touch-
ing at a point; the prefrontals usually touching the second supra-
ocular, occasionally separated from it; supraciliaries normally five,
rarely six; seven supralabials four preceding the subocular labial;
adpressed limbs overlap, the longest toes reaching palm, wrist, or
nearly half-way to elbow.

All forms seen from Thailand have the modified group of scales
just above the ankle. The scales are pointed (often partially erect),
offering refuge to chigger mites.

There are normally thirty scalerows around the body.

The material available does not suffice to treat of the more
western populations of this species. The presence of the "chigger-
mite refuges" on the legs seemingly has not been recorded and
actually may be absent in western forms.

Mabuya macularia quadrifasciata subsp. nov.

Type: EHT-HMS No. 31802 (RE 5621), Phu Nam Lang (mt.),
1780 m. elev., Ban Khok (subvillage), Na Phung (village), Dan
Sai (district), Loei (province), Thailand; coll. June 10, 1955 by
Robert E. Elbel.

1104 The University Science Buixetin

Diagnosis: Four dark dotted dorsal lines on back; scales on chin
and throat with brown borders; most scales of sides each with a
lighter area; an axillary pocket; anterior loreal usually divided; area
of specialized "chigger" scales on leg above ankle.

Description of type: Rostral in contact with frontonasal; supra-
nasals separated; prefrontals separated from each other and from
second supraocular; frontonasal touches frontal; no typical true post-
nasal but first loreal divided transversely leaving a scale behind
supranasal; two or three presuboculars; six enlarged palpebral scales
on eyelid; frontal longer than its distance from tip of snout, and
longer than combined parietals; eight supralabials, five anterior to
large subocular ( one scale on left side abnormal in size ) ; three pri-
mary and three secondary temporals; common frontoparietal suture
two thirds of the length of the scales; one pair of nuchals; infralabials
seven; mental border on lip much larger than that of rostral; a large
azygous postmental; first two pairs of chinshields widened, sepa-
rated narrowly from each other; five supraciliaries, four supraoculars,
two touch frontal; thirty scalerows about middle of body; 15-16
lamellae under fourth toe; a distinct axillary pocket, floored with
tiny scales, infested with chigger mites; tail partly regenerated; ear
small, rounded, less than half size of a dorsal scale; scales on back
quinquecarinate, denticulate behind; on the tail quadricarinate, the
two median keels heavier and more widely spaced; scales on leg
above ankle especially modified, somewhat pointed, semierect, the
area strongly infested with chigger mites.

Color: A dim narrow continuous or dotted dorsolateral light line
bordered above by a row of black flecks covering parts of two
scalerows; two median scalerows each with a more or less con-
tinuous black line, the black extending onto the paravertebral row;
laterally a dim broad dark band on side of body and tail bordered
below on side of neck by a blue-white dotted line that is more or
less continuous with the light line or row of white spots on supra-
labials; most scales of lateral dark stripe and of rows on side below
stripe each with a larger or smaller bluish-white spot or area, pro-
ducing a polka-dot effect; top of head brown; labial sutures brown
or blackish; most scales on chin with brown posterior borders; front
part of chin pure white.

Measurements in mm.: Snout to vent, 57; tail, 70 (regenerated);
head length, 15; head width, 10; snout to ear, 13; snout to arm in-
sertion, 24; axilla to groin, 25; arm, 17; leg, 25.

Remarks: Most of the dorsal scales are denticulated behind, the
number of points depending on the number of keels on the scale.

Contribution to Herpetology of Thailand 1105

It would appear that this, like postnasalis is a mountain form.
The type locality is at an elevation of approximately 5800 ft. The
type locality of postnasalis is at about 8000 ft. elevation. A speci-
men of Mabuya KUMNH 40114 from Phu Lorn Lo (mt.) 2100 m.
elev., Kok Sathon (village), Dan Sai (district), Loei (province),
Thailand, has been badly injured and most scales are missing from
the body. There is an axillary pocket; prefrontals rather widely
separated; no postnasal; 14 lamellae under fourth toe; the first chin-
shields barely touch mesially. Loss of dorsal scalerows prevents
knowledge of the marking on the back but a few scales present on
sides show the light spotting. The first loreal on one side is divided.
We are unable to offer a satisfactory identification of the specimen.
The third form, macularia malcolmi, seems to be a lowland form
also lacking the postnasal.

Mabuya macularia postnasalis subsp. nov.

Type: KUMNH No. 40110. From Phu Lorn Lo (mt.), 2100m.
elev., Kok Sathon (village), Dan Sai (district), Loei (province),
Thailand; collected by Robert E. Elbel, Mar. 30, 1954.

Paratypes: Nos. 40109, 40111, 40112 all with same data.

Diagnosis: A postnasal present; a specialized group of scales on
leg above ankle forming a chigger-mite refuge; no axillary "chigger
pocket." A dark lateral band present bordered below by a light
line; a few scales on rump and those on lower lateral rows with
small blackish spots some of which may have a tiny cream spot in
center; one or two presuboculars.

Description of type: A small species the limbs overlapping when
adpressed; rostral and frontonasal touching; prefrontals narrowly
separated from each other and touching the second supraocular;
supranasals small, separated; anterior loreal single, higher but nar-
rower than second; a true postnasal present; no scale behind supra-
nasal; one presubocular; usually four enlarged palpebral scales;
frontal about equal to its distance to the end of the snout, shorter
than combined length of parietals; one pair of nuchals; seven supra-
labials; three primary and three secondary temporals; frontoparietal
suture two thirds of the length of the scales, longer than the inter-
parietal which separates parietals; eight infralabials; mental border
on mouth wider than that of rostral; five supraciliaries; four supra-
oculars, two touching frontal; 30 scalerows around body; 15-16
lamellae under fourth toe; no shallow pocket in axilla; tail complete
with 71 subcaudals; ear about half size of a dorsal scale, with some
tiny lobules bordering the anterior edge.

1106 The University Science Bulletin

Area above foot on hind limb with specialized pointed scales
strongly infested with chigger mites; scales of body three or five-
keeled, or sometimes seven-keeled, becoming four-keeled on base
of tail, the two inner keels stronger and a little wider apart, the
median keel absent; scales not or scarcely denticulate behind; scales
on sides of neck, on arm and leg, with two or three keels, more dis-
tinct on neck and leg. Ventrals smooth and lower lateral scalerows
smooth or nearly so.

Two pairs of enlarged chinshields touching labials, the first pair
very narrowly separated, the second pair separated by a scale; third
chinshields small, separated from labials by an elongate scale;
median preanals larger than outer ones; subcaudals slightly wider
than adjoining scales.

Color: Above brown or brown olive with a very dim darker
lateral band on side, most distinct on neck, bordered above by a
slightly lighter line scarcely visible behind neck; there are eight
scales on back between darker bands; numerous small dark equal-
sized flecks scattered on rump, each with a tiny lighter middle part;
along the lateral stripe all scales with a discrete black spot; below
this stripe all scales with black dots tending to form rows; one or
two similar lines of dots along tail; a distinct light line from tip of
snout along supralabials below ear (but including lower edge) and
continuing very dimly along side below dark stripe; chin without
dark marks; seven median ventral rows of scales uniform, probably
white or yellow in life.

Measurements in mm. and scale data on type and paratypes of Mabuya macu-

laria po.stnasalis, respectively

Numbers 10109 10110 10111 10112

Length, snout to vent 58 55 56 55

Tail — 80 85 72 broken

Head length 15.5 14 15 —

Head widtn 10.5 10.2 10.2 —

Snout to ear 12.3 11.2 12 —

Snout to arm 20.5 19.8 18 —

Axilla to groin 29 25.5 24.4 25.2

Arm 19 18.2 17 16.8

Leg 25.5 22.5 23 23

Postnasal Yes Yes Yes Yes

Prefrontals separated Yes Yes Yes Yes

Prefrontal touches 2nd supraocular . . Yes Yes No Yes

First chinshields separated Yes No Yes No

Supraciliaries 5 5 5 5

Axillary pocket No No No No

Chigger ' area" on leg Yes Yes Yes Yes

Scalerows 30 30 30 30

Contribution to Herpetology of Thailand 1107

Mabuya macularia malcolmi subsp. nov.

Type: EHT-HMS No. 31774, from Phu Phak Khi Nak (mt.), near
Ban Nam Yen (subvillage), Kok Sathon (village), Dan Sai (district),
Loei (province), Thailand, Mar. 18, 1955. Robert E. Elbel, coll.

Diagnosis: No trace of an axillary pit, no postnasal, but posterior
part of nasal somewhat narrowed; frontal shorter than combined
parietal but equal or a little greater than its distance from snout
tip, wider than the frontonasal, but touching it; second loreal rela-
tively short, much less than twice as wide as first loreal; a chigger
mite refuge on the lower part of leg; leg reaches to elbow when
adpressed; first chinshields separated mesially; transverse rows of
scales, parietal to above vent, 38; from mental to vent, 47; 30 scale-
rows about body.

Color: Above generally olive, each dorsal scale with a slightly
darker central area; head darker olive; a dorsolateral stripe on side
of head and neck continued on to the anterior part of body, and
bordered below by a lighter line from the upper lip; four anal scales
whitish.

Mabuya multifasciata multijasciata (Kuhl)

Scincus maltifasciatus Kuhl, Beitr. Zool. vergl. Anat., 1820, p. 126 (no type
locality stated).

It seems reasonably certain that the specimen described by Kuhl
came from the Dutch East Indies, and perhaps most probably from
Java. In view of the necessity of fixing a type locality so as to study
variation in this widespread species, we propose to fix the locality
at Batavia (Djakarta), Java.

The collection contains several specimens of this large form from
the following localities:

EHT-HMS No. 31803 (RE 3755), Phu Phan (mt), Sakon Nakhon
(district and province), 550 m. (approx. 104° 05'; 16° 55'), June 20,
1954, Robert E. Elbel and Dr. Boonsong Lekagul. No. 31807 (RE
3947), Phu Kho (mt.), 522 m., Kan Luang (village), Na Kae (dis-
trict), Nakhon Phanom (province), (104° 22'; 16° 49'), July 28,
1954, R. E. Elbel and Dr. Boonsong Lekagul. No. 31804 (RE 4485),
Ban Muang Khai (subvillage), Tha Li (village), Tha Li (district),
Loei (province), Thailand, Jan. 15, 1955, elev. approx. 600 m., R. E.
Elbel. 31806 (RE 5245), Ban Bo (subvillage), Na Haeo (village),
Dan Sai (district), Loei (province), Thailand, May 16, 1955, ap-
prox. 1780 m., same mountain range as Phu Nam Lang (int.), but
farther north, Robert E. Elbel.

1108 The University Science Bulletin

KUMNH No. 40055 (BL 20133) Phatthalung (district and prov-
ince), Thailand, Nov. 1954, Dr. Boonsong Lekagul, coll. KUMNH
No. 40056 (BL 20012) Nakhon Si Thammarat (district and prov-
ince), peninsular Thailand, Sept. 1953, Dr. Boonsong Lekagul,
collector.

KUMNH No. 40113 (RE 3484), Phu Lorn Lo (mt.) 2100 m. elev.,
Kok Sathon (village), Dan Sai (district), Loei (province), Thai-
land, Mar. 26, 1954, Dr. Boonsong Lekagul and Robert E. Elbel,
collectors.

No. 31807 is a gravid female. The dorsolateral line is scarcely
discernible in color from the remainder of the dorsum. There are
seven dark-brown dorsal stripes on the lateral edges of the dorsal
scalerows. The outer scalerow does not extend beyond the shoul-
der. On the sides there is a brown stripe in which the outer parts
of the scales are darker than the median parts. On the area above
the arm insertion and posterior to it, the scalerows are directed up-
wards and backwards. Somewhat farther back on the sides of the
body and tail, as well as on the sides of the neck, there are small
bluish-white marks heavily bordered by black above and below.
The sutures of many of the cephalic scales are edged with black.

One of the males, No. 31804, differs in pattern somewhat. There
is a distinct dorsolateral light line bordered above by a light-olive
line, the two covering two scalerows and including a row of small
triangular dark spots, the second upper row bordered by a distinct
black line. The four scalerows separating the black lines display
no black except between the shoulders. The head of this specimen
is gray-olive.

The keels in this form are normally three. Occasionally the
nuchals show four or five keels but sometimes, especially in the
very young and females, the nuchals may be smooth. Most speci-
mens and especially females have the keeling reduced on the sides
and some specimens have only dim keels on the arm. No. 40055
is striking in having the three keels placed very close together near
the middle of the scute leaving the outer third on each side smooth.
Thus the space between the series of keels is occupied by wide
smooth strips.

The following variable characters obtain: scalerows around
middle of body, three with 29, four with 30; transverse rows be-
tween parietals and above vent, 41-43; prefrontals touch; the supra-
nasals separated; the prefrontals touch the second supraocular (50
percent); a postnasal invariably present; lobules invariably on front

Contribution to Herpetology of Thailand 1109

border of ear-opening; the first loreal lower than second; supra-
labials, 7-7; the fifth widened, below eye ( on one side, in one speci-
men, eight supralabials with five preceding the snbocular labial);
four or five large quadrangular scales on the translucent part of the
eyelids; supraciliaries, usually 6-6.

The female contains six nearly full-time embryos each still with a
bit of the yolk present in the yolksack. One embryo measures 35
mm. snout to vent, while the tail is approximately 47 mm. in length.
(No. 31806, a recently born specimen in the collection measures
36 mm. ) The mother has a snout-vent length of 120 mm. while
No. 40056 has a snout-vent length of 125 mm. which is about the
maximum size for the species.

Mabuya longicaudata Hallowell

Euprepis longicaudata Hallowell, Amer. Philos. Soc, ser. 2, vol. 11, p. 77,

pi. IV, fig. 1 (type locality, "Siam." Here restricted to Bangkok, Thailand).
Eumeces siamensis Giinther, Rept. Brit. India, 1864, p. 91 (type locality,

"Siam." Here restricted to Bangkok, Thailand).
Mabuya longicaudata Smith, Fauna of British India including Ceylon and

Burma; Reptilia and Amphibia, vol. 2, Sauria, Feb. 7, 1935, pp. 270-271

( "The whole of Siam" ) .

KUMNH No. 31397 (RE 520131), Ban Lat (subvillage), Ban
Kaeng (village), Phukhieo (district), Chaiyaphum (province),
Thailand, Jan. 31, 1952. KUMNH No. 31398 (RE 520132), Ban-
pong (city and district). Rat Buri (province), Thailand, Apr. 8,
1952; both collected by Robert E. Elbel.

EHT-HMS No. 31792 (RE 3947), Phu Kho (mt), 522 m. elev.,
Kan Luang (village), Na Kae (district), Nakhon Phanom (prov-
ince), Thailand, July 28, 1954; collected by Dr. Boonsong Lekagul
and Robert E. Elbel.

EHT-HMS No. 23 (BL 1979), Bangkok, Thailand; collected by
Dr. Boonsong Lekagul.

Diagnosis: Dorsal scales with two dim or fairly distinct keels, a
median (third) keel dimly evident on some scales; supranasals in
contact; prefrontals in contact and touching the second supraocular;
supralabials seven; 28 scalerows about body; 49 transverse series be-
tween parietals and a point above vent; normally the supranasals
separate rostral from the frontonasal; subcaudals, 127, 128; leg
reaches halfway to elbow of adpressed arm; tail long, often two and
one-half times head-body length.

The general color of all is as follows: there is an olive or olive-
brown dorsum, and a broad brown stripe from eye to base of tail
covering most of three scalerows, bordered above by a dorsolateral

1110 The University Science Bulletin

bluish white stripe covering one or one and one-half scalerows.
The edges of the dorsal scales are slightly brownish and there ap-
pear to be seven dim dorsal brownish lines. On one specimen the
dorsolateral light line is invaded by black above and below leaving
the line bordered above and below by blackish triangular dots for
a part of its length.

All four specimens have 28 scalerows about the middle of the
body, and 49 transverse rows between parietals and a point above
vent. The lamellae under the fourth toe vary between 24 and 26.

We propose to fix the type locality of this species at Bangkok,
Thailand.

Measurements of Mabuya longicaudata

EHT-HMS KUMNH KUMNH EHT-HMS
Number 23 31397 31398 31792

Snout to vent 92 115 95 —

Tail 227 tip miss, tip miss. 233

Head length 22 30 21 —

Head width 15 17.5 14 —

Snout to ear 19 24 18 —

Snout to arm 31 36 32 —

Axilla to groin 46 52 48 44

Arm 29 33 30 28

Leg 39 43 39 41

Eumeces quadrUineatus ( Blyth )

Plestiodon quadrilineatum Blyth, Jour. Asiat. Soc. Bengal, vol. 22, 1853, p. 652,
tvpe locality, China [Hong Kong?]; Taylor, Univ. Kansas Sci. Bull., vol. 23,
1935 ( 1936, Aug. 15), pp. 452-457.

Eumeces quadrivirgatus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1860,
p. 502 (type locality, Hong Kong).

Diagnosis: Palatine bones not meeting on midline of palate; eye-
lids scaly; nostril in single nasal; supranasals present; two fronto-
parietals; pentadactyl limbs; pterygoids toothed; two dorsolateral
stripes separated by two scalerows and the edges of two others;
three pairs of nuchals.

A specimen of this species, EHT-HMS 31791 (RE 5621) from
Phu Nam Lang (int.), 1780 m. elev., Ban Khok (subvillage), Na
Phung (village), Dan Sai (district), Loei (province), Thailand,
June 10, 1955, Robert E. Elbel, coll., has the following characters:
the portion of the rostral visible above equals the greatest length of
the internasals. The latter scales are diagonally elongate touching
the postnasal behind the nasal, and in contact mesially. The fronto-
nasal is about a fourth wider than long, touching the first loreal.
The prefrontals are slightly wider than long, forming a common
suture. The frontal is a little longer than its distance from the

Contribution to Herpetology of Thailand 1111

tip of the snout, as well as longer than the distance from the nuchal.
Its posterior border is rounded rather than angular. The fronto-
parietals are separate, forming a median suture less than a third
the length of the scales. The interparietal is nearly a half longer
than wide, and is enclosed narrowly by the parietals which are
followed by three pairs of nuchals, of nearly equal size.

Three supraoculars border the frontal, the second longer but
not quite as wide as third while both are larger than the first. The
nostril is pierced in a small nasal. Also, there are present a post-
nasal, two loreals of which the anterior is higher but only about half
the length of the second, seven supraciliaries, two presuboculars,
four postsuboculars, eight supralabials of which five are in front of
the subocular, and seven infralabials. The mental has a larger labial
border than the rostral and is followed by two unpaired postmentals,
which in turn are followed by three pairs of chinshields only the
first of which is in contact. The last pair of chinshields are fol-
lowed by an elongate postgenial scale bordering labials, three times
as long as wide. The first labial touches the nasal and postnasal.
The lower eyelid has an opaque area covered with quadrangular
scales larger than those adjoining them. One primary temporal,
two large secondary, and two smaller tertiary temporals are present.
The ear-opening is small, with three lobules on the anterior border.
There are 22 scalerows around the middle of the body. Dorsal
scales are large, only two scalerows and the edges of two others
lying between the white dorsolateral lines. There are two enlarged
preanal scales. The subcaudals, much widened, number 87, the
last 16 being on a regenerated part of the tail; 53 dorsal scales from
parietals to above the vent. The adpressed limb reaches approx-
imately to the elbow. A greenish or bluish white stripe following
the second scalerow passes from the rostral onto tail where it
widens and with its fellow practically encompasses the tail poste-
riorly. A lateral line from the labials passes through the lower part
of the ear and continues as far as the groin.

Remarks: The longitudinal scalerows vary from 20 to 22; the
transverse rows from parietals to above vent vary between 48 and
54. There are seven or eight supralabials, and the second post-
mental may be single or divided. The digits of the leg may reach
to the base of the fingers or to the elbow of the arm when adpressed.

The amount of known variation in the characters of this species is
relatively small, but it may prove to be geographical. One speci-
men partially digested was taken from the stomach of a Natrix

1112 The University Science Bulletin

in the collection. The subcaudals are much widened and thus
differing from most Eumeces. There is only a single divided scale
following the vent. The widened subcaudals in the skinks are
usually separated from the vent by several rows of small scales.

Dasia olivacea Gray

Dasia olivacea Gray, Ann. Mag. Nat. Hist., vol. 2, 1838, p. 331 (type locality,

Penang I. Malaya).
Euprepis olivaceus Giinther, The fauna of British India, 1864, p. 80, pi. X,

fig.D.

Diagnosis: Dorsal scales finely striate, the striae somewhat sinu-
ous; keels almost obsolete anteriorly; on posterior part of back scales
bearing three, five, or seven dim keels; ear-opening greatly reduced,
almost covered by large flat scales; second supraocular widely
separated from the prefrontal; prefrontals not in contact; 28 scale-
rows around body; 43 transverse scale series from parietals to above
vent; no postnasal; two anterior temporals. Transverse irregular
black bands on body, each scale bearing a bluish-white spot or fleck.

Description of species: Rostral large, its labial border distinctly
less than that of rostral, moderately in contact with the frontonasal;
latter broader than long (4.9 mm. x 3.7 mm.), and narrowly in con-
tact with the frontal; prefrontals large, separated, touching both
loreals; nasals single, much longer than wide, rectangular; supra-
nasals separated, narrower than nasals; six supraciliaries; four supra-
oculars, three touching frontal; interparietals form a suture for half
of their length; interparietal separates the parietals; a pair of
nuchals; two presuboculars; two or three postsuboculars; seven
supralabials, the fifth below middle of eye, in direct contact with
eye granules; four large palpebrals on lower eyelid; two anterior
temporals, three secondary and four tertiary temporals; six infra-
labials; mental followed by a large azygous postmental which is in
turn followed by two pairs of chinshields, the first pair in contact;
third chinshield small, separated from labials by an elongate scale;
frontal longer than combined parietals and longer than its distance
from tip of snout. Scalerows, 28 around middle of body; about 44
transverse rows between parietals and a point above vent; about ten
preanal scales, the two median enlarged; ear very small, the scales
in front large, flat, nearly covering opening; the scales over dorsum
and sides of body very finely striate, the striae forming wavy rather
than straight lines — from 50 to 100 on each scale; scales of body
and sides with low flat-topped keels varying from two to eight on
scales in various parts of body, most numerous on scales of the

Contribution to Herpetology of Thailand 1113

rump; scales of the posterior part of forearm usually dimly tri-
carinate, the fine striae continued on scales to the tips of the digits.

When limbs are adpressed the digits overlap; palm covered with
rounded moundlike scales; the lamellae broad, not keeled; no en-
larged scale at wrist; scales on sole similar to those of palm but a
pair of enlarged scales at heel; subdigital lamellae broad, unkeeled,
19 under longest toe; tail broken, the subcaudals widened; scales
on venter smooth.

Color: Body brownish gray with a series of dark irregular trans-
verse bars across the neck and body each scale of which has a
silvery or bluish-white fleck or spot; head with small somewhat
symmetrical marks, pronounced in occipital region and on the supra-
oculars; limbs with some darker flecks suggesting bands.

Measurements in mm.: Snout to vent, 100; tail broken and partly
lost; head width, 14.5; head length, 24.2; snout to ear, 20; snout to
foreleg, 33; axilla to groin, 50; arm, 28; leg, 36.

Remarks: The specimen described is KUMNH No. 40057 from
Nakhon Si Thammarat, peninsular Thailand, south of the Isthmus of
Kra. While the type locality is relatively near (about 200 mi.
south and on a coastal island) there are certain differences. The
ear differs in its general make-up and appearance; the frontonasal is
longer than broad. We have found no mention in the literature of
the curious striation of the scales of this species. It is possible that
it has been overlooked, yet it is also possible that it differs from the
typical species of Dasia in this character.

It may be noted that Mabuija longicaudata has developed ex-
tensive striation on scales while most species of Mabuya lack the
striae altogether.

Sphenomorphus indicus indicus (Gray)
(Fig. 16)

Hinulia indicum Gray, Ann. Mag. Nat. Hist., ser. 2, vol. 12, 1853, p. 388
(type locality, "Himalayas." The type locality is restricted to "Sikkim,
Himalayas" by Stejneger, loc. cit.).

Sphenomorphus indicus Stejneger, U. S. Nat. Mus. Bull. 58, 1907, pp. 216-218,
pi. 17, fig. 1 (from Boulenger).

Lygosoma indicum Boulenger, A vertebrate fauna of the Malay Peninsula from
The Isthmus of Kra to Singapore . . . Reptilia and Batrachia, 1912,
p. 87, fig. 24; Smith, Jour. Nat. Hist. Soc. Siam, vol. 2, June, 1916, p. 55
("Hills near Pre [northern Siam]"); Bull. Raffles Mus., no. 3, April, 1930,
pp. 33-34.

This species unfortunately had no type locality given except
"Himalayas." The generalized locality "Himalayas" has been fixed
by Stejneger (loc. cit.) to Sikkim, Himalayas, which should suffice

1114

The University Science Bulletin

Fig. 16. Sphenomorphus indicus indicus (Gray).
KUMNH No. 40108, Phu Lorn Lo (nit), 2100 m.,
Kok Sathon, Dan Sai, Loei (province), Thailand.
Actual total length, 199 mm.

Contribution to Herpetology of Thailand 1115

since the species occurs in this rather small Indian State, lying in be-
tween the countries of Nepal and Bhutan. From here the species
ranges across southern China to Formosa, and south through Burma,
Assam, Thailand, Indo-China and parts of Malaya. Knowledge as
to whether or not it has developed subspecific forms in this vast
range other than now recognized must await a review of consider-
able material. It is not a common species and seemingly it is not
evenly distributed in Thailand. It is believed that S. i. zebratus
Boulenger (1887) from Tenasserim should be recognized as a sub-
species.

Two specimens of S. i. indicus are in the collection (KUMNH No.
40107-40108 (RE 3536-3567) from 2100 m. elev., on Phu Lorn
Lo (mt.), Kok Sathon (village), Dan Sai (district), Loei (province),
Thailand, March 30, 1954, Dr. Boonsong Lekagul and Robert E.
Elbel, collectors.

Diagnosis: No supranasal; frontonasal touches rostral; divided
frontoparietals; prefrontals touch; nostril in a single nasal, no post-
nasal; lower eyelid scaly; parietals enclose interparietals; only four
supraoculars; a dark lateral band from eye to groin, indefinite on
lower side.

Description: (KUMNH No. 40107): Rostral convex, not flat-
tened, forming a broad suture with the frontonasal which is much
broader than long; prefrontals separated; the frontal shorter than
the combined parietals, nearly a half longer than its distance from
tip of snout; parietals enclose interparietal; a pair of nuchals sepa-
rated from secondary temporals by a pair of tertiary temporal scales;
four supraoculars, last touching large upper secondary temporal;
eight or nine supraciliaries; two loreals, second widest, slightly
lower than first; no postnasal; two preoculars, lower large; nine
scales form a pre-, sub- and postocular series extending to the tem-
poral; two or three small scales between last supraocular and the
postoculars (sometimes described as small supraoculars); supra-
labials seven, the fifth and sixth below eye; infralabials seven; a pair
of superimposed anterior temporals; a pair of larger secondary tem-
porals, the upper touching the parietal, three tertiary, the upper
lying behind parietals; a pair of large preanals.

Mental wider than rostral; postmental single, followed by two
large chinshields in contact and these by a second pair separated by
one scale, a third pair following separated by five scales; the chin-
shields all in contact with the labials; ear-opening moderate (1.6
mm. x 2 mm.), granules but no lobules on anterior border; lower

1116 The University Science Bulletin

eyelid scaly without transparent disc; scalerows around body, 36;
transverse scale series from parietals to above vent, 78; lamellae
under fourth toe, 19-20 (95 subcaudals in No. 40108).

Color: Above olive-gray on head and dorsum; a dark lateral band
from eye, passing above ear-opening then widening and continuing
to groin, the upper edge well defined and bordered by a dim light
line, its lower edge ill defined, generally covering much of three
scalerows. All ventral surfaces immaculate except for some dim
gray areas under sides of neck. A few dark flecks on dorsal part
of neck, shoulder region, and a few near base of tail. Tail repro-
duced.

Measurements in mm. (of KUMNH Nos. 40107, 40108, respec-
tively ) : Snout to vent, 95, 79; tail, — , 120; snout to ear, 17.5, 14.5;
head length, 20, 18; head width, 15, 12.5; snout to arm insertion, 30,
26; axilla to groin, 53, 43; arm, 22, 20; leg, 33, 32.

Remarks: The second specimen agrees with the one described in
most characters, but the supralabials have more or less distinct black
spots on the sutures. The tail has a narrow dorsolateral black line
to tip, and the olive back has more numerous but smaller dark flecks.
On the median dorsal line of the tail there is a row of black dots.
Toward the end of the tail most of the scales have blackish dots.

The hind limb in the young and in males reaches to the elbow; in
large adult females they barely overlap. Both specimens have
nuchal scales separated from the large secondary temporals by an
enlarged tertiary temporal scale.

Boulenger states "nostril in a nasal or between nasal and post-
nasal." We would suspect that the latter condition betokens a
different form from indicns. There is not a trace of "large spots in
flanks" mentioned by Boulenger (Cat. vol. 3, pp. 241-242, 1887).

Sphenomorplms maculatus (Blyth)

(Fig. 17)

Lissonota macula' a Blyth, Journ. Asiat. Soc. Bengal, vol. 22, 1853, p. 653,
(type locality, Assam).

A specimen of an adult gravid female lizard (EHT-HMS No.
31781 [BE 3887] ) has the following characters that ally it with
maculatus:

The rostral has a distinct concavity just in front of its straight
transverse suture with the frontonasal. The prefrontals are widely
separated. Of the five supraoculars, three touch the frontal. There
are 34 scalerows around the middle of the body. While there are
no distinct nuchals the median scales on the neck are usually
widened.

Fig. 17. Sphenomorphus maculatus (Blyth). Upper
figure, EHT-HMS No. 31784, actual total length, 161
mm. Lower figure, EHT-HMS No. 31781, actual total
length, 176 mm. Both from Phu Kho (int.), 522 m.
elev., Kan Luang, Na Kae, Nakhon Phanom (province),
Thailand.

1118 The University Science Bulletin

A stripe from the rostral passes through the eye, then widens and
continues along the side. It is black anteriorly with a few small
white spots, then more olive posteriorly, tending to form four nar-
row dark lines and three olive lines. The stripe continues along
the side of the tail to tip but there it is nearly a uniform olive,
edged with a scalloped brown line. A discreet white stripe from be-
hind eye passes above the ear to the hind leg where it is interrupted,
but continues somewhat indistinct for some distance on the tail.
Below this the labials and sides of neck have some small dark spots
or flecks, while below the lateral white line there is an indefinite
darker stripe, broken anteriorly. The chin, venter, underside of
limbs, and the subcaudal area are immaculate whitish-flesh. The
dorsum is olive with dim paired dark dots on either side of the
median line, and its outer border has a more or less distinct very
narrow lighter line.

In a young specimen from the same locality and lot, the median
paired dark spots are somewhat indistinct but the dorsal olive color
is separated from the deep black lateral stripe by a narrow indefinite
light line. A lateral white line is present, the edges irregular but
clearly defined. Below this white line the black is arranged in a
row of 18 spots, and the series is more or less continuous in front
of thigh and tibia.

The two males (Nos. 31783-31784) lack the distinctive white
lateral stripe. Their legs are longer and they show some other small
differences in the color pattern. Seemingly the subcaudal count
is lower and scalerows around the middle of the body higher (38).

The leg reaches to the elbow on the female distended with eggs.
In males the leg reaches the axilla or a little beyond.

The specimens are from the following localities: EHT-HMS No.
31780 (RE 3755), Phu Phan (int.), 550 m, Sakon Nakhon (prov-
ince and district), Thailand, (104° 05', 16° 55'), June 20, 1954, R. E.
Elbel and Dr. Boonsong Lekagul, colls. 31781-31784 (RE 3887),
Phu Kho (mt. ), 522 m., Kan Luang (village), Na Kae (district),
Nakhon Phanom (province), Thailand, (104° 22', 16° 49'), July 19,
1954, Dr. Boonsong Lekagul and R. E. Elbel, colls.

Measurement in mm. and scale data for Sphenomorphus maculatus

Snout-

White

Number

vent

Tail

Total

Dorsals

Sub-

Scalerows

lateral

and sex

length

length

length

to vent

caudals

middle

line

31781 9 .

64

112

176

79

132

34

yes

31783 5 .

57

111

168

77

124

38

no

31784 5 ..

59

102

161

75

123

38

no

31782? .

28

80

38

yes

31780 9

60

80

36

yes

Contribution to Herpetology of Thailand 1119

Genus Leiolopisma Fitzinger

The species of Leiolopisma * may be recognized by the following
characters: rostral and frontonasal in contact; parietals enclose
interparietal; four large supraoculars; six or seven supralabials; no
supranasals; lower eyelid with an undivided transparent disc; limbs
pentadactyl; prefrontals touch (in Thailand forms) usually making
a substantial suture.

The following species are represented in the collection: Leio-
lopisma eunice Boulenger, L. rupicola Smith, L. siamensis sp. nov.
Besides these the following occur:

Leiolopisma kolitaoensis Cochran (spots on throat; 30-32 scale-
rows; one or more nuchals; 16 lamellae under toes; limbs of adults
not touching when adpressed ) . An island form which may or may
not occur in peninsular Thailand (see Malcolm Smith, Fauna Brit.
India, Bept. Amph. vol. 2, p. 297.

Leiolopisma v. vittigerum Boulenger [=? Leiolopisma pranensis
Cochran] (with a median white or golden stripe from snout to tail,
usually bordered with black; in juveniles often three light stripes
present on body; 28-30 scalerows).

Leiolopisma doriae Boulenger (three or four pairs of nuchals;
28-32 scalerows; six scalerows across back; dorsals much larger than
laterals; prefrontals barely touching).

? Leiolopisma mclanostictum Boulenger (34-38 scalerows; back
dotted with black; a broken lateral stripe present).

Leiolopisma tavesae Smith (26 scalerows; longitudinal streaks on
back and neck.

Leiolopisma r. reevesi Gray (eight scalerows above on back; 28
scalerows around body; several pairs of nuchals present).

Leiolopisma rupicola (M. Smith)

(Fig. 18)

Lygosoma rupicola M. Smith, Jour. Nat. Hist. Soc. Siam, vol. 2, 1916, p. 46
( type locality, "Chong Kae, near Paknampo, central Siam" ) .

Two specimens, EHT-HMS Nos. 31786-31787 (BE 3887) are in
the collection from Phu Kho (nit. ) 522 m. elev., Kan Luang (vil-
lage), Na Kae (district), Nakhon Phanom (province), Thailand,
July 19, 1954, Dr. Boonsong Lekagul and Bobert E. Elbel, collectors.
These specimens have the following characters:

* Malcolm Smith, in a paper published in Records of the Indian Museum, vol. 39,
pt. 3, Sept. 1937, pp. 213-234 (p. 223), treats of Leiolopisma as a section under the
genus Lygosoma (if we interpret his arrangement correctly). He states, "Under Leio-
lopisma are included a number of species that cannot clearly be assigned to any genus
. The majority of the Leiolopismids have no doubt been derived from Lygosoma
by the simple change in the eyelid, others from Emoia by loss of the supranasal shield
through fusion with the nasal."

1120

The University Science Bulletin

The general markings are shown in the figure. The color is vari-
able light brown on dorsum and sides growing much lighter towards
the venter. The spots are black, the lateral stripe broken by white
or yellow spots. The venter is yellowish or whitish. The number

Fig. 18. Leiolopisma rupicola (M. Smith). Left figure, EHT-
HMS No. 31787, actual total length, 45 mm.; right figure, EHT-
HMS 31786, actual total length, 87 mm. Both specimens from
Phu Kho (mt. ), 522 m. elev., Kan Luang, Na Kae (district), Nak-
hon Phanom (province), Thailand.

of scalerows around the middle of the body is 34, the lamellae
under fourth toe 18 or 19. There are 117 subcaudals in No. 31786,
the median series slightly enlarged. Under a lens, scales indefinitely
roughened on surface by from ten to twenty fine irregular longi-

Contribution to Herpetology of Thailand 1121

tudinal lines covering all but front edge of the scale; venter scales
smooth while the greater part of the tail is likewise smooth.

The snout-vent length of No. 31786 is 32 mm., the total length 87
mm.; of No. 31787, 29 mm.; tail lost. The leg reaches the elbow
of the adpressed arm or a little farther.

Leiolopisma eunice Cochran
(Fig. 19)

Leiolopisma eunice Cochran, Proc. Biol. Soc. Washington, vol. 40, Dec. 2, 1927,
pp. 187-188 (type locality, Bang Suk near Pak Jong,* Thailand), USNM
No. 72180 (Dr. Hugh Smith, coll.); Proc. U. S. Nat. Mus., vol. 77, 1930,
p. 18, fig. 3 (Doi Angka, 700 ft., and Pak Jong * ).

Leiolopisma reevesi reevesi Smith, The fauna of British India including Ceylon
and Burma; Beptilia and Amphibia, vol. 2, Sauria, 1935, pp. 295-296.

Diagnosis: A small form reaching 56 mm. snout to vent; anterior
loreal much higher and narrower than second; eight supralabials,

Fie. 19. Leiolopisma eunice Cochran. EHT-HMS
No. 31785, Ban Muang Khai, Tha Li (village), Tha
Li (district), Loei (province), Thailand. Actual
snout-to- vent length, 43 mm.

the sixth and seventh largest, below eye; one large anterior tem-
poral; three secondary and three tertiary temporals; one distinct pair
of nuchals; ear three or four times size of the palpebral disc; 32
scalerows around middle of body; two much enlarged anals flanked
by two smaller scales; toes of adpressed limbs overlap two to three
millimeters; four supraoculars.

A specimen (EHT-HMS No. 31785 [RE 4390] ) from Ban Muang
Khai (subvillage), Tha Li (village), Tha Li (district), Loei (prov-
ince), Thailand, elevation approx. 600 m., Dec. 29, 1954, R. E. Elbel,
coll., has the following characters: rostral convex reaching back to

* Pak Jong = Pakchong, Doi Pia Fai Mts.

1122 The University Science Bulletin

a line connecting the front edge of nostrils, broadly in contact with
the frontonasal, which is much wider than long; prefrontals broadly
in contact, each nearly as large as the frontonasal; frontal slightly
longer than its distance from the tip of snout, more than a third
shorter than the combined parietals; frontoparietals paired, each
larger than the interparietal; latter enclosed by parietals; temporal
formula, 1 + 3 + 4; supralabials eight, five preceding first subocular
labial; a row of small suboculars below eye, the series broken
medially so the first subocular labial is in contact with small gran-
ules of eyelid; a transparent eye-disc one third or one fourth of the
area of ear-opening; labial borders of mental and rostral nearly
equal; infralabials six, the last nearly concealed by last supralabial,
first small; a large undivided postmental; three pairs of chinshields,
the first in contact, second separated by a scale, third pair by three
scales, fourth pair small separated by seven scales; all four of the
chinshields touching labials. Most of the tail missing. Transverse
scale series from parietals to above vent, 70; the distance between
snout-tip and arm-insertion ( 15.4 mm. ) is contained in the axilla-
to-groin distance (20.2 mm.) 1.31 times. There are 17 lamellae
under the fourth toe; eight scalerows between dorsolateral lines.

The color above gray-brown with very numerous fine black flecks,
the larger flecks tending to be more numerous towards the middle
line of the back; the dorsolateral black line, bordered by a dim
line of very light tan, is more or less broken up by cream vertical
bars into about twenty spots; on neck the line is nearly solid black
posteriorly but it narrows and becomes broken behind eye; a black
line across edges of eyelids, and a dark line across loreal region to
rostral; arm and leg marbled or reticulated above; sides with a few
fine scattered flecks; sutures of labials with indefinite dark spots.

Remarks: Dr. Cochran has suggested that Leiolopisma eimice
has several pairs of nuchals. In the several species the scales on
the neck are widened in the four dorsal rows. This is true in the
described specimen but only the first pair are modified enough to
be considered nuchals. Dr. Cochran's statement is "several pairs
of feebly enlarged nuchals." The presence of 34 scalerows would,
in my opinion, preclude association with the Leiolopisma reevesi
from Ningpo. Malcolm Smith, however, has referred L. eimice
to the synonymy of L. reevesi reevesi as a supspecies.

Contribution to Herpetology of Thailand 1123

LeioJopisma siamensis sp. nov.

(Fig. 20)

Type: EHT-HMS No. 31788 (RE 3936) from Phu Kho (mt),
522 m., Kan Luang (village), Na Kae (district), Nakhon Phanom
(province), Thailand, (104° 22'; 16° 49'), collected by Dr. Boon-
song Lekagul and Robert E. Elbel, July 26, 1954.

Diagnosis: Prefrontals meet at a point; nostril in a nasal; no supra-
nasals; large undivided disc on lower eyelid, less than one third of

Fig. 20. Leiolopisma damensis sp. nov. EHT-HMS No.
31788, Kho Mt, 522 m., Kan Luang, Na Kae (district),
Nakhon Phanom (province), Thailand. Actual total length,
116 mm.

the size of the ear; tympanum deeply sunk, three lobules barely
indicated on border; sixth labial separated from large upper tem-
poral by two temporal scales; fifth labial separated from it by one
temporal scale; snout to arm insertion 19.5 mm.; axilla to groin, 24
mm.; 34 scalerows about middle of body; hind leg reaches elbow
of arm; 20 lamellae under fourth toe; subcaudals not or but slightly
widened except on regenerated part, lateral body scales as large as
dorsals; a double frontoparietal; enlarged preanals; small median
pair of nuchals, flanked by a larger one laterally; ear-opening 1.5

1124 The University Science Bulletin

mm. x 1.1 mm. No black markings of any sort; brown above, grow-
ing light low on sides; venter, chin, throat, subcaudal region,
and lower part of sides immaculate, without trace of pigment.

Description of species: Rostral large, length of the part visible
on top of snout minutely less than its distance from frontal, broadly
in contact with frontonasal; latter twice as wide as long, touching
anterior loreal and nasal laterally; prefrontals meeting mesially at
a point as do the frontal and frontonasal; frontal a third longer
than its distance from tip of snout, shorter than interparietal and
frontoparietals together; parietals broadly in contact behind inter-
parietal; nostril in a single nasal, without a postnasal; two loreals,
the anterior narrower and higher than second; two unequal super-
imposed preoculars; four large supraoculars, two touching frontal;
8-9 supraciliaries, first largest; an undivided transparent disc in
lower eyelid; seven supralabials, the fifth and sixth largest below
eye but separated from orbit by two irregular rows of suboculars;
a wedge-shaped scale inserted partially between the fourth and
fifth supralabials; two anterior temporals followed by two secondary,
the upper very large; tertiary temporals four, the upper lying be-
hind the parietal, separated from its fellow by a pair of nuchals;
anterior temporals separated from eye and last supraocular by two
or three rows of postoculars; tympanum deeply sunk; ear-opening
large, rounded, oval, about size of eye-opening, three or four times
size of the disc in the eyelid; about 65 scales from parietals to above
vent; 34 scalerows about middle of body; subcaudals not or but
slightly wider than adjoining scales except where tail is regenerated
in which part the subcaudals are greatly widened.

Color: Generally light brown above with a faint indication of a
lighter line on dorsolateral nuchal area separated from its fellow
by six scalerows and disappearing back of shoulder; indication of
a faint darker line from behind eye, passing above tympanum and
above arm below dorsolateral line; faint indications of darker areas
continue some distance on sides. Venter and all undersurfaces im-
maculate whitish lacking any trace of pigment except on hand and
foot. Top of head a little darker than body.

Measurements in mm.: Snout to vent, 51; tail (regen.), 65; snout
to arm insertion, 19; axilla to grain, 24.5; length of head, 13; width
of head, 7.3; length of arm, 14; leg, 20.

Remarks: This species belongs in that group of the genus hav-
ing the lateral and dorsal scales subequal, and a very large ear-
opening which approaches the open eye in size. It differs from

Contribution to Herpetology of Thailand 1125

L. reevesi in having the limbs overlap a greater distance ( to elbow ) ;
more scalerows around body, no black marks on body; a pair of
nuchals flanked by an enlarged outer pair of tertiary temporal
scales, and three anterior temporals, instead of two.

From melanostictum it differs in having a greater overlap in the
adpressed limbs, much enlarged preanals, the absence of distinct
black markings on body, and in the distance from snout to arm in-
sertion. The distance from axilla to groin is slightly less proportion-
ally.

Leiolopisma etmice Cochran, a species described from Bang Suk
near Pakchong, Siam, differs in having a short snout, the limbs when
adpressed failing to meet and the distance between axilla and groin
is double that from tip of snout to arm insertion.

Suborder SERPENTES
Family Typhlopidae *

Typhlops diardi mulleri Schlegel

Typhlops mulleri Schlegel, Abbildungen amphibiens, 1839, p. 39, pi. 32, figs.

25-28 ( type locality, Padang, Sumatra ) .
Typhlops diardi Schlegel, Abbildungen amphibiens, 1839, p. 39 (type locality,

Indes Orientales); Smith, The Fauna of British India, Ceylon, Burma.

. . . Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 51-52, fig. 15

( synonymy and literature ) .
Typhlops schneideri Jan, Iconographie generale des ofidiens, tome 1, livr. 9,

1864, p. 20, pi. I, fig. 3, (type locality, Bangkok, Siam).
Typhlops siamensis Giinther, Reptiles of British India, 1864, p. 175, pi. 16,

fig. D (3 figs.) (type locality, Siam).
Typhlops diardi mulleri Brongersma, Zool. Meded., Leiden, vol. 17, 1934,

p. 193.

The single specimen of Typhlops diardi mulleri in the collection
is KUMNH No. 31436, from Chantuk (village), Sidiu (district),
Khorat (province), Thailand; Robert E. Elbel, coll., Aug. 31, 1952.
It agrees with the subspecies mulleri as to the color markings.
There are 26 scalerows around the body near the middle, the eleven
dorsal scalerows being dark brown, the remaining fifteen rows
cream-white in color. Farther forward the white color impinges on
the adjoining row until only nine dorsal rows are dark brown, and
seventeen are cream-white. There are 305 transverse scalerows
from the mental to the vent, and eight rows on the tail, counting
the terminal spine. The total length is 332 mm., the tail 4.5 mm.

The nasals are incompletely divided. They form a very narrow
contact behind the rostral (which is not mentioned as a character

* We do not follow the recently proposed idea that the Typhlopidae are not serpents.
It is not improbable that the Serpents are diphyletic in origin.

1126 The University Science Bulletin

in either of the subspecies). The eye is distinct and lies wholly
below the surface of the ocular scale. All anterior head scales have
minute tubercles. The anterior nasal scale covers a depression
below the nostril.

Held in certain lights the scales show a most elaborate criss-
crossing striation on the undersurface of their exposed free parts.
This is more evident when the scale is removed and viewed from
below. This condition has also been observed in other species of
TijphJops.

Family Dipsadidae

Parens macularius Theobald

(Fig. 21)

Pareas macularius Theobald, Jour. Linn. Soc, vol. 10, 1868, p. 54 (type local-
ity, Martaban, South Burma).

The single specimen of Pareas macularius in the collection is
EHT-HMS No. 31798 $ . It is definitely from Thailand but the
exact locality is uncertain owing to the disintegration of the tag.

Diagnosis: This species, which resembles Pareas margaritophoras
Jan somewhat in color and markings, differs from it, however, in
having a more compressed body, keels on the seven median scale-
rows, and a somewhat larger series of ventrals and subcaudals.
It seems to replace P. margaritophorus in Burma, Bengal, northern
Thailand and Indo-China. The following characters are evident in
the specimen at hand :

Description: Bostral large, slightly wider than high, its upper
edge a straight transverse line; internasals more than twice as wide
as long, notching upper line of the nasal, narrower than prefrontals;
latter wider than long, touching preocular and loreal, entering orbit;
frontal short, six-sided, its length greater than its width, longer than
its distance from tip of snout, shorter than parietals; latter scales
longer than their distance from tip of snout; nostril in a single nasal
somewhat triangular; loreal about as wide as high; one preocular,
one postocular; a large narrow curving subocular; supraocular not
longer than eye; temporals two and three, however, they are irregu-
lar in second and third rows; seven supralabials, none entering eye,
four or three touching subocular, last much the largest; mental
small, touching one of the first pair of chiushields; eight (seven)
infralabials, four (or five) touch first chiushields; second and third
pairs of chinshields transverse, followed by first ventral, which is
larger than succeeding scales. Eye distinctly larger than its dis-
tance to mouth.

Contribution to Herpetology of Thailand 1127

Scales on back of head 20; 15 scalerows on neck, middle of body,
and in front of vent. Ventrals 163, subcaudals 55, paired; anal
single. Total length, 210 mm. Tail, 49 mm. Mandibular teeth, 29,
the anterior 5 times as large as the posterior.

Color: The top of the head heavily speckled with brown; a pair
of black-brown spots behind parietals; a diagonal brown line from
upper level of eye to mouth angle; on seventh labial a suggestion

Fig. 21. Pareas macularius Theobald. EHT-HMS No. 31798 9, "Thai-
land." Actual total length, 210 mm.

of a second line from lower part of eye across the suture of the
sixth and seventh labials; brown flecking on labials; a light nuchal
band from which a process runs forward behind eye, and joins the
light color of chin and throat behind jaw angle. General body
color brown with approximately 60 transverse rows of small black
dots or spots, each bordered in front by white on the same scale,
the rows irregular and often incomplete. Outer part of ventrals and

4—8050

1128 The University Science Bulletin

subcaudals flecked with brown. A deep black W-shaped mark on
neck preceded by an irregular neckband, yellowish (in life).
A photograph is shown.

Pareas carinattis Boie

(Fig. 22)
Pareas carinattis Boie, Isis, 1828, p. 1035 (type locality, Java).

The two specimens of Pareas carinattis are, EHT-HMS No. 31796
(RE 5246) Ban Po (subvillage) approx. 1780 m. elev. ("in the
same range as Phu Nam Lang ( mt. ) but farther north" ) , Na Haeo
(village), Dan Sai (district), Loei (province), Thailand, May 16,
1955, and EHT-HMS No. 31797 (RE 5610), Phu Nam Lang (mt.),
1780 m. elev., Ban Khok (subvillage), Na Phung (village), Kan Sai
(district), Loei (province), Thailand, June 10, 1954, both collected
by Robert E.Elbel.

No. 31797 5 . The body is strongly compressed. The rostral of
this species is much narrowed and visible above only as a point.
The internasals, longest mesially, are one and a half times as wide
as long. The prefrontals bend down on the sides of the head but
do not enter orbit, the suture between them being a little longer
than the internasal suture. The frontal is six-sided, the sides sub-
parallel, longer than its distance to the tip of the snout and equally
as large as the parietals. The supraocular is longer than the eye.
The nasal is as high as long, the loreal distinctly higher than wide.
The first pair of infralabials touch behind the mental. The first pair
of chinshields is not longer than the second or third transverse pairs.

The color is nearly uniform brown, finely peppered with darker
pigment. The venter is whitish with a row of irregular dark flecks
on outer edge of ventrals and a discontinuous row of flecks in the
middle of venter. A narrow line runs from eye back to the scale-
row bordering the median row, and is lost almost immediately. The
chin and neck are white, the head peppered with black flecks.
Some trace of the dorsal markings, obvious in the male, may be dis-
cerned if the specimen is submerged in clear liquid.

No. 31796 J displays some 78 narrow dark slightly diagonal trans-
verse markings, each having a width of less than a single scale, ex-
tending laterally to the ventrals. There are 7-7 supralabials, and
8-9 infralabials. Otherwise it agrees generally with the female in
squamation. There are 77 dim dark bands on body formed by
scales having a little black pigment on their edges. The median
scalerow is nearly uniformly colored in dark gray with the centers
of the scales a little lighter because of less gray pigment. The eye

Contribution to Herpetology of Thailand

1129

Fig. 22. Pareas carinatus Boie. EHT-HMS No.

31796c? , Ban Bo, Na Haeo, Dan Sai, Loei (province), Thai-
land. Actual total length, 430 mm.

Infra-
labials

Maxil-
lary
teeth

Mandib-
ular
teeth

Total
length

8-9
8-5

5-5
5-4

23
23-21

430
522

1130 The University Science Bulletin

is one-half times greater in diameter than its distance from the
mouth, and equals the length of the snout in front of the eye. The
head markings differ from the female in having a threadlike line
running below the eye to the seventh supralabial. Three series
of ventral spots or flecks are present. The median scalerow in the
posterior part of the body is somewhat enlarged. The pupil is
vertical.

There are 5 subequal maxillary, and 23 mandibular teeth, the
anterior ones two to three times the size of the posterior. The
distal half of the maxillary teeth have some distinct striations.

Data on Pareas carinatus Boie

Scale- Sub- Supra-

Number rows Ventrals caudals labials

31796 . 19,15,15,15 181 26+ 7-7

31797 . 21,15,15,15 178 71 6-7

In both there are 1-1 preoculars, 1-1 postoculars, 3 - 3 suboculars.
The temporals are 3 + 4 + 3, and 3 + 3 + 4, respectively.

Family ColubridaE

Subfamily Sibynophinae

Sibynophis coUaris triangularis subsp. nov.

(Fig. 23)

Type: KUMNH No. 33520 (RE 233), Nongko (village), Siracha
(district), Chon Buri (province), Thailand, Dr. Boonsong Lekagul,
collector, Aug. 18, 1953.

Diagnosis: Related to Sibynophis coUaris but differs in having the
black nuchal band reduced to a triangle bordered on two sides by
cream lines two scales wide, which are continuous with the cream
line on labials; a series of cream or yellow spots chiefly on the fourth
scalerow.

Description of subspecies: Rostral three-fifths times as high as
wide, narrowly visible above; internasals a little wider than long,
their posterior borders curving, laterally narrowed, touching loreal;
frontal a little wider than supraocular (4 mm. x 2.5 mm.), longer than
its distance from end of snout; parietals longer than wide, about
equal to distance from rostral; nasal scale at least partially divided
with a vertical depression following nostril, the posterior portion
higher than anterior, touching two supralabials; loreal small, longer
than high, touching two supralabials; a large preocular higher than
wide; ten supralabials, the fourth, fifth and sixth enter orbit, the
sixth and seventh touch lower postocular but the single anterior

Contribution to Herpetology of Thailand

1131

Fig. 23. Sibynophis collaris triangularis subsp. nov. Type.
KUMNH No. 33520, Nongko (village), Siracha (district), Chon
Buri (province), Thailand. Actual total length, 327 mm.

1132 The University Science Bulletin

temporal touches only the eighth; tenth and last supralabial much
the largest of the series; two poctoculars; the parietal touching only
the upper postocular which is nearly square; the temporal formula
is 1 -f-jx]; nine infralabials, the fourth largest, the first four touch-
ing the first chinshields which are larger than second pair; scales
smooth; scalerows, 17, 17, 17; ventrals, 160; anal divided; subcaudals
(incomplete), 49.

Color: Generally grayish brown with a fawn line beginning on
neck at level of the eleventh ventral, following the fourth and part
of fifth scalerows; somewhat farther back it becomes discontinuous
forming elongate or rounded spots on the upper half of the fourth
sca^row, sometimes including a part of the adjoining scale in the
fifth row; pigment surrounding these spsts becoming somewhat
blackish; on the middorsal line a row of black spots continues for a
short distance; farther back a discontinuous series of white dots be-
gins and continues posteriorly; these finally become obsolete but
they can be traced onto the tail; ventral surface whitish with a dis-
creet rounded black spot on each ventral and subcaudal; the outer
turned-up part of the ventrals pigmented like the lateral body scales.

Head variegated olive with brownish-black spots, one or two on
each head scale; an indefinite transverse band across back of frontal
and supraoculars. A second somewhat curving indefi :ite band
bordering parietals and temporals, joining the deep black triangular
nuchal spot and thus enclosing an olive area; a cream line from
rostral passes back across the labials and meets its fallow on the
median nuchal region; this bordered above on labials with black;
on the lower part of some labials black spots present; the anterior
part of infralabials light brown with darker-edged cream spotting.

Dentition: Maxillary teeth, 49-47; mandibular teeth, 53-52; pala-
tine-pterygoid series, 53-53.

Measurements in mm.: Snout to vent, 239; tail (mutilated), 88;
head width, 8.15; head length, 14.

Remarks: The snakes of this group (Sibynophinae) show a large
percentage of individuals with mutilated tails. The senior author
has recently pointed out that at least one species of the genus
Scaphiodontophis also a member of the Sibynophinae, is capable
of casting off the tail to escape. This is a habit common in lizards.
Autotomy possibly is a subfamily character and may account for
the mutilation so common in species of both genera of the subfamily.
Possibly it is a primitive character of serpents harking back to some
Saurian ancestor. The senior author has found a certain popula-

Contribution to Herpetology of Thailand 1133

tion of gartersnakes (Thamnophis) with mutilated tails but in that
particular case it was obviously brought about by disease. No such
diseased condition has been reported in any sibynophid snake.
Malcolm Smith has observed this form, and in his Serpentes,
vol. 3 of the "Fauna of British India" calls attention to it as follows:
"Specimens from Siam and Annam may have a lateral series of

Fig. 24. Acrochordus granulatus (Schneider). KUMNH No. 40064 yg., Phet
Buri, Thailand. Actual total length, 283 mm.

1134 The University Science Bulletin

yellow spots on scalerow 4 and 5, and the yellow border on the nape
may be chevron-shaped the apex pointing backwards."

Bourret has listed three forms of Sibynophis for French Indo-
China, two of which, sinensis and grahami, are regarded as sub-
species of collaris. Both Pope (1935) and M. Smith (1943),
however, have regarded these as of specific rank. In either case
they are not to be confused with the form here described.

Subfamily Acrochordinae
Acrockordus granulatus (Schneider)

Hydras granulatus Schneider, Historiae Amphibiorum naturalis et literariae,
fase. primus, 1799, pp. 243-244.

Three specimens of Acrockordus granulatus (KUMNH No. 40064-
40066 are from Phet Buri, Thailand, collected by Dr. Boonsong
Lekagul. One of the two smaller specimens (No. 40065) differs
rather considerably by having the body much compressed and
proportionally higher than the others, the black and cream bands of
nearly equal width and extending completely around the body. The
scalerows at the widest part of the body number from 130 to 140.

The largest specimen, No. 40066, has 53 black bands on body and
13 on the tail. The black bands are widened dorsally and narrowed
somewhat laterally.

No. 40064 is longer and slenderer than 40065, the largest number
of scalerows about the body being 103. There are 62 dark bands
about the body, 13 on the tail, the bands narrowing and fading on
the sides, and in some cases not crossing the venter. The body is
less compressed than in No. 40065. In all three specimens there are
eleven or twelve scales in a row between the eyes.

Subfamily Colubrinae
Elaphe radiata (Schlegel)

Coluber radiatus Schlegel, Essai sur la physionomie des serpens, vol. 2, 1837, p.
135, pi. V, figs. 5, 6 (type locality, Java); Smith, Fauna of British India,
Ceylon and Burma; Reptilia and Amphibia, vol. 3, Serpentes, Dec. 1943, pp.
146-148, text figs. 44 a, b, c, d, and 46.

The following specimens are in the collections: KUMNH No.
31445, Khorat (city and province), Thailand, Aug. 17, 1953, Robert
E. Elbel coll. 31430 (RE 969) Khon San (village), Phukhieo (dis-
trict), Chaiyaphum (province), Thailand, R. E. Elbel coll. Dec. 25,
1952. 40073 * "Thailand"; 40074, Phukhieo (district), Chaiyaphum

* This appears to be RE 950 Khon San (village), Phukhieo (district), Chaiyaphum
(province), Thailand, Dec. 23, 1952, R. E. Elbel coll. The tag is lost.

Contribution to Herpetology of Thailand 1135

(province), Thailand, 1952; 40075 yg. (bad state), Kok Kong (vil-
lage), Chanuman (district), Ubon (province) "from sand by
Mekong River," May 31, 1954. R. E. Elbel coll. EHT-HMS No.
31719 (RE 5609), Phu Nam Lang (int.), elev. 1780m., Ban Khok
(subvillage), Na Phung (village), Dan Sai (district), Loei (prov-
ince ) , Thailand, June 10, 1953. R. E. Elbel coll.
The following table shows variation in our series.

Table of data on Elophe radiata (Schlegel)

Total

Tail

Ven-

Sub-

Supra-

Scale-

Maxillary

Infra-

Number

length

length

trals

caudals

labials

rows

teeth

labials

31430 5

1457

276

231

97

9-9

19-19-17

23

10-10

31445 5

1320

252

236

98

9-9

19-19-17

22

10-11

31719 5

1840

358+

232

90+

8-9

19-19-17

22

10-10

40073 5

?

296

97

9-9

19-19-17

21

10-10

40074 $

1845

362

238

101

9-8

19-19-17

21

11-10

40075 9 yg.

9-9

19-19-17

21

9-9

This form may be readily diagnosed by the four black lines on
the anterior part of the body beginning on the neck some distance
back of the head. The posterior part of the body is nearly uniform
fawn or sometimes light olive. There are three radiating lines, one
below, two behind the eye, the upper touching a transverse black
postparietal line, from both ends of which narrow black lines run
back to connect with black spot on the side of the neck.

Ptyas mucosus (Linnaeus)

Coluber mucosus Linnaeus, Mus. Ad. Frid. I, 1754, p. 37, pi. 23; Systema Na-
turae, 10th ed., 1758, p. 226 (type locality, India).

The specimen, KUMNH No. 31444 (RE 723), is from Lam Phaya
(village), Nakhon Pathom (district and province), Thailand, col-
lected Apr. 18, 1952 by Robert E. Elbel.

The following characters obtain: supralabials, 7-8, with third and
fourth, or fourth and fifth entering eye; 9-9 infralabials; ventrals,
199; subcaudals, 117; scale formula, 23 (head), 17, 15, 17. There
are 32 tail bands, the last four or five touching a continuous dorsal
line; labial sutures black; head brown; scales with two apical pits.
Maxillary teeth 10 + 2 grooved fangs.

A second young specimen is KUMNH No. 40070 (BL 20148)
from Thailand without specific locality data from the Dr. Boonsong
Lekagul collection. Ventrals, 197; subcaudals, 120; scale formula,
25, 17 ( 18 ) 16 14. There is faint indication of keels on the posterior
part of the body. There are light black-edged bands on latter half
of the body the black continued on venter as black edges on
ventrals. The anterior part of the body is nearly uniform olive
(olive brown in preservative)

1136 The University Science Bulletin

Ptyas korros (Schlegel)

Coluber korros Schlegel, Essai sur la physionomie des serpens, vol. 2, 1837, p.
139 (type locality, Java).

Four specimens of this species are in the collection, KUMNH No.
31434 (RE 951) from Khon San (village), Phukhieo (district),
Chaiyaphum (province), Dec. 24, 1952; No. 31446 (RE 998), 6 km.
N Chaiyaphum (city), Chaiyaphum (district and province), Jan.
3, 1953, and KUMNH No. 40080 (RE 3644), Ban Hua Bua (sub-
village), Um Mao (village), Yangtalat (district), Kalasin (prov-
ince), all collected by R. E. Elbel, May 6, 1954; KUMNH No. 40069
(BL 20040) "Siam", Dr. Boonsong Lekagul coll.

Table of data for Ptyas korros

Sex or

Snout-vent

Tail

Number

age

Ventrals

Subcaudals

Scalerows

length

length

31434

9

177

141

15, 13, 11

935

533

31446

$

170

153

15, 15, 11

1036

?

40069

yg

171

127+

15, 16, 11

329

143

40080

$

163

?

15, 15, 11

1100

P

Lycodon capucinus Boie

Lycodon capucinus Boie, Isis, 1827, p. 551 (based on Russell, Indian Serpents,
vol. 2, pi. 37, (type locality Trivandrum, India).

One typical specimen, BL coll. No. ? Rat Buri, Thailand, was
examined. We do not regard this as a subspecies of Lycodon
aulicus ( Linnaeus ) since each of these forms occupies a large part
of the range of the other.

Oligodon cyclurus (Cantor)

(Fig. 25)

Coronella cyclurus Cantor, Proc. Zool. Soc. London, 1839, p. 50 (type locality,
?); M. Smith, Jour. Nat. Hist. Soc. Siam, vol. 4, 1920, p. 96.

Simotes fasciolatus Giinther, Reptiles of British India, 1864, p. 218, pi. XX, fig.
B ( type locality "Petchabun," SE Siam ) .

Simotes smithi Werner, Sitz. Akad. Wiss. Wien, vol. 143, p. 58 (type locality,
Siam).

The several varieties of this species were regarded by Malcolm
Smith (1943) merely as color varieties that intergrade with one
another. Certain of these variations are represented in the few
specimens in the collections as follows :

KUMNH No. 31429 (RE 909), $ from Ban Lat (subvillage),
Ban Kaeng (village), Phukhieo (district), Chaiyaphum (province),
Thailand, Dec. 16, 1952, Robert E. Elbel, coll. This specimen has
the following characters: supralabials, 8-8, the fourth and fifth
border orbit; infralabials, 9-9; scale formula, 26 (head), 21, 21, 17;
ventrals, 175, somewhat angulate laterally; subcaudals, 43; the anal

Contribution to Herpetology of Thailand 1137

single; two pre- and two postoculars; temporals, 2 + 2; 10-9 maxil-
lary teeth, three enlarged.

The triangular head markings are rather narrow, not drawn out
anteriorly. The body markings are as follows: the thirteen black
circular spots are separated from each other by three dark blotches
or marks,. the median one most distinct; two circular spots on tail.

Fig. 25. Oligodon cyclurus (Cantor). KUMNH No. 40067 "Thailand."

Actual total length, 510 mm.

A row of small dark spots on the fourth and fifth scalerows; immac-
ulate below except for some pigment on the outer edges of the
ventrals. This color variety differs somewhat from those listed by
Smith (1943, pp. 202-206).

EHT No. 31700 <? (RE 4215) from Ban Na Muang ( subvillage ) ,
Na Haeo (village), Dan Sai (district), Loei (province), Thailand,

1138 The University Science Bulletin

elev. approx. 1780 m., Oct. 23, 1954, collected by Robert E. Elbel,
gives the following scale data: supralabials, 8-8; infralabials, 9-9;
scalerows, 24 (head), 21, 21, 17, 15; scales smooth without pits;
temporals, 2 -4- 2; frontal six-sided; preoculars, 1-2; postoculars, 2-2;
loreal present; ventrals, 185; subcaudals, 56; anal single.

EHT-HMS No. 31701 (RE 3909), from Phu Kho (mt.) 522 m.
elev., Kan Luang (village), Na Kae (district), Nakhon Phanom
(province), Thailand, July 24, 1954, collected by Dr. Boonsong
Lekagul and Robert E. Elbel, yields the following data: supralabials,
8-8, fourth and fifth enter orbit; infralabials, 9-9, four or five touch
first chinshields; scale formula, 26 (head), 21, 21, 17, 17; ventrals,
161; subcaudals, 43; anal single; eleven maxillary teeth, last two
large; head and neck pattern complete (see Smith, loc. cit., p. 203).
Between the ten large brown spots on body there are three, or more
rarely four, indefinite transverse lines similar to the figure given by
Smith, fig. D, p. 205, but more intense. Venter with some pigment
on outer edge of ventrals.

KUMNH No. 40067, "Thailand" and EHT-HMS No. 31702 (RE
3817) from Phu Phan (mt.) 550 m. elev., Sakon Nakhon (district
and province), June 28, 1954, R. E. Elbel and Dr. Boonsong Lekagul
collectors, are also in the collection.

Table of data from Oligodon cijclurus

Scalerows Sub- Supra- Infra- Pre- Post- Total Tail

Number at middle Ventrals caudals labials labials oculars oculars length length

31700 <$ 21 185 56 8-8 9-9 1-2 2-2 812 136

31701$ 21 161 43 8-8 9-9 2-2 2-2 579 86

31702i 21 166 54+ 8-8 9-10 2-2 2-2 109

31429 9 21 175 43 8-8 9-9 2-2 2-2 685 90

40067 5 21 166 54 8-8 9-9 2-2 2-2 510 111

Oligodon cinereus Giinther

(Fig. 26)

Oligodon cinereus Giinther, Reptiles of British India, 1864, p. 215 (type lo-
cality, Cambodia); Smith, Fauna British India, Ceylon and Burma. . . .
Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 215-217.

Simotes inornatus Boulenger, Jour. Nat. Hist. Soc. Siam, vol. 1, 1914, p. 69
(type locality, Sriracha, SE Siam).

Oligodon violaceous Smith, Jour. Nat. Hist. Soc. Siam, vol. 4, 1920, p. 96.

One specimen in the collection, EHT-HMS No. 31699 5 (RE
4532), resembles the body pattern of Malcolm Smith's var. II, loc.
cit., 1943, p. 216. It is from Ban Muang Khai (subvillage), Tha Li
(village), Tha Li (district), Loei (province), Thailand, Jan. 20,
1955, Robert E. Elbel, coll. The following characters obtain: 8-7
supralabials, the fourth and fifth or the third and fourth entering
eye (the third and fourth fused on one side); temporals, 2 -f 2; two

Contribution to Herpetology of Thailand

1139

Fig. 26. Oligodon cinereus Gunther. EHT-HMS No.
31699 9, Ban Muang Khai, Tha Li, Tha Li (district), Loei
(province), Thailand. Actual total length, 600 mm.

1140 The University Science Bulletin

preoculars, the lower much smaller than upper, which is quad-
rangular, more than twice as wide as high; scale formula, 19 (head),
17, 17, 17, 15; ventrals, 190; subcaudals, 39; anal single; 12 maxillary
teeth.

The general color is dark gray with a series of diagonally placed,
dotted dark lines often discontinuous along the edges of diagonal
scalerows or on skin between; alternate scalerows have a dashed
cream line along their edges often extending on the skin between.
Tail with two or three indefinite small spots but the characteristic
markings of body are absent. The ventrals are generally white with
their outer ends dark gray or ash, and with large black spots on
alternate ventrals. The subcaudals are white except on the ends
and there are no spots evident. A well-defined interocular band is
present.

Oligodon quadrilineatus (Jan)

(Fig. 27)

Simotes quadrilineatus Jan, Nouv. Arch. Mus. Paris, vol. 2, pt. 7; Iconographie

general des ofidiens, tome 1, livr. 12, 1865, pi. 4, fig. 3 (type locality, Siam);

Smith, The Fauna of British India, Ceylon and Burma; Beptilia and

Amphibia, vol. 3, Serpentes, Dee. 1953, p. 210.
Simotes taeniatus Boulenger, Catalogue or the snakes in the British Museum

(Natural History), vol. 2, pp. 227-228 (part.).

The name Oligodon quadrilineatus was resurrected by Malcolm
Smith in 1943 for this species. It had long been considered a
synonym of O. taeniatus. The two specimens in our collection help
confirm the distinctness of quadrilineatus. These are KUMNH Nos.
40068, from Thailand (probably from near Bangkok), and No.
31437, from Ban Lat ( subvillage ) , Ban Kaeng (village), Phukhieo
(district), Chaiyaphum (province), Thailand, Jan. 17, 1952, Robert
E. Elbel, coll.

The general characters of these two specimens, Nos. 31437 $ and
40068 $ respectively, are as follows: supralabials, 7-8, 8-8; infra-
labials, 9-9, 9-9; labials enter orbit, 4th & 5th, 4th & 5th; preoculars
and postoculars, 1-2, 1-2; scale formula, 22 (head), 19, 19, 17, 15;
22, 19, 19, 17, 15; suboculars, 0, 0; temporals, 1 + 1 + 2, 1 + 1+2;
ventrals, 161, 151; subcaudals, 40, 44; anal, 1, 1. Total length in
mm.: 354, 411; tail, 56, 68.

The specimens also agree in color. There is a pair of dark lines
dorsally, separated by a narrow lighter line less than the width of a
scalerow. The two dark lines meet on the nape of the neck, then
widen, terminating on the parietal. On the head a pair of dark
diagonal stripes run from the parietal across the angle of the mouth
and there are two or three other transverse stripes across the snout.

Contribution to Herpetology of Thailand

1141

Some scattered small dark marks are present on the head scales, in-
cluding a diagonal line below the eye and a shorter one following it
behind. The dorsal ground color is grayish tan. A narrow lateral
brown line covers parts of the third and fourth scalerows and
terminates at the level of the vent. The median pair of brown lines

Fig. 27. Oligodon quadrilineatus Jan. KUMNH No. 31437, Ban Lat,
Ban Kaeng, Phukhieo, Chaiyaphum (province), Thailand. Actual total length,
354 mm.

1142 The University Science Bulletin

continues to the tip of the tail. A very indefinite brownish line
borders the outer edge of the subcaudals.

The venter is coral red in life (whitish in preservative) with
triangular or quadrangular brown marks irregularly placed on ven-
trals. Usually a scale without marks alternates with one bearing
one or two brown spots. The chin and subcaudal regions are im-
maculate.

Ahaetulla ahaetalla ahaetulla (Linnaeus)

Coluber ahaetulla Linnaeus, Systema Naturae, ed. 10, 1758, p. 228 (part).
Ahaetulla ahaetulla ahaetulla Smith, Fauna of British India, Ceylon and Burma;
Reptilia and Amphibia, vol. 3, Serpentes, 1943, pp. 239-244, fig. 85.

There has been very considerable disagreement as to the proper
name for this species.* It was long known under the designation
Dendrophis pictus and so appears in the major part of the literature
dealing with the species.

One specimen (KUMNH No. 40076 [RE 1691]) from Pak-
nampho (city), Nakhon Sawan (district and province), Thailand,
was collected by the junior author.

The following characters obtain: maxillary teeth, 22; the four
posterior largest; one preocular touching frontal; two postoculars
the upper elongate; supralabials, 9-9, the 4th, 5th and 6th entering
orbit; temporals, 2 + 1 + 2 (1 + 1 + 2); one loreal more than
twice as long as high; scales following parietals and temporals some-
what enlarged; scale formula, 19 (head), 15, 15, 9; median row
enlarged but definitely somewhat less wide than outer scalerow.
The ventrals are 172/2; subcaudals 130, anal divided. Scales with
a single apical pit, the surface of the epidermis finely striate, not
discernible when epidermis is shed.

There is a strong black line from behind eye onto neck; a cream
line on the two outer scalerows with a well-defined dark line
below it covering part of outer scalerow and outer edge of the
ventrals. Type locality unknown.

Subfamily Boiginae

Boiga multimaculata (Boie)

Dipsas multimaculata Boie, Isis, 1827, p. 549 (type locality, Java).

Our specimen, KUMNH No. 31439 (RE 890), is from Non Khun
(village), Phukhieo (district), Chaiyaphum (province), Thailand;
collected Dec. 12, 1952, by Robert E. Elbel.

* Besides Malcolm Smith, loc. cit., see also discussion by James Oliver, Bull. Amen
Mus. Nat. Hist., vol. 92, art. 4, 1948, pp. 167-190, 267-272; J. M. Savage, Bull. Chicago
Acad. Sci., vol. 9, 1952, pp. 203-216. 8

Contribution to Herpetology of Thailand 1143

This specimen has the following scale counts: supralabials, 8-8,
the third, fourth and fifth entering the eye; infralabials, 11-11; pre-
ocular, 1; postoculars, 2; temporals, 2 -f- 2; scale formula, 27 (head),
19, 19, 13; ventrals, 223; anal single; subcaudals, 99. Body slender,
compressed. Total length, 565 mm.; tail, 113 mm.

Two dark stripes begin on the posterior temporal areas and run
forward diagonally and meet on the anterior part of the frontal,
continuing on to the middle of the prefrontals; a dark stripe from
angle of mouth runs forward to eye, and then continues in front of
eye to nasal scale. Back and tail with a series of dark brown spots,
often irregular in outline, running from neck to tail, separated
mesially by a narrow line of the gray of the general coloration.
Chin unspotted; anteriorly, dark marks present on outer part of
ventrals; more posteriorly the flecks become more numerous extend-
ing over whole ventral region.

Boiga cyanea (Dumeril, Bibron and Dumeril)

(Fig. 28)

Triglyphodon cyaneum Dumeril, Bibron and Dumeril, Erpetologie generale,
vol. 7, pt. 2, 1894, pp. 1079-1080 (type locality unknown).

Two specimens are at hand, EHT-HMS No. 31693 (RE 4016)
from Ban Na Muang (subvillage), elev. approx. 1780 m. "in same
range as Phu Nam Lang (mt. ) but farther north," Na Haeo
(village), Dan Sai (district), Loei (province), Thailand; Sept. 29,
1954, Robert E. Elbel coll., and EHT-HMS No. 31826 (RE 4689)
from Phu Lorn Lo (mt), elev. 2100 m., Kok Sathon (village), Dan
Sai (district), Loei (province), Feb. 18, 1955, R. E. Elbel, coll.

No. 31693 5 has the following characteristics: supralabials, 8-8;
the third, fourth, and fifth entering eye; infralabials, 11-11, four (or
five) touching first pair of chinshields; one preocular, reaching upper
surface of head but not touching frontal; eye very large, its diameter
equal to its distance from nearest point of rostral; two postoculars;
temporals, 3 + 3 ( median divided ) + 4 on both sides; scale formula,
29 head, 21, 21, 15, 15, and the median row somewhat enlarged;
ventrals, 248; subcaudals, 136; anal single. Body compressed. The
subcaudals have an indistinct lateral keel; no gland visible through
the posterior labials. Total length, 1558 mm.; tail, 311 mm.

The color was probably uniform green in life. In preservative it
is bluish to bluish gray, the edges of the scales darker. The chin-
shields are cream.

No. 31826 is a young specimen and we are not wholly convinced
that it is correctly referred to this species. The following characters
obtain: supralabials, 8-8; the third, fourth, and fifth entering orbit;

1144 The University Science Bulletin

11-11 infralabials, four touching first chinshields; one preocular
reaching upper surface of head, but not touching frontal; scale
formula, head 28, 22, 21, 15, the median row somewhat enlarged;
ventrals, 250; subcaudals, 124, with an elongate terminal shield,

Fig. 28. Boiga cyanca (Dumeril, Bibron and Dumeril). EHT-HMS
No. 31826, Phu Lorn Lo (mt.), 2100 m. elev., Kok Sathon, Dan Sai, Loei
(province), Thailand. Actual total length, 700 mm.

rounded at tip, grooved on both sides; anal single; temporals,

2 + 3 + 3 (5
tail, 161 mm.

2 + 3 + 3 (2+^+3); loreal as high as long; total length, 720;

Contribution to Herpetology of Thailand 1145

The head is uniform smoky black, probably green in life; three
posterior supralabials semitransparent showing a cream-colored
gland lying below them. Body with dark markings as shown in the
figure.

The species has been reported from Nakon Lampang, Dong Rek
Mts., and from certain islands in the Gulf of Siam.

Malcolm Smith and others have pointed out that the young are
not green but are variously colored and marked. The squamation
is practically identical to that of the adult specimen reported above.
One specimen examined in the Boonsong Lekagul collection (No.
222) is from Khao Khansong (village), Sriracha (district), Chon
Buri (province), Thailand, Aug. 1953, Dr. Boonsong Lekagul, coll.

Boiga cynodon Boie
Boiga cynodon Boie, Isis, 1827, p. 549 (type locality, Sumatra).

Our single specimen is EHT-HMS No. 31698 (RE 4099), from
Ban Na Muang (subvillage, elevation 1780 m.), Na Haeo (village),
Dan Sai (district), Loei (province), Thailand; October 6, 1954,
Robert E. Elbel, coll. The following characters obtain: supra-
labials, 8-8, the third, fourth and fifth entering orbit; infralabials,
12-12, the first four (or five) touching the first pair of chinshields;
second chinshields larger than first pair; frontal broader than long;
eye equals its distance from nostril; temporals, 2 + 3, 3 -f- 3.

Scale formula, (head) 26, 23, 23, 15, 15; ventrals, 261, anal single,
subcaudals, 118. Maxillary teeth, 11 + 2 grooved fangs.

Above clouded and indefinitely blotched with brown, fawn, and
black. A series of light (fawn) blotches low on the sides surrounded
by black, situated chiefly on the outer part of ventrals. Remainder
of venter gray-brown.

Dryophiops rubescens (Gray)

Dipsas rubescens Gray, Illustrations of Indian Zoology, vol. 2, 1834, pi. 84, fig.
2 (type locality, ? Malay Peninsula).

One specimen seen is BL Coll. No. 20099, collected by Dr. Boon-
song Lekagul, from "Siam." This widespread, usually arboreal
species is relatively rare in collections. The senior author has taken
it on Penon de Coron, western Philippines, a limestone island cov-
ered sparsely with small shrubs and plants, but lacking trees owing
to the almost complete absence of soil on the rock.

1146 The University Science Bulletin

Psammodynastes pulverulentus Boie
Psammodijnastes pulverulenta Boie, Isis, 1827, p. 547 (type locality, "Java").

This variable and widespread form is represented by three speci-
mens: EHT-HMS No. 31793 (RE 3817), from Phu Phan (mt.)
550m. elevation, Sakon Nakhon (district), Sakon Nakhon (prov-
ince), Thailand; collected by Dr. Boonsong Lekagul and Robert
E. Elbel June 28, 1954. No. 31794 (RE 4689), a beheaded speci-
men, Phu Lorn Lo (mt.) 2100 m. elev., Kok Sathon (village), Dan
Sai (district), Loei (province), Thailand; Feb. 18, 1955, Robert E.
Elbel, coll. No. 31795 (RE 5002), Phu Nam Lang (mt.) 1780m.
elev., Ban Khok (subvillage), Na Phung (village), Dan Sai (dis-
trict), Loei (province), Thailand, June 2, 1955, R. E. Elbel, coll.

No. 31794 § had recently eaten a young E.umeces quadrilineatus.
The specimen is gravid, containing seven eggs. There are 49 sub-
caudals and 17 scalerows.

The following scale data are taken from No. 31795 $ : supra-
labials, 8-8; infralabials, 8-8; five pairs of chinshields, the second
and third about equal, three infralabials touching the first pair;
one loreal, two pre- and two postoculars; supraoculars wider than
frontal; ventrals, 163; subcaudals, 53; anal single; scalerows, head,
25, neck, 17, body, 17, in front of tail, 17, all without pits.

No. 31793 5 has the following scale counts: supralabials and
infralabials, 8-8, the third, fourth, and fifth supralabials border orbit;
three infralabials touch the first chinshields; one preocular; two post-
oculars; the temporals, 2 + 3 + 3. The second and third pairs of
chinshields are longer than the first; scalerows, 24, 17, 17, 15; ven-
trals, 149; subcaudals, 52. The maxillary teeth vary greatly in size,
the last two being grooved.

The species occurs also in Indonesia, and the senior author has
collected it in the mountains of central eastern Mindanao, P. I.

Chrysopelea ornata (Shaw)

Coluber ornatus Shaw, General Zoology, 1802, vol. 3, p. 477 (type locality,
East Indian Islands, based on Seba, Thesaurus, vol. 1, pi. 94, fig. 7; vol. 2
pi. 7, fig. land pi. 61, fig. 2).

Chrysopelea ornata Brongersma, Zool. Meded, Leiden, vol. 20, 1938, p. 241;
M. Smith, The fauna of British India, Ceylon and Burma, including the
whole of the Indo-Chinese sub-region; Beptilia and Amphibia, vol. 3, Ser-
pentes, Dec. 1943, pp. 251-254, text figs. 86, 87.

Two forms of the arboreal C. ornata are recognized by Malcolm
Smith, one occurring in Thailand, one in Ceylon and India.

The former is characterized as having "the black cross-bars much
less conspicuous and may be entirely absent; the mesial streak on

Contribution to Herpetology of Thailand 1147

each scale may then give the appearance of black longitudinal
lines." The head markings consist of five or six golden yellow trans-
verse bars following scale sutures between rostral, internasals, pre-
frontals, frontal and following parietal while one crosses the middle
of the frontal. There are usualy several golden dots between the
lines. The lines may be apparently broken when crossing longi-
tudinal sutures. In the latter form the black and yellow bars al-
ternate.

Five specimens are at hand in the collection as follows: KUMNH
Nos. 31431 (RE 1600), Paknampho (city), Nakhon Sawan (district
and province), Thailand, Jan. 18, 1953, and 31432, Banpong (city
and district), Rat Buri (province), June 16, 1952, both collected by
Robert E. Elbel.

EHT-HMS Nos. 31817 (RE 3756), Phu Phan (mt.), 550 m., 104°
05'; 16° 55', Sakon Nakhon (district and province); 31818 (RE
3846), Phu Kho (mt), circa 522 m. elev. (104° 22'; 16° 49'), Kan
Luang (village), Na Kae (district), Nakhon Phanom (province).
Collected June 20 and July 16, 1954, respectively by Dr. Boonsong
Lekagul and Robert E. Elbel.

In the form occurring in Thailand the row of scales bordering
the parietals and temporals behind are definitely enlarged; the ven-
trals, anals, and subcaudals are strongly keeled; in the region on
sides and above anus the scales are usually more or less keeled.
In all there are two postoculars, the temporals, 2 -f- 2 + 2, and the
scalerows are 24 (25) head, 17, 17, 13. The infralabials are ten
or eleven with five scales usually bordering the first chinshields.

Chrysopelea paradisi occurs in the peninsular area of Thailand,
and also in Indonesia and the Philippines.

Scale data on Chrysopelea ornata Shaw

Preocular

reaches

Number

Sex

Preocular

frontal

Ventrals

Subcaudals

Anals

31817

9

1-1

no

218

107 +

2 2 %

31818

9

1-1

no

222

117

2 2 (?)

31819

$

1-1

yes

223

138

2 2

31431

S

1-1

no

224

141 +

2 2

31432

$

1-1

no

222

140

2 % 2

Dryophis prasinus Boie

Dryophis prasinus Boie, Isis, 1827, p. 545 (type locality, Java); Malcolm Smith,
Fauna of British India, Ceylon and Burma, including the whole of the Indo-
Chinese sub-region. Beptilia and Amphibia, vol. 3, Serpentes, Dec. 1943, pp.
375-386, text fig. 120.

Malcolm Smith's (loc. cit.) treatment of the genus calls attention
to the fact that Dryophis xanthozonia Boie should bear the name

1148 The University Science Bulletin

Dryophis mycterizans (Linnaeus) while the species long known
in literature under the latter name should bear the name Dryophis
pulverulentus Dumeril and Bibron and Dumeril. This is mentioned
here since mycterizans occurs in Thailand also. It may be separated
from the present species by its undivided anal plate, fewer ventrals *
and subcaudals.

A single specimen of Dryophis prasinus, KUMNH No. 40077 (RE
3474) is in the collection. It was captured on Phu Lorn Lo (mt. )
2100 m. elev., Kok Sathon (village), Dan Sai (district), Loei (prov-
ince), Mar. 26, 1954, by Robert E. Elbel.

The following characters obtain: no proboscis, the nasals not
in contact; sharp canthus; nasal single, elongate, the nostril in pos-
terior part; two or three loreals separate prefrontals from labials;
one preocular, longer than high, with a sharp canthus, broadly
touching the frontal; supralabials, 9-9, the fourth, fifth and sixth
enter orbit; two postoculars; frontal three times as wide anteriorly
as posteriorly; temporals 2 + 2 + 3 + 3, the lower anterior very
small; anterior chinshields broader and shorter than the very narrow
posterior ones; infralabials, 9-9, the first four touch first chinshields.

Scale formula: 21 head, 15, 15, 11, the scales in the region of vent
with keels or tubercles, the middle line of the scales with series of
pits, four to five on some scales. Ventrals ? (body broken); sub-
caudals, 196; anal divided.

Dryophis nasutus Lacepede

Dryophis nasutus Lacepede, Histoire naturelle des serpens, vol. 2, 1789, Dis-
course and Table Methodique, p. 100. Also, pp. 277-279, pi. 4, fig. 2 (type
locality, Ceylon, Guinea, Carolina); Smith, The fauna of British India, Cey-
lon and Burma, including the whole Indo-Chinese subregion. Reptilia and
Amphibia, vol. Ill, Serpentes, Dec. 1943, pp. 376-378 (synonymy and litera-
ture), fig. 119 (head, three views).

Of the type localities listed by M. le Compte de Lacepede for this
species, one only, Ceylon, has the species. In consequence the
locality Ceylon is hereby designated as the revised type locality.

One specimen, KUMNH No. 31438 (RE 783) from Paknampho
(city), Nakhon Sawan (district and province), Thailand, Jan. 30,
1953, Robert E. Elbel, collector, is in the collection. There are 202
ventrals, the last (preceding divided anal) also divided. The
divided anal has a small intercalated median scute which may be
an abnormality. The subcaudals are 163. The scalerows about the
back part of the head are 20, about neck 15, about middle of body

* 186-195 ventrals, 132-156 subcaudals fide Malcolm Smith while prasinus has 194-235
ventrals, 165-187 subcaudals fide Malcolm Smith.

Contribution to Herpetology of Thailand 1149

15, while in front of aims the number reduces to 11. There are two
pre- and two postoculars.

The tooth series on the maxillary consists of three spaced teeth
anteriorly followed by a fanglike tooth, which, after a diastema, is
followed by four small teeth, and two enlarged grooved fangs.

The nasal appendage is formed entirely from the rostral. The
loreal is absent.

Total length, 662 mm.; tail, 227 mm.

Subfamily Natricinae
Pseudoxenodon macrops macrops (Blyth)

(Fig. 29)

Tropidonotus macrops Blyth, Asiat. Soc. Bengal, vol. 23, 1854, p. 296 (type
locality, Darjeeling, Bengal, India).

Two specimens are in the collection: EHT-HMS No. 31703
(RE 4521) is from Ban Muang Khai ( subvillage ) , Tha Li (village),
Tha Li (district), Loei (province), Thailand, Jan. 19, 1955, and
EHT-HMS No. 31704 (RE 5533) juv., Phu Nam Lang (mt), (elev.
1780 m.), Ban Khok (subvillage), Na Phung (village), Dan Sai
(district), Loei (province), Thailand, June 4, 1955; both collected
by Robert E. Elbel.

There is a series of white ( ? ) transverse bands across the mid-
line extending throughout the body to the tip of the tail, the edges
of each band in front and behind bordered by large dim dark spots
formed of scales with black edges and grayish centers. Laterally
the light bands terminate at black spots which form a row on the
third to fifth scalerows. The space on the dorsal area between each
two dark spots may be light (whitish) but is never as distinct as
the primary white marks. There are quadrangular dark spots on
the ventrals.

The following scale characters obtain: scalerows, 19, 15, 15;
anterior temporals, 2; supralabials, 8; infralabials, 8; ventrals, 171;
subcaudals, 63; anal, 2.

The young specimen has 174 ventrals and 67 subcaudals. The
supralabials are 8-8, with dark sutures. The infralabials are 9-9.
On the back there are 50 white transverse spots. The black chev-
ron on the neck is preceded by a lighter line. Total length, No.
31793: 882 mm.; tail, 160; No. 31704: 263 mm.; tail, 47.5.

The larger contains 10 ovarian eggs.

1150

The University Science Bulletin

Genus Natrix

The genus Natrix is a heterogeneous assemblage and efforts to
break up this unnatural group have not been successful. One group
of species, characterized by very long posterior ungrooved fangs,
is thrown in with species having little or no enlargement of the

Fig. 29. Pseudoxenodon macrops macrops (Blyth). EHT-HMS
No. 31704, Phu Nam Lang (mt.), 1780 m., Ban Khok, Na Phung, Dan
Sai, Loei (province), Thailand. Actual total length, 263 mm. (Note
that the head is proportionally large in this young specimen.)

posterior teeth. We are of the opinion that the entire matter is in
strong need of re-examination.

In this paper we propose to recognize the large-fanged species
under the generic name Rhabdophis. The following species of

Contribution to Herpetology of Thailand 1151

Natrix (scnsu lata) have been reported in Thailand: deschauenseei,
groundwateri, irias, modesta, piscator piscator, trianguligera, per-
carinata, subminiata subminiata, subminiata helleri, stolata, nigro-
cincta, chrysarga. The last five forms are typical Rhabdophis, R.
subminiata being the type of the genus. Certain other forms re-
ported from Thailand may be misidentifications or are synonyms.
Such a case is Bourret's Rhabdophis tigrina lateralis which Malcolm
Smith places as a synonym of piscator.

Natrix deschaaenseei, described by the senior author from Chiang
Mai, Thailand, has also been placed as a synonym of Natrix modesta
by Dr. Malcolm Smith. It, however, seems to differ from that Indian
species in having 36 maxillary teeth (28 to 32 recorded for mo-
desta); nasal divided; two preoculars, two postoculars (usually
three ) ; temporals, 1 + 2; only two ( fifth and sixth labials ) entering
orbit instead of three; ventrals, 159, subcaudals, 137 — total, 296,
higher than reported for modesta. The body is very slender with
a proportionally longer tail. The color pattern differs likewise in
details (see figures of the type and compare with Boulenger, vol. 1,
pi. 9, fig. 3). Dr. Smith states that he has not seen a specimen of
deschauenseei. It is possible that collections from the region of
Chiang Mai, the type locality, will provide data suggesting the
distinctness of the species or confirming Dr. Smith's opinion as to
synonymy. No material is available in this collection. Bourret
has placed the species in the genus Macropophis Boulenger.

Natrix piscator flavipunctata (Hallowell)

Amphiesma flavipunctatum Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1860,
p. 503 ( type locality, Kwantung province, China ) .

Two specimens are referred to the subspecies Natrix piscator
flavipunctata: KUMNH No. 31441 J (BE 965), Khon San (vil-
lage), Phukhieo (district), Chaiyaphum (province), Thailand, Dec.
24, 1952; No. 31440 J (BE 967), same locality, Dec. 25, 1952.
Bobert E. Elbel, collector.

The following characters obtain in the two specimens, 31441 and
31440 respectively: snout pointed; internasals pointed anteriorly;
body with a tendency to form a slight median ridge; supralabials,
9-9, 9-9; labials entering orbit, 4 and 5, 4 and 5; 4, 4 and 5; pre-
oculars, 1-1, 1-1; postoculars and suboculars, 3-4, 3-3; infralabials,
10-10, 10-10, five touching the anterior chinshields which are shorter
than second pair; scalerows, head, 25, 19, 19, 17; 24, 19, 19, 17;
two outer scalerows smooth (3 anteriorly), others weakly keeled;

1152 The University Science Bulletin

anterior temporals, 2, 2; ventrals 134 $ , 127 $ ; subcaudals, 59 -j-,
85.

The head is black above in No. 31441 the color extending onto the
neck; a diagonal black line along suture of sixth and seventh labials;
another black line from behind eye runs down across the mouth
angle then curves back up onto side of neck bordering the dark
head mark; a distinct black spot on each side of the neck. Nearly
uniformly lavender-brown with the merest trace of dark markings
and numerous dashlike cream or yellow indistinct marks chiefly
confined to interstitial skin or edges of scales; ventrals and sub-
caudals are dark edged. In No. 31440 some lateral dark marks
running up on sides diagonally, ventrals and subcaudals cream or
whitish, their edges barred with black.

The maxillary teeth in 31441, 24, the last four enlarged and
strongly compressed; teeth in No. 31440, 26-24, the last four com-
pressed and enlarged, the last a little larger than the others. There
is no diastema between the larger and the smaller teeth.

While we follow Malcolm Smith in treating this form as a sub-
specific form of Natrix piscator there seems room for doubt that this
is its true relationship.

Rhabdophis subminiata helleri (Schmidt)

Natrix helleri Schmidt, Amer. Mus. Novitates, no. 157, 1925, p. 3 (type locality,
Tengyueh, 5500 ft. altitude, province of Yunnan, China ) .

The following series of specimens are referred to this form:
EHT-HMS No. 31765 (RE 5504), Phu Nam Lang (mt), 1780 m.,
Ban Khok (subvillage), Na Phung (village), Dan Sai (district),
Loei (province), Thailand, June 2, 1955; EHT-HMS No. 31766 (RE
4323), 31812 (RE 4311), 31813 (RE 4305), 31814 (RE 4256),
31815 (RE 4256), 31816 (RE 4310)— all from Ban Nong Wai (sub-
village), 1780 m., Na Phung (village), Dan Sai (district), Loei
(province), Thailand, Nov. 15-20, 1954, all collected by Robert E.
Elbel. KUMNH No. 40072 (RE 3189), Ban Muang Khai (sub-
village), Tha Li (village), Tha Li (district), Loei (province), Thai-
land, about 600 m. elev., Dec. 4, 1953; collected by Robert E. Elbel,
and Dr. Boonsong Lekagul.

The following table presents data from the above series of speci-
mens.

Contribution to Herpetology of Thailand 1153

Data on Rhabdophis subminiata helleri

Maxillary

Supra-

Labials

Infra-

teeth

Sub-

Number

Sex

labials

enter orbit

labials

and fangs

Ventrals

caudals

31766

s

8-8

4, 5

10-10

J25+2
123+2

155

89+

31811

$

8-8

3, 4, 5

10-10

22+2

153

84

31812

9

8-9

/3, 4, 5

14, 5, 6

10-10

22+2

151

84

31813

9

9-8

)4, 5, 6
13, 4, 5

10-10

22+2

152

78+

31814

$

8-8

4, 5

9-10

22+2

158

81

31815

9

8-8

4, 5

10-10

22+2

155

83

31816

9

8-9

(3, 4, 5
14, 5, 6

10-10

23+2

163

?

Young specimens have a large discrete black spot covering the
back of the head and extending for ten to twelve scale-lengths be-
hind parietals; it is bordered by a lighter line about one and one-
half scale-lengths wide. There follow several rows of indefinite
dark spots more or less connected and forming a reticulum. The
enclosed scales have red interstitial skin, the color sometimes cover-
ing more than half of the scale. Farther back the color is uniform
gray olive (somewhat blackish in preservative) with usually two
series of single or double paired yellow dots on the sides; sometimes
these pairs will be connected by small dashlike marks across the
back. When the skin is distended more yellow marks are dis-
cernible on interstitial skin or on lower edges of scale. A few
specimens show traces of black marks usually, appearing as black
edges of certain adjoining scales.

The maxillary teeth are small anteriorly gradually attaining a
slightly greater size, then, following a short diastema, are two greatly
enlarged fangs directed nearly backwards. Ventral coloration is
light, probably yellowish (or reddish) in life, the outer edges of
the ventrals gray, often with small discrete or diffuse dots near their
ends; supralabials light with a strongly defined diagonal bar below
back part of eye. Dorsal head markings are dim or obsolete in
old specimens.

Rhabdophis stolaia (Linnaeus)

(Fig. 30)

Natrix stolata Linnaeus, Systema Naturae, 10th ed., 1758, p. 219 (type locality,
Asia).

A single specimen, EHT-HMS No. 31761 s (RE 3952), is from
Ban Na Phua (subvillage), Kan Luang (village), Na Kae (district),
Nakhon Phanom (province), Thailand, collected July 28, 1954,
by Robert E. Elbel and Dr. Boonsong Lekagul. The markings
are typical of the eastern (Chinese) specimens. The following

1154

The University Science Bulletin

characters obtain: supralabials, 8-8; infralabials, 10-10; temporals,
1 + 1, or 1 -r- 2; scale formula, 19, 19, 17; ventrals, 144; subcaudals,
50; preocular, 1; postoculars, 3; maxillary teeth, 21 -f 2 (enlarged).
Snout-vent length, 356 mm.; tail, 92 mm.

There is a light chevron-shaped band across the neck. From
this two stripes begin, covering the fifth scalerow and halves of the
adjoining rows. These stripes, grayish at first, become white farther
back and extend to the tip of the tail.

Fig. 30. Rhahdophis stolata Linnaeus. EHT-HMS No. 31761, Ban Na Phua,
Na Kae, Nakhon Phanom (province), Thailand. Actual total length, 448 mm.

Contribution to Herpetology of Thailand 1155

Rhabdophis nigrocincta (Blyth)

Tropidonotus nigrocinctus Blyth, Jour. Asiat. Soc. Bengal, vol. 24, 1856, p. 717
(type locality, Pegu, Burma).

There are two specimens in the collections: EHT-HMS No.
31760 <y (RE 5517) from Phu Nam Lang (mt), 1780 m., Ban Khok
(snbvillage), Na Phung (village), Dan Sai (district), Loei (prov-
ince), Thailand, June 3, 1955, Robert E. Elbel, coll.; and KUMNH
No. 40071 $ (RE 3541) from Phu Lorn Lo (mt), 2100 m. elev.,
Kok Sathon (village), Dan Sai (district), Loei (province), Thai-
land, Mar. 30, 1954, Dr. Boonsong Lekagul and Robert E. Elbel,
collectors.

The following characters obtain in No. 31760: maxillary teeth,
29-30, followed by two suddenly enlarged fangs lacking grooves,
several times larger than nearest tooth; ventrals, 167; subcaudals,
99; anal divided; scale formula, 27 head, 19, 19, 17; scales except
outer row keeled; two anterior temporals; one preocular, three post-
oculars; supralabials, 9-9, the fourth to sixth enter orbit; internasals
truncate anteriorly; a distinct series of 8 pairs of nuchal glands;
on each side of median line of neck a short indistinct groove. Total
length in mm.: 615; tail, 179.

A narrow band (six to eight rows wide) following the parietals
is lighter than ground color, suggesting a juvenile light band; this
is followed by a deep-black chevron pointing backward; general
ground color dark gray with a series of 46 narrow transverse black
bars on the body, sometimes broken mesially and alternating; tail
banded and flecked with black. Venter with anterior part ivory,
the median third gray, the posterior part with dark blackish-gray
or brown clouding or blotching; underside of tail dark with white
flecks; an immaculate white or cream spot on labials between a
diagonal postocular stripe and a short subocular stripe.

No. 40071 is a larger specimen but it is injured anteriorly so that
an accurate measurement cannot be made. It agrees in general
with the above specimen. There are two anterior temporals, one
preocular and four postoculars; 9-9 supralabials, 10-11 infralabials,
five or six bordering the anterior chinshields which are shorter than
posterior; latter separated by small scales; ventrals, approximately,
166, subcaudals, 88; anal divided. Scale formula, 28 head, 19, 19, 17.
There are 31 + 2 maxillary teeth.

The stomach contains a specimen of a frog (Megophrys) partially
digested.

1156 The University Science Bulletin

Rhabdophis chrysarga (Schlegel)

Tropidonotus chrysargus Boie, Isis, 1827, p. 534 (Java), nomen nudum.
Schlegel, Phys. Serp., vol. 2, 1837, p. 312, pi. 12, figs. 6-7.

A specimen (BL No. 20090 from "Thailand") was examined.
This species is widespread, extending through southeastern Asia,
Indonesia and into the Philippines.

Parahelicops boonsongi sp. nov.

(Fig. 31)

Holotype: EHT-HMS 31707 (RE 5500), Phu Nam Lang (int.),
1780m., Ban Khok (subvillage), Na Phung (village), Dan Sai
(district), Loei (province), Thailand, June 2, 1955; Robert E.
Elbel, collector.

Diagnosis: A natricine snake, with 30 maxillary teeth, the last
three suddenly much enlarged but not grooved; internasals nar-
rowed; a single prefrontal; three (or two) postoculars; two sub-
oculars; one preocular; nasal partially divided; anal divided; with
a divided scale also preceding it; all dorsal scalerows finely striate
and heavily keeled, outer ones less so except outermost which is
smooth anteriorly and faintly keeled posteriorly; posterior edges of
scales notched, bearing a fine transparent fringe. Uniform gray-
olive on dorsal parts (in preservative), growing gradually lighter
laterally, the outer scalerows almost cream-white anteriorly; venter
cream-white or yellowish; subcaudal region gray.

Description of type: Rostral once and a half as wide as high,
very narrowly visible above; internasals elongate, narrowed an-
teriorly, rounded posteriorly, their length slightly greater than
length of the prefrontal; latter twice as wide as long; frontal five-
sided, slightly wider anteriorly than elsewhere, a little shorter
than supraoculars, equal to its distance from rostral, a third shorter
than parietals; nasal elongate; a suture from nostril to second supra-
labial; loreal as high as long; a large preocular not reaching frontal;
three postoculars, two suboculars, the anterior reaching forward
beyond middle of eye; temporals, 2 -f- 3 -f- 4; nine supralabials, the
first three touching nasal, third only touches loreal, fourth and fifth
enter orbit, seventh largest; mental wider than deep, as wide as
rostral; infralabials ten, five touching first chinshields which are dis-
tinctly shorter than second pair; latter separated, narrowly so an-
teriorly.

Scalerows across back of head 25, at neck 19, middle of body 19,
in front of vent 17. Ventrals, 141; anals, 2; subcaudals, 33 -f- (tail

Contribution to Herpetology of Thailand 1157

broken and lost). Eye rather large, equal to its distance from nos-
tril; pupil round.

Color: Nearly uniform gray-olive growing a little lighter laterally,
becoming nearly cream-white on outer two or three scalerows an-
teriorly on side of neck; posteriorly two outer scalerows and to
some extent the outer edge of ventrals with some grayish pigment;
supralabials and suboculars cream; an indefinite fumate spot back

Fig 31. Parahelicops Boonsongi sp. nov. EHT-HMS No. 31707, Phu Nam
Lang (mt.), 1780 in., Ban Khok, Na Phung, Dan Sai, Loei (province), Thai-
land. Actual total length 621 mm.

1158 The University Science Bulletin

of jaw angle; tail incomplete; venter generally cream-white; sub-
caudal region with a median gray line and many scales with an in-
definite pigmented area (on portion of tail remaining).

When the specimen is held in certain lights or immersed in clear
liquid it is possible to discern dimly certain indefinite darker areas
on the anterior half of the body.

Measurements in mm.: Snout to vent, 520; tail, 101 + ; total
length 621 + ; length of head, 26; width of head, 18; width of neck,
13.

Remarks: The genus Paralielicops was proposed by Bourret for
a species having a single prefrontal, the scales lacking apical pits,
hypapophyses developed throughout the vertebral column, 25 max-
illary teeth, the last two enlarged but not grooved. The present
species differs from the described type of the genus Paralielicops
annamensis Bourret in having a totally different pattern of colora-
tion, more maxillary teeth (30 as compared with 25); the nasal
partially instead of wholly divided; the scalerows 19, 19, 17 instead
of 17, 17, 15; the anal double (preceded by a paired scale), instead
of a single anal; eye larger, and the ventrals 141 (compared with
169 in annamensis). The character of the temporals (abnormal in
annamensis) are 2 + 3-f-4 (as compared with 1 + 1 + 2 on the
normal ? right side while on the left side there is one long fused
scale followed by three ) . There are three instead of one ( or two )
suboculars.

The species is named for Dr. Boonsong Lekagul who has so
generously provided the University of Kansas with Thailand am-
phibians and reptiles.

Pope (Rept. China, 1936, p. 125, and M. Smith, vol. 3, Fauna
Indian Reptilia and Amph., 1943, p. 330), have placed Paralielicops
in the synonymy of Opisthotropis together with seven or eight other
generic names, thus bringing together a rather heterogeneous group
of species. There are smooth and keeled forms; forms with maxil-
lary teeth varying from 20 to 40, equal or "subequal" or having
two or three much enlarged fanglike posterior teeth; groups with
19, 17, or 15 scalerows at midbody, groups with or without sub-
oculars, etc. The one common character is the presence of a single
prefrontal. Whether Giinther's atra the type of the genus Opis-
thotropis is African as stated by Boulenger or is actually South
Asian cannot be determined by us. We are recognizing Bourret's
Paralielicops and placing this species in that genus rather than in
Opisthotropis. It is significant that Opisthotropis spenceri M. Smith
has been described from Thialand ("Muang Ngow, N. Siam") a

Contribution to Herpetology of Thailand 1159

species having 17 smooth scalerows throughout, lacking the sub-
oculars, and lacking the enlarged posterior maxillary teeth.

It is surprising that certain forms of both Porahelicops and
Opisthotropis are nearly unicolored above and yellowish or whitish
below. One suspects that this may be an adaptation to a stream
habitat. However, but little is known of the ecology of these snakes.

Subfamily Homalopsinae
Enhydris plumbca (Boie)
Homalopsis plumbea Boie, 1827, p. 560 (type locality, Java).

Two specimens are at hand: EHT-HMS No. 31705 yg. (RE 3818)
from Phu Phan (mt. ), 550 m. elev., Sakon Nakhon (district and
province), Thailand, June 28, 1954, Dr. Boonsong Lekagul and
Robert E. Elbel, colls., and EHT-HMS No. 31706 (RE 541128),
Kam (village), Khon Kaen (district and province), Nov. 28, 1954,
collected by Robert E. Elbel.

These two have the following characteristics respectively: supra-
labials, 8-8, 8-8; infralabials, 10-10, 10-10; scale formula, 23, 19, 19,
15; 23, 19, 19, 13; ventrals, 127, 125; subcaudals, 39, 37; anals, 2, 2.
Total length in mm., 220, 455; tail 31, 52.

No. 31706 contains 14 enlarged ovarian eggs.

Enhydris enhydris (Schneider)

Hydriu enhydris Schneider, Historiae amphibiorum naturalis et literariae,
fasc. primus, 1799, pp. 245-246 (type locality, India orientalis, fide Russell).

One specimen, EHT-HMS No. 31759 (RE 541022) is at hand
from near Kam (village), Khon Kaen (district and province),
Thailand, collected Oct. 22, 1954, by Robert E. Elbel. It appears
to be typical of the species. The following characters obtain:
all scales smooth without pits; scale formula: 34, 21, 21, 21, 19;
ventrals, 150; subcaudals, 47+ (tail broken); anal divided. An-
terior chinshields shorter than second pair.

Dorsal scalerows nearly uniform brownish gray, the three outer
scales at least partly without pigment. Ventrals outlined in brown;
a median brown subcaudal line. The stomach contained eight small
fish and parts of others. "Collected on road."

Homalopsis buccata (Linnaeus)

(Fig. 32)

Coluber buccatuj Linnaeus, Mus. Ad. Frid., 1754, p. 29, pi. 19, fig. 3; Systema

Naturae, 10th Ed., 1758, p. 217 (type locality, India).
Homalopsis buccata Smith, Jour. Nat. Hist. Soc. Siam, vol. 1, 1914, p. 101;

idem ibid., vol. 1, no. 4, p. 162 (Bangnara, Patani); ibid, idem, vol. 1, p. 213

5_8050

1160

The University Science Bulletin

Fig. 32. Homalopsis buccatus (Linnaeus). EHT-HMS No. 31697,
Phu Phan ( mt. ), 550 m., Sakon Nakhon (district and province), Thailand.
Actual total length, 772 mm.

Contribution to Herpetology of Thailand 1161

(listed); the fauna of British India, Ceylon and Burma, including the whole
of the Indo-Chinese sub-region; Reptilia and Amphibia, vol. 3, Serpentes, pp.
390-392, fig. 123 (after Boulenger, 1890) (rivers, canals, ponds of Burma
south of lat. 17°, Siam, Cambodia, Cochin-China, Malay Peninsula; Indo-
nesia, usually found not far above tidal limits); Boulenger, A vertebrate
fauna of the Malay Peninsula; Reptilia and Batrachia, 1912, pp. 162-163,
fig. 50; Bourret, Les serpents de 1'Indochine, tome 2, 1936, pp. 293-294,
fig. 117 (listing of literature).

This widely distributed serpent is represented by four specimens:
EHT-HMS Nos. 31694-31697 (RE 3818), all collected on Phu Phan
(int.), 550 m. elev., 104° 05'; 16° 55', Sakon Nakhon (district
and province), Thailand, June 28, 1954, by Robert E. Elbel and
Dr. Roonsong Lekagul. These vary in age and to some extent
in color pattern. In No. 31695, the smallest specimen, (489 mm.),
the body is brownish black on the back and sides, with 21 narrow
(two or three scales wide) creamy-white transverse bands on body
and 13 on tail. The bands on the back part of the body in some
cases alternate with very narrow light lines which tend to divide the
intervening dark blotches. An irregular cream line beginning at
the mouth-angle runs back along the third and fourth scalerows
for some distance, then becomes broken and some of the individual
spots resulting are placed transversely on the venter. The venter
is generally brownish black with many small rounded cream spots
and anteriorly a few transverse bars, none completely crossing the
venter.

A cream line from the angle of the mouth runs forward to below
the eye to the nasal scale, while another begins at the same place,
runs along the lower lip a short distance, then crosses the chin.
From the middle part of the chin two cream lines run back onto
the throat where they disappear. A pair of large rounded spots
are present on the occiput.

In general this same detailed pattern is repeated in two other
specimens, a male and female both larger than the described speci-
men. In a still larger specimen, No. 31696, the dorsal color is nearly
brown and the transverse bands have become almost obsolete. The
venter is brownish black showing small cream spots. The head
markings are constant although dim in the largest specimen.

Measurements and scale data on Homalopsis buccata

Total

Chin

Sub-

Number

length

Tail

scales

Ventrals

Anals

caudals

Scalerows

31695 6

489

114

11

157

2

84

48, 39, 27

31694 $

573

118

10

154

2 + 2

67

54, 39, 26

31696 £

782

185

8

160

2 + i

82

53, 38, 27

31697 3

772

188

8

160

2

80

49, 39, 27

1162 The University Science Bulletin

Malcolm Smith ( 1943 ) reports a Siamese specimen from "Eastern
Siam, B. M. Coll." that has the whole lower part grey, thickly spotted
with black. The largest known specimen measures 1310 mm.,
the tail being 285 mm. in length.

Family Elapidae
Bungarus candidus (Linnaeus)

Coluber candidus Linnaeus, Systema Naturae, 10th ed., 1758, p. 223 (type
locality, India).

A specimen, KUMNH No. 31435 (BE 1063), was collected by the
junior author Aug. 26, 1952, at Khorat ( city and province ) , Thailand.

The following characters obtain: supralabials, 7-7, the third and
fourth entering orbit; infralabials, 7-7; scale formula: 19, 15, 15, 15,
the median row much enlarged; ventrals, 221; anal single; sub-
caudals, 42 -j- ( tail incomplete ) . Total length, 1442; tail, 140 mm.

There are 23 light dorsal bars widening on the side, separated by
23 saddlelike black blotches, narrowing on the sides, reaching down
and covering outer part of ventrals. The scales of the light areas
have flecks or bars of dark color, the median row with two dark
marks on each scale. A wide cream stripe follows the upper lip
covering all or the greater part of all the labials. The top of the
head is uniformly dark.

This is, we believe, the fifth specimen reported from Thailand.

Naja naja kaouthia Lesson

Naja kaouthia Lesson, in Ferrusae, Bull. Sci. Nat., vol. 25, 1831, p. 122 (type

locality, Bengal).
Naja damensis Laurenti, Specimen medicum, exhibens synopsin reptilium

emendata . . ., 1768, p. 91 (tvpe locality, Thailand), based on Seba,

vol. 2, pi. 89, figs. 1-2.

There are three specimens of this cobra in the collections ob-
tained by the junior author. These are KUMNH Nos. 40081-40082
(BE 2624) Khao Khat (mt.), approx. 500 m. elev., Paknampho
(city), Nakhon Sawan (district and province), Thailand, June 4,
1953.

The following characters obtain in 31433: scale formula 27
(neck), 21, 14-15; ventrals, 161, subcaudals 34+; anal single.
Total length 987 mm.; tail, 115 mm. (tip incomplete). This is from
the same locality as the other two.

Contribution to Herpetology of Thailand 1163

Family Hydrophiidae
Enhydrina schistosa (Daudin)

Hydrophis schistosus Daudin, Histoire naturelle generate et particuliere des

reptiles, vol. 7 (an XI), 1803, p. 386 (type locality, "Coromandei" ) .

A single young specimen of E. schistosa (KUMNH No. 40060)
from Phet Buri, Thailand, was collected by Dr. Boonsong Lekagul.

The following scale characters obtain: rostral higher than wide,
angulate above, the median lower part fitting into an indistinct
notch in front part of lower jaw; nasals in contact, narrowed and
angulate anteriorly, partly divided; one preocular, one postocular;
prefrontals widened anteriorly, narrowed posteriorly, not in contact
with labials; frontal small, narrower than a supraocular, angulate
anteriorly and posteriorly, shorter than its distance from anterior
end of nasals; eight supralabials, the four anterior enlarged, the
fourth one entering orbit, four posterior ones small.

Temporals 1 -f- 1 + 2 + 4, the second touches the postocular;
mental a very slender elongate scale nearly hidden in a groove; first
supralabials narrow, much elongate, not in contact behind mental;
two pairs of chinshields, the anterior largest in contact with three
supralabials. Eight infralabials, the three or four posterior ones
small.

Scale formula: 46 (head), 42, 47, 58, 35. Grayish white or cream
with 34 brown spots on back and 4 on anterior part of tail (latter
half of tail black). The specimen measures, total length, 325 mm.;
tail, 38 mm.

Thalassophina viperina (Schmidt)

Thalassophis viperina Schmidt, Abh. Nat. Ver. Hamburg, Band 2, 1852, p. 79,
pi. 3, (type locality, "lava").

The one specimen examined is BL No. 20063, Thailand (probably
from the Gulf near Bangkok), collected by Dr. Boonsong Lekagul.

Lapemis hardwicki Gray

(Fig. 33)

Lapemis hardwicki Gray, Illustrations of Indian Zoology, vol. 2, 1834, pi. 87, fig.
2 (type locality, Penang); Smith, Monograph of the sea snakes (Hydro-
phiidae), London, Nov. 27, 1926, pp. 108-112, fig. 32, (Gulf of Siam).

A typical specimen of this widespread species (KUMNH No.
40063) from Phet Buri, Thailand, collected by Dr. Boonsong Leka-
gul, May, 1954, is in the collection. The total length is 646 mm.;
tail, 66 mm.

1164

The University Science Bulletin

Fig. 33. Lapemis hardtvicki Gray. KUMNH No. 40063, Phet Buri (province),
Thailand. Actual total length, 646 mm.

Hydrophis mamUlarls Daudin

(Fig. 34)

Hydrophis mamillaris Daudin, Historic naturelle generale et particnliere des
reptiles, vol. 7, an XI=1803, p. 340 (based on Russel, vol. 1, p. 49, pi. 44
from Vizagapatam, India).

KUMNH No. 33517. We are associating with this species a
specimen of sea-serpent from Bangphn (village), Bangkok, Thai-
land, collected by Dr. Boonsong Lekagul. The following characters

Contribution to Herpetology of Thailand

1165

obtain: ten maxillary teeth following grooved tangs; 1 (2) pre-
ocnlars, 1 postocular; supralabials, 7, second touches prefrontal on
one side only; the fourth only enters orbit, last three small; infra-
labials, 10-10, three only touch the first chinshields, the second and
third on one side separated from the lip by two or three intercalated

Fig. 34. Hydrophis mamillaris Daudin. KUMNH No. 33517, Bangphu, Bang-
kok, Thailand. Actual total length, 655 mm.

small scales; posterior infralabials small. A single large anterior
temporal followed by a still larger scale, and the latter touched
by another enlarged temporal bordering the parietal. This last
scale separated from its fellow by four small scales.

Scale formula, 36 (head), 33, 39, 32; ventrals. approximately * 325,

* Occasional small body scales are intercalated between ventrals making the count
somewhat uncertain.

1166 The University Science Bulletin

not double width of adjoining scales. Two preanals much en-
larged, separated from each other by two and one half pairs of
scales, the last pair extending somewhat behind them. A somewhat
enlarged pair of postanal scales. Head small, scarcely wider than
neck.

Head and neck about one-third times greatest diameter of body.
Head black with a yellow streak on each side behind eye. Body
blackish with 55 dark bands on body, and seven on flattened tail,
wider dorsally, narrowing much on sides; except for the first eight,
the white bands encircle the body, or occasionally fail to meet on
venter by a very narrow margin. The dorsal and lateral scales
of the light bands may have considerable pigment forming indefinite
dark spots on the scales; most of the dorsal series of scales anteriorly
have longer or shorter keels which more posteriorly become short-
ened into tubercles, the scales of the median dorsal row sometimes
have two such tubercles, one following the other; ventrals usually
with two lateral tubercles often indistinct.

The presence of a form of mamillaris in the Gulf of Siam is un-
expected. There are some differences and with sufficient material
a subspecific form might be recognized. The most significant
differences are, only a single labial entering eye (the fourth); 33
scalerows around the neck; only a single postocular; only three
scales touch the first small chinshields, the second pair distinguish-
able but not large. The head has a yellow streak on each side from
nasal to eye and from eye to the first transverse white spot. There
are two small scales bordering the lip between the anterior labials
excluding the third labial on one side, and three small scales on the
other side excluding both third and fourth from lip.

The species shows some similarity to H. fasciatus atriceps M.
Smith and holbrookci Giinther but the maxillary teeth are twice
as numerous. It would appear that the maxillary teeth are less
variable than other characters.

From H. caerulescens it differs in a smaller number of teeth,
smaller head and anterior body, a longer frontal, only one instead
of two supralabials touching orbit, and a smaller number of scale-
rows on body.

The specimen is gravid, containing three very small embryos
attached to large egg masses.

Contribution to Herpetology of Thailand

1167

Hydrophis caerulescens (Shaw)

(Fig. 35)

Hydrus caerulescens Shaw, General Zoology, vol. 3, 1802, p. 561 (type locality,
Indian Ocean).

Two young specimens, KUMNH Nos. 40061, 40062 from Phet
Buri (province), Thailand, were collected by Dr. Boonsong Leka-

Fig. 35. Hydrophis caerulescens Shaw. KUMNH No. 40062, Phet Buri,
Thailand. Actual total length, 537 mm.

1168 The University Science Bulletin

gul. They have been assigned to H. caerulescens chiefly on the
basis of the large number of teeth behind the poison fangs. In
No. 40062 there are 14-16; in 40061 there are 14 teeth. The follow-
ing characters obtain:

No. 40061: 1 preocnlar, 1 postocular, eight supralabials, the third
and fourth enter orbit, the second touches prefrontal; the fifth
labial is large and is possibly fused to the anterior temporal; last
three labials small; infralabials seven, the first four small touch-
ing the anterior chinshields, last three small. Scale formula, 34
(neck) — 42-35. Anal scales rather large, separated by three paired
scales, the last pair almost completely behind them. Ventrals, 293;
subcaudals, 410. Black bands on body and tail, 63, laterally taper-
ing very slightly; those on tail connected below. Head black with
some yellow lateral marks.

No. 40062 differs in some points. There are two postoculars on
one side, the temporal is not fused to the labial, the anterior tem-
poral contacts the eye at one point. There is a small scale on lip
border between the third and fourth infralabials. There are 61
bands on body and tail; head without light marks; the light bands
tend to be greatly narrowed, and interrupted ventrally; dorsally
the white bands contain much dark pigment. Ventrals, 407; sub-
caudals, 36.

Family Crotalidae

Agkistrodon rhodostoma (Boie)
Trigonocephalus rhodostoma Boie, Isis, 1827, p. 561 (type locality, Java).

One specimen, EHT-HMS No. 31718 (BE 3817) $\ (bad state)
is in the collection from Phu Phan (mt.), 550 m. elev. (104° 05';
16° 55'), Sakon Nakhon (district and province), Thailand; June 28,
1954; Bobert E. Elbel and Dr. Boonsong Lekagul, colls.

The following characters obtain: supralabials, 7-7; infralabials,
11-11, four touch the first chinshields; temporals, 2-4-1-4-2; two
labials touch the single subocular; ventrals, 150; subcaudals, 53;
scale formula, 19, 21, 21, 17.

Key to Thai species of Trimeresurus

1. First labial fused partially or completely with the nasal 2

First labial completely separated from the nasal scale 3

2. Upper head scales small subequal, tubercular or granular; supra-

oculars narrow (sometimes broken) separated by from twelve to
fifteen scales; internasals two to three times size of adjoining scales,
usually separated by a single scale; supralabials, 11-13, separated
from the elongate subocular by from two to three rows of scales;
scale formula, 25, (25, 27), (19, 21), strongly keeled. Usually

Contribution to Herpetology of Thailand 1169

uniform purplish-brown or olive with spots; tail spotted with

brown Trimeresurus p. purpureomaculatus

Upper head scales small, subimbricate; supraoculars narrow, not
broken or divided, separated by from eight to twelve scales; inter-
nasals two to four times size of adjoining scales, usually in contact;
10-12 supralabials separated from large subocular by one or two
rows of scales; scale formula, (21, 23), (21, 19), (15, 17). Green
above, yellow or white below, males with a ventrolateral cream
stripe, females with a row of light flecks on oviter scalerow,

Trimeresurus albolabris

3. An elongate subocular present 4

Supraoculars divided; no white or cream line along outer scalerow;

supraoculars divided transversely, separated by eight scales; inter-
nasals separated by a single scale; ten supralabials, a single series
of scales between labials and the elongate subocular; temporal
scales small with obtuse keels; scale formula, 19, 19, 15; ventrals,
152; subcaudals, 42, tail prehensile. Brownish gray with a series of
brown spots and smaller ones on sides . Trimeresurus kanhuriensis

4. Supraoculars not divided transversely, usually their length equals

twice their width, separated by from ten to thirteen subimbricate
scales; internasals separated; temporals more or less strongly
keeled; scale formula, 21 (23), 21, 15 (17). Green above, usually
whitish below; a light stripe bordered below by orange or chocolate
along flank and base of tail usually on first scalerow; a light post-
ocular stripe. Hemipenes without spines . . Trimeresurus popeorum
Similar to preceding but hemipenes spinose . Trimeresurus stejnegeri

Trimeresurus albolabris Gray

Trimeresurus albolabris Gray, Zool. Misc., p. 48, 1842 (type locality, "China");
Smith, Fauna of British India, Ceylon and Burma, including the whole of
the Indo-Chinese Sub-Region; Reptilia and Amphibia. Vol. Ill, Serpentes,
Dec. 1943, pp. 523-525.

Six specimens are referred to this species despite considerable
variation. They are EHT-HMS No. 31768 (RE 4259) from Ban
Nong Wai (subvillage), approx. 1780 m. elev., Na Phung (village),
Dan Sai (district), Loei (province), Thailand, Nov. 15, 1954; EHT-
HMS No. 31769 (RE 541118), Ban Sang Kho (subvillage), Khok
Phu (village), Sakon Nakhon (district), Sakon Nakhon (province),
Thailand, Nov. 18, 1954; EHT-HMS No. 31770 (RE 4028), Ban
Na Muang (subvillage), approx. 1780 m. elev., Na Haeo (village),
Dan Sai (district), Loei (province), Thailand, Sept. 30, 1954;
EHT-HMS No. 31801 (RE 5501), Phu Nam Lang (mt.) 1780 m.,
Ban Khok (subvillage), Na Phung (village), Dan Sai (district),
Loei (province), Thailand, June 2, 1955; KUMNH No. 40083 (RE
3217), Ban Muang Khai (subvillage), Tha Li (village), Tha Li
(district), Loei (province), Thailand, Dec. 7, 1953. All specimens

1170 The University Science Bulletin

collected by Robert E. Elbel except last, collected by Dr. Boonsong
Lekagul and Robert E. Elbel.

A young specimen, EHT-HMS No. 31769 (RE 541118) shows the
male markings clearly. It is nearly uniform greenish above
(greenish blue with a large series of brown marks on each side
of the body in preservative), the venter white growing olive on
outer edges of ventrals. A cream-white line originating on pre-
oculars passes below the eye and over temporals covering most
of two whole rows of temporals, then, crossing mouth angle, it
continues along the outer scalerow covering more than half of each
scale; on the tail it passes onto the outer part of the much widened
subcaudals. The prehensile tail is more or less spotted with brown.
Chin and to some extent throat, white; rostral and two first supra-
labials partly white.

The following scale characters obtain: supraoculars large, their
length three times their width, separated from each other by nine
scalerows, the scales flat, smooth, imbricate, the temporals not or
but very indistinctly keeled. Internasals large, in contact, three to
four times size of the adjoining canthal; supralabials, 11-12, first
fused to nasal; long curved subocular; two postoculars; third supra-
labial touches the subocular; fourth separated from it by one scale-
row, succeeding ones by two rows; infralabials, 12-14; two pairs of
chinshields, the second small, in contact, separated from ventrals
by four pairs of scales; scale formula: 46 (head), 21, 21, 15; scales
not discernibly keeled; ventrals, 163; anal single, subcaudals, 73 -f-
1; total length, 251 mm.; tail, 45 mm.

The coloration of this specimen is quite different from that re-
ported by other workers for the species. The specimen is very
young and this coloration possibly is lost in the adult. The absence
of keels on the scales is possibly owing to the youth of the specimen.

No. 31768 5 . The following characters obtain: internasals in
contact, three to four times as large as the two canthals; supraocu-
lars four times as long as wide, separated by ten or eleven smooth,
subimbricate scalerows; three postoculars, one long subocular; the
fourth and fifth scales separated from it by one scale; rostral one
and two-thirds times as wide as high; two pairs of chinshields,
followed by four paired scales; infralabials, 12-11; ventrals, 157;
anal single; subcaudals, 75 -f- 1; scale formula, 44 head, 23, 21, 15,
scales completely smooth; no ventrolateral line and no line behind
or under eye; whitish on chin; a row of light flecks on outer scale-
row. Tail uniformly colored.

Contribution to Herpetology of Thailand 1171

No. 31770 5 . Internasals one and one-third times as long as
wide, separated by two scales, one following other; two canthals
less than .half size of internasals; loreal present, reaching or just
failing to reach canthal edge; 11-10 supralabials, the first more than
half fused with nasal, the third touching subocular, fourth separated
by one scale, fifth by two; two postoculars; one long subocular
widened anteriorly; supraoculars four times as long as wide, sepa-
rated by nine or ten small subimbricate scales; posterior temporals
definitely ridged or keeled; one pair of chinshiclds followed by six
paired scales, none separated by small scales; scale formula, 44
(head), 21 (22), 21, 15; ventrals, 167; subcaudals, 58 + 1. A row
of cream flecks on outer scalerow. Tail brown spotted.

No. 40083. Internasals rectangular, about two and one-half times
longer than wide, separated by a single scale; supraoculars about
two and one-half times as long as wide, separated by 11-12 scales,
not keeled but somewhat thickened, sometimes moundlike; supra-
labials, 12-12, first completely fused to nasal, third touches the sub-
ocular; other labials separated from it by two rows of scales; infra-
labials, 14-14; two pairs of chinshiclds, followed by four paired
scales none separated by small scales; two postoculars; scale formula,
48 (head), 21, 21, 15. The ten to fifteen dorsal scalerows well
keeled; ventrals, 163; subcaudals, 52 -f- 1.

Color (in preservative): uniform bluish-black, grayish on tail;
venter bluish or bluish black, the posterior edges of the ventrals
fight; a row of light flecks on outer scalerow; tip of lower jaw black;
light areas on chin and beginning of throat. Tail grayish, not
brown spotted.

When skin is distended there appears to be transverse rows of
light flecks, the light color being on skin between scales; these sepa-
rated by similar dark areas. This occurs more or less distinctly
in all the specimens (in 31769 the whole "light" area is brown).

Trimeresurus popeorum Smith

(Fig. 36)

Trimeresurus popeorum Smith, Jour. Bombay Nat. Hist. Soc, vol. 39, p. 730
(original spelling popeiorum, a clerical error, was corrected by the author)
(restricted type locality, Khasi Hills, Assam).

Two specimens are referred to this species. They are EHT-HMS
Nos. 31799 (RE 5503) and 31800 (RE 5620) from Phu Nam Lang
(mt.), 1780 m. Ran Khok (subvillage), Na Phung (village), Dan
Sai (district), Loei (province), Thailand, June 2, and June 10, 1955,
respectively. Robert E. Elbel, collector.

1172

The University Science Bulletin

The second specimen, 31800, has the following characters: rostral
angular, slightly wider than high, narrower than mental; two loreals
superimposed; eye relatively large, the length of snout a little less
than twice its greatest diameter; internasals large, double the size
of two enlarged contiguous canthals, but three to four times larger
than other scales adjoining, separated from each other by one

Fig. 36. Trimcrestirus popeorum M. Smith. EHT-HMS No. 31800, Phu
Nam Lang (mt. ), 1780 m. elev., Ban Khok, Na Phung, Dan Sai, Loei (prov-
ince), Thailand. Actual total length, 412 mm.

Contribution to Herpetology of Thailand 1173

elongate scale; supralabials, 10-9, the first completely free from
nasal, the second high forming anterior border of the pit, the third
labial largest separated from the elongate subocular by one scale,
the fourth labial separated from same by one (or two) scales;
infralabials, 12-12, three touch chinshields; only one differentiated
pair of chinshields, followed by five pairs of scales preceding the
first ventral, the two anterior pairs separated each by a pair of
small scales; supraoculars three to four times as long as wide sepa-
rated from each other by 12-13 irregular rows of smooth subimbri-
cate scales; temporals bordering labials largest. Scale formula,
45, head, 21, 21, 15; the dorsal 15 series vaguely keeled; posterior
temporals with short posterior elevations or keels; subocular narrow,
elongate, curving; two postoculars. Ventrals, 167, anal single, sub-
caudals, 64+1. Tail prehensile, slender, and practically the same
diameter for much of its length. Total length, 412 mm.; tail 69
mm.

Dorsally nearly uniform green (dark blackish in preservative);
a light ventrolateral line, cream above, orange along its lower edge,
covering all of the outer scalerow, the lower two fifths of second
row, and the extreme outer edge of ventrals; on the tail only part
of the outer row is involved, the stripe covering also outer part of
subcaudals. Tail flecked and spotted with brown and black, except
terminal part which is uniform light brown. Beginning back of
eye a cream line bordered by orange above runs diagonally back,
crosses mouth-angle and is continuous with the ventrolateral stripe.

The second specimen is a young female. It agrees in general
with the characters given. The following characters obtain: supra-
oculars three times as long as wide separated by 10-12 scales; length
of snout not double eye diameter; internasals separated by two
scales (abnormal); two loreals small, superimposed; supralabials,
10-10, first separated from nasal, third separated from eye by one
scalerow, the fourth and succeeding labials by two rows; infralabials,
12-12; one pair of chinshields, followed by 5 pairs of scales, the
first two separated by small scales. Scale formula, 48, 21, 21, 15;
ventrals, 165; anal single; subcaudals, 57 + 1.

Head and body uniform bluish black except for a narrow bluish
white line confined almost entirely to middle of the outer scalerow;
on tail the outer edges of the subcaudals are somewhat involved.
Total length, 285 mm.; tail, 44 mm.

Remarks: Malcolm Smith founded the name, T. popeorum on
T. gramineus (Shaw) Pope and Pope. No type locality was men-
tioned by Smith.

1174 The University Science Bulletin

The material treated by Pope and Pope under T. gramineus had
a range from "Upper Burma southward into the Malay Archi-
pelago". The exact localities were Darjeeling, Bengal; Mergui,
Tenasserim; Prov. Wellesley, Malay Peninsula; Pinang, Malay Pen-
insula; "Sungei Kumbang, Korinchi", Sumatra; Lao Mts. Cochin-
China; Saiap, Kina Balu, Borneo.

A detailed description of one specimen, (British Museum Regis-
ter No. 72.4.17.137, Khasi Hills, Assam, Jerdon collector), is given
by Pope and Pope. We propose to designate this specimen as the
holotype of T. popeorum, and to restrict the type locality to Khasi
Hills, Assam.

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1176 The University Science Bulletin

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Leyden, Mus., vol. 13, 1891, pp. 255-256.

LoNNBERG, E.

1916. Zoological results of the Swedish zoological expeditions to Siam,
1911-1912 and 1914. II Lizards. Stockholm Vet. Akad. Handl,
vol. 55, no. 4, 1916, pp. 1-12.
Nieden, Fritz

1926. Amphibia, Anura II, Engystomatidae. Das Tierreich, Lief. 49,
pp. i-xvi; 1-110.
Noble, G. K.

1926. An analysis of the remarkable cases of distribution among the Am-
phibia, with descriptions of new genera. Amer. Mus. Novitates,
New York, no. 212, 1926, pp. 1-24, figs. 1-9.
Parker, H. W. A.

1934. A monograph of the frogs of the family Microhylidae. London,
1934, pp. i-viii, 1-208; figs. 1-65.
Rai, K.

1936. Die Beeinflussung des Blutzuckers von Kaninchen durch die Gifte
Siamesischer Schlangenarten. Jour. Med. Ass. Formosa, vol. 35, 10,
1936, pp. 2195-2209.
Robinson, H. C.

1905. Further additions to the batrachian fauna of the Malay Peninsula
with a list of the species at present known to occur therein. Jour.
Fed. Malay States Mus., vol. 1, pp. 19-24.

Contribution to Herpetology of Thailand 1177

SCHAUENSEE, RODOLPHE MEYER DE

1928. Random notes on snakes in Siam. Bull. Antiven. Inst., Glenolden
Pa., vol. 2, 1928, no. 3, pp. 76-77.
Smith, Malcolm

1914. The snakes of Bangkok. Jour. Nat. Hist. Soc. Siam, vol. 1, 1914,
pp. 5-18, 3 pis.

1914a. The distribution of Ancistrodon rhodostoma the Malayan viper in

Siam. Jour. Nat. Hist. Soc. Siam, vol. 1, 1914, pp. 57-58.
1914b. Large-banded krait. Jour. Nat. Hist. Soc. Siam, vol. 1, no. 1, Feb.

14, 1914, pp. 58-59.
1914c. Occurrence of the krait (Bungarus candidus) and the small-spotted

coral snake Callophis macidiceps in Siam. A new color variety of

the latter. Jour. Nat. Hist. Soc. Siam, vol. 1, Aug. 1914, pp. 123-

125.
1914d. On the breeding habits of Hypsirhina enhydris and Herpeton ten-

taculatum (the tentacle snake). Jour. Nat. Hist. Soc. Siam, vol. 1,

Aug. 1914, pp. 126-127.
1914e. Note on a rare lizard (Lygosoma isodactylum). Jour. Nat. Hist.

Soc. Siam, vol. 1, 1914, pp. 127-128.

1915. The snakes of Bangkok. Jour. Nat. Hist. Soc. Siam, vol. 1, 1915,
pp. 173-185.

1915a. List of the snakes at present known to inhabit Siam. Jour. Nat.

Hist. Soc. Siam, vol. 1, 1915, pp. 211-215.
1915b. Miscellaneous notes. Jour. Nat. Hist. Soc. Siam, vol. 1, no. 4,

pp. 252-256.
1915c. On the breeding habits and color changes in the lizard Calotes

mystaceus. Jour. Nat. Hist. Soc. Siam, vol. 1, 1915, pp. 256-257.
1915d. Notes on some snakes from Siam. Bombay Jour. Nat. Hist. Soc,

vol. 23, 1915, pp. 784-789.
1915e. In K. G. Gardiner, Notes on the fauna and flora of Ratburi and

Ratchaburi districts. Reptiles and Batrachians. Jour. Nat. Hist.

Soc. Siam, vol. 1, no. 3, Mar. 1915, pp. 153-156.

1916. On reptiles and batrachians from the coast and islands of south-east

Siam. Jour. Nat. Hist. Soc. Siam, vol. 1, no. 4, 1916, pp. 239-249.
1916a. Description of five tadpoles from Siam. Jour. Nat. Hist. Soc. Siam,

vol. 2, no. 1, 1916, pp. 37-43, two pis.
1916b. Description of three new lizards and a new snake from Siam. Jour.

Nat. Hist. Soc. Siam, vol. 2, no. 1, 1916, pp. 44-47, 1 pi.
1916c. A list of the crocodiles, tortoises, turtles, and lizards at present

known to inhabit Siam. Jour. Nat. Hist. Soc. Siam, vol. 2, no. 1,

1916, pp. 48-57.
1916d. On a collection of reptiles and batrachians from peninsular Siam.

Jour. Nat. Hist. Soc. Siam, vol. 2, 1916, pp. 148-171.
1916e. On the frogs of the genus Oxyglossus. Jour. Nat. Hist. Soc. Siam,

vol. 2, 1916, pp. 172-175, 1 pi.
1916f. Note on a rare sea snake (Thalassophis anomalus) from the coast

of Siam. Jour. Nat. Hist. Soc. Siam, vol. 2, 1916, pp. 176-177, 1 pi.

1917. Description of new reptiles and a new batrachian from Siam. Jour.
Nat. Hist. Soc. Siam, vol. 2, no. 3, May, 1917, pp. 221-225, 2 pis.

1178 The University Science Bulletin .

1917a. A list of the batrachians at present known to inhabit Siam. Jour.

Nat. Hist. Soc. Siam, vol. 2, no. 3, May, 1917, pp. 226-231, 256.
1917b. On tadpoles from Siam. Jour. Nat. Hist. Soc. Siam, vol. 2, 1917,

pp. 261-275, 2 pis.
1917c. A new snake and a new frog from Siam. Jour. Nat. Hist. Soc. Siam,

vol. 2, 1917, pp. 276-278.
1917d. Preliminary diagnosis of four new sea snakes. Jour. Nat. Hist. Soc.

Siam, vol. 2, Dec, 1917, pp. 340-342.

1918. Description of a new snake (Opistliotropis spenceri) from Siam.
Jour. Nat. Hist. Soc. Siam, vol. 3, 1918, p. 13.

1919. Remarks on Col. Wall's identification of Hijdrophis cyanocinctus.
Bombay Jour. Nat. Hist. Soc. 1919, pp. 682-683.

1919a. The lizards of the genus TropidopJiorus in Siam, with descriptions
of two new species. Jour. Nat. Hist. Soc. Siam, vol. 3, 1919, pp.
223-228.

1919b. Crocodilus . iamensis. Jour. Nat. Hist. Soc. Siam, vol. 3, 1919, pp.
217-221, pis. 4-6.

1920. On sea snakes from the coasts of the Malay Peninsula, Siam and
Cochin China, Jour. Fed. Malay Mus. Kuala Lumpur, vol. 10,
no. 1, 1920, pp. 1-63, pi. map.

1921. A new name for the frog Rana pullus. Jour. Nat. Hist. Soc. Siam,
vol. 4, 1921, p. 193.

1922. Notes on reptiles and batrachians from Siam and Indo-China
(no. 1). Jour. Nat. Hist. Soc. Siam, vol. 4, 1922, pp. 203-214, pi. 8.

1922a. The frogs allied to R. doriae. Jour. Nat Hist. Soc. Siam, vol. 4,
1922, pp. 215-219, 1 pi.

1923. Notes on reptiles and batrachians from Siam and Indo-China
(No. 2). Jour. Nat. Hist. Soc. Siam, vol. 6, 1923, pp. 47-53, pi. 5.

1923a. A review of the lizards of the genus TropidopJiorus on the Asiatic
mainland. Proc. Zool. Soc. London, 1923, pp. 775-781.

1924. New tree frogs from Indo-China and the Malay Peninsula. Proc.
Zool. Soc. London, pt. 1, 1924, pp. 225-234, pis. 1-3.

1926. Monograph of the sea snakes Hydrophiidae. British Museum,
London, 1926, pp. 1-17, 1-130, pis. 1-2, text figs. 1-35.

1926a. The function of the "funnel" mouth of the tadpoles of Megalophrys,
with a note on M. aceras Boulenger. Proc. Zool. Soc. London, 1926,
pp. 983-988, 2 text figs.

1927. Contributions to the herpetology of the Indo-Australian Region.
Proc. Zool. Soc. London, 1927, pp. 199-225, pis. 1-2; text figs. 1-4.

1928. Two vipers new to Siam. Jour. Nat. Hist. Soc. Siam, vol. 7, no. 3,
1928, pp. 194-195.

1930. The Reptilia and Amphibia of the Malay Peninsula . . ., Bull.
Raffles Mus. no. 3, April, 1930, pp. 1-149, figs. 1-13.

1930a. Ovoviviparity in sea snakes. Nature, no. 3180, 1930, p. 568.

1931. The fauna of British India including Ceylon and Burma. Reptilia
and Amphibia, vol. 1, Loricata, Testudines, Mar., 1931, pp. i-xxvi,
1-185, 41 text figs., 2 pis.

1932. Some notes on the monitors. Jour. Bombay Nat. Hist. Soc, vol. 35,
1932, pp. 613-619, figs.

Contribution to Herpetology of Thailand 1179

1933. Remarks on some Old World Geckoes. Rec. Ind. Mus., vol. 35,
1933, pp. 9-19, figs.

1935. The fauna of British India including Ceylon and Burma. Reptilia
and Amphibia. Vol. II. Sauria, Feb. 7, 1935, pp. i-xiii, 1-440,
94 text figs., 1 pi. map.

1935a. The sea snakes ( Hydrophiidae) Dana Report No. 8. Copenhagen,
July 10, 1935, pp. 1-6, text fig., map.

1937. Draco blanfordi and its allies. Bull. Raffles Mus., Singapore, no.
13, 1937, pp. 75-76, pi. 8.

1937a. A Bangkok python. Jour. Siam Soc. Nat. Hist. Suppl., vol. 11,

1937, pp. 61-62.
1937h. A review of the genus Lygosoma ( Scincidae, Reptilia) and its allies.

Rec. Ind. Mus., vol. 29, 1937, pp. 213-234, text figs.
1939. A revision of the Acrochordinae (Snakes). Ann. Mag. Nat. Hist.

Soc, London, 1939, ser. 11, vol. 3, pp. 393-395.
1943. The fauna of British India, Ceylon and Burma, including the whole

Indo-Chinese Sub-region. Reptilia and Amphibia. Vol. Ill, Ser-

pentes, Dec. 1943, pp. i-xii, 1-583, 163 text figs.
Suvatti, C.

1938. A checklist of aquatic fauna in Siam. Bangkok, 1938. Amphibia
Testudines Crocodilia, pp. 81-97.

1950. Fauna of Thailand. Bangkok ( Dept. of Fisheries ) 1950, pp. 1-1100.
Steindachner, Franz

1867. Reise der Osterreichischen Fregatte Novara, um die Erde in 1857,
1858, 1859. Amphibien, Zool. Theil, Band 1, 1867, pp. 1-70, pis.
1-5.
Taylor, Edward H.

1934. Zoological results of the third de Schauensee Siamese Expedition.
Ft. 3.; amphibians and reptiles. Proc. Acad. Nat. Sci., Philadelphia,
vol. 86, June 13, 1934, pp. 281-310, text figs. 1-4, pi. 17.

Tweedie, M. W. F.

1952. The paradise green snake. Malay Nat. Jour. vol. 7, no. 2, 1952,
p. 67, 1 pi.
Wall, F.

1926. The reticulate python, Python reticulatus (Schneider). Jour. Bom-
bay Nat. Hist. Soc, vol. 31, 1926, pp. 84-90.
Werner, F.

1926. Neue oder wenig bekannte Schlangen aus dem Wiener natur-
historischen Staatsmuseum, Teil 3, Sitz. Ber. Ak. Wiss. Wien,
Abt. 1, Bd. 135, 1926, pp. 243-257.

Contribution to Herpetology of Thailani

Mar-

k

Wf4

5 1958
TY

INDEX

PAGE

abbotti, Goniocephalus 1042

Acanthosaura 1095

armata 1095

crucigera 1095

aceras, Megophrys 1077, 1079

aceras, Megophrys montana .... 1079

Acrochordinae 1134

Acrochordus granulatus 1134

javanicus 1134

aenea, Rana 1039

Aepyurus eydouxi 1045

affinis, Cnemaspis 1041

agaardi, Hydrophis torquatus . 1046

Agama armata 1095

versicolor 1098

Agamidae 1083

Agkistrodon rhodostoma 1168

Ahaetulla ahaetulla ahaetulla . . 1142

caudolineata 1044

cyanochloris 1044

formosa 1044

subocularis 1044

ahaetulla, Ahaetulla ahaetulla 1142

ahaetulla, Coluber 1142

albiceps, Typhlops 1043

Aleuroscalabotes felinus 1041

albolabris, Trimeresurus 1169

allapallensis, Mabuya 1102

alticola, Rana 1039

amboinensis, Cuora 1040

Amphiesma flavipunetatum 1151

Aniyda cartilagenea 1041

angularis, Cyrtodactylus 1041

anguinoides, Ophioscincus . . 1043

Aniliidae 1043

annamensis, Microhyla 1040

annamensis, Parahelicops 1158

annandalei, Kolpophis 1045

annandalii, Hieremys 1041

annectens, Microhyla 1040

anomalus, Thalassophis 1045

armata, Acanthosaura 1095

armata, Agama 1095

armatus armatus, Goniociphalus, 1095
armatus crucigerus,

Goniocephalus 1095

PAGE

asper, Bufo 1075

asper, Philautus 1040

atra, Opisthotropis 1158

harroni, Oligodon 1044

baska, Batagur 1041

Batagur baska 1041

belliana belliana, Liolepis 1100

belliana, Uromystax 1100

bengalensis nebulosus, Varanus, 1101

berdmorei, Engystoma 1070

berdmorei, Microhyla 1070

berdmorei, Tropidophorus .... 1043

bibroni, Pelochelys 1041

bimaculatus, Philautus 1040

bimaculatus, Rhacophorus ... 1040

Bitia hydroides 1045

bivirgata, Maticora 1045

blanfordi, Draco 1083, 1086, 1091

blythi, Rana macrodon 1053

boa, Haplopeltura 1043

bocourti, Enhydris 1045

Boiga cyanea 1143

cynodon 1145

dendrophila dendrophila 1044

multimaculata 1142

Boiginae 1142

boonsongi, Parahelicops 1156

borbonica, Bufo 1039

bowringi, Riopa 1043

braminus, Typhlops 1043

brevis, Euprepis 1102

brevis, Mabuya 1102

brevipalmata, Cyrtodactylus . 1041

brookei, Hydrophis 1046

buccata, Coluber 1159

buccata, Homalopsis 1159

Bufo asper 1075

borbonica 1039

macrotis 1039

melanostictus 1076

parvus 1039

penangensis 1039

Bufonidae 1075

Bungarus candidus 1162

butleri, Microhyla 1040

Caecilia glutinosa 1047

1182

The University Science Bulletin

PAGE

Caeciliidae 1047

caerulescens, Hydrophis 1167

caerulescens, Hydrus 1167

Calamaria pavimentatus 1044

Callophis gracilis hughi 1045

maculiceps 1045

Calluella guttulata 1040

Callula pulchra 1072

Calotes 1096

Calotes cristatellus 1096

emma 1097, 1099

flowed 1097

mierolepis 1096, 1097

mystaceus 1097, 1100

versicolor 1097, 1098

cancrivora cancrivora, Rana . . . 1039

cancrivora raja, Rana 1039

candidus, Bungarus 1162

candidus, Coluber 1162

capucinus, Lycodon 1136

carinatus, Pareas 1128

carinatus, Zaocys 1044

carinense, Megophrys 1077

cartilagenea, Amyda 1041

Caudata 1038

caudolineata, Ahaetulla 1044

Cerberus rhynchops 1045

clialconota, Rana 1064

Chelonia mydas 1040

Cheloniidae 1040

chinensis, Enhydris 1045

Chitra indica 1041

Chirixalus doriae 1040

chrysarga, Rhabdophis . 1151, 1156

chrysargus, Tropidonotus 1156

Chrysopelea ornata 1146

paradisi 1044

cinereus, Oligodon 1138

Cnemaspis afflnis 1041

siamensis 1041

cocincinus, Physignathus 1101

collaris triangularis, Sibynophis, 1130

colletti, Rhacophorus 1040

Coluber ahaetulla 1142

buccatus 1159

candidus 1162

korros 1136

mucosus 1135

ornatus 1146

PAGE

radiatus 1134

Colubridae 1130

Colubrinae 1134

colubrina, Laticauda 1045

condenarius indochinensis,

Psammophis 1044

coriacea, Dennochelys 1040

Coronella cyclurus 1136

Cosymbotus craspedotus 1041

platurus 1082

crassicollis, Siebenrockiella .... 1041

cristatellus, Calotes 1096

Crocodilus siamensis 1081

Crocodylidae 1081

Crocodylus siamensis 1081

palustris 1081

porosus 1081

Crotalidae 1168

crucigerus, Goniocephalus

armatus 1042

cubitalis, Rana 1058

cyanea, Boiga 1143

cyaneum, Triglyphodon 1143

cyanochloris, Ahaetulla 1044

cyanocinctus, Hydrophis 1046

cyanolaemus, Draco .... 1083, 1085

cyanophlyctis, Rana 1039

Cyclemys amboinensis 1040

dentata 1040

mouhotii 1040

cyclurus, Coronella 1136

cyclurus, Oligodon 1136

Cylindrophis rufus rufus 1043

cynodon, Boiga 1145

Cyrtodactylus angularis 1041

brevipalmatus 1041

consobrinoides 1041

intennedius 1081

oldhami 1041

peguensis 1041

pulchellus 1081

Damonia subtrijuga 1041

Dasia olivacea 1112

davidsonii, Dryocalamus 1044

dawsoni, Lygosoma 1102

dendrophila dendrophila, Boiga, 1044

Dendrophis pictus 1142

dentata, Cyclemys 1040

Dermochelidae 1040

Contribution to Herpetology of Thailand

1183

PAGE

Dermochelys coriacea 1040

deschauenseei, Natrix 1151

diadema, Hydrophis torquatus, 1046

diardi, Typhlops 1125

diardi miilleri, Typhlops 1125

Dipsadidae 1126

Dipsas multimaculata 1142

rubescens 1145

divergens, Draco . . 1083, 1084, 1085

Dogania subplana 1041

doriae, Chirixalus 1040

doriae, Goniocephalus 1042

doriae, Leiolopisma 1119

doriae, Rana 1039

Draco 1083

Draco blanfordi 1083, 1086, 1091

cyanolaemus 1083, 1085

divergens 1083, 1084, 1085

Bmbriatus 1084, 1085

formusus 1083, 1086

haasei 1083, 1084, 1088

maculatus, 1083, 1084, 1085, 1089

major 1091

melanopogon . 1084, 1086, 1093

microlepis 1083, 1086

nigriappendiculata 1093

punctatus 1083, 1086

quinquefasciatus .... 1083, 1084

taeniopterus 1083, 1086

volans 1083, 1085

whiteheads 1083, 1084, 1085, 1086

Dryocalamus davidsoni 1044

Dracunculus maculatus 1089

Dryophiops rubescens 1145

Dryophis mycterizans 1148

nasutus 1148

prasinus 1147

pulverulentus 1148

xanthozonia 1147

dumerilii dumerilii, Varanus . . 1101

Elachyglossa gyldenstolpi 1039

Elaphe rlavolineata 1044

porphyracea porphyracea . . . 1044

radiata 1134

taeniura 1044

Elapidae 1162

elongata, Testudo 1041

emma, Calotes 1097, 1099

emys, Testudo 1041

Engystoma berdmorei 1070

PAGE

Enhydrina schistosa 1163

Enhydris bocourti 1045

chinensis 1045

enhydris 1159

jagori 1045

plumbea 1159

smithi 1045

enhydris, Enhydris 1159

enhydris, Hydrus 1159

Eretmochelys imbricata 1040

erythraea, Rana 1064

Eumeces quadrilineatus 1110

quadrivirgatus 1110

siamensis 1109

eunice, Leiolopisma, 1119, 1121, 1125

Euprepis brevis 1102

longicaudata 1109

macularius 1102

olivaceus 1112

eydouxi, Aepyurus 1045

fasciatus atriceps, Hydrophis . . 1046

fasciatus, Rungarus 1045

fasciatus, Lycodon 1044

fasciolatus, Simotes 1136

feae, Megophrys 1077

felinus, Aleuroscalabotes 1041

fimbriatus, Draco 1084, 1085

flaveseens, Varanus 1101

flaviceps, Rungarus 1045

flaviceps, Macropisthodon 1045

flavipunctata, Natrix piscator . . 1151
flavipunctatum, Amphiesma . . . 1151

flavolineata, Elaphe 1044

flowed, Calotes 1097

formosa, Ahaetulla 1044

fonnosus, Draco 1083, 1086

fowleri, Microhyla 1070

fremitus, Hemidactylus 1082

garnoti, Hemidactylus 1041

gecko, Gekko 1083

Gekko gecko 1083

smithii 1041

gecko, Lacerta 1083

Gekkonidae 1081

geminatus, Sibynophis 1043

Geoemyda grandis 1041

spengleri 1040

spinosa 1040

trijuga edeniana 1041

glandulosa, Rana 1040

1184

The University Science Bulletin

PAGE

glutinosus, Ichthyophis 1047

Glyphoglossus molossus 1072

Goniocephalus 1095

Goniocephalus abbotti 1042

armatus armatus 1095

armatus crucigerus 1095

lepidogaster 1042

doriae 1042

Gonyosoma o.xycephalum 1044

gracilis gracilis,

Microcephalophis 1045

gracilis hughi, Callophis 1045

grahami, Sibynophis 1134

gramineus, Trimeresurus 1173

grandis, Geoemyda 1041

granulatus, Acrochordus 1134

granulatus, Hydrus 1134

groundwateri, Natrix 1151

guttulata, Calluella 1040

gyldenstolpei, Elachoglossa . . . 1039

gyldcnstolpei, Ophioscincus . . . 1043

Gymnodactylus 1081

intermedins 1081

pnlchellus 1081

haasei, Draco 1083, 1084, 1088

haasei, Rana 1040

hamptoni, Pareas 1043

Hamadryas hannah 1045

hannah, Hamadryas 1045

hascheana, Rana 1039

hardwicki, Lapemis 1163

hasselti hasselti, Megophrys . 1077

hasselti, Leptobrachinm 1077

helleri, Natrix 1152

Hemidactylus frenatns 1082

garnoti 1041

Hemiphyllodactylus typus typus, 1041

herberti, Riopa 1043

Herpeton tentaculatum 1045

heymonsi, Microhyla 1040

Hieremys annandalii 1041

Hinnlia indicum 1113

Homalopsinae 1159

Homalopsis buccata 1159

plumbea 1159

horridus, Philautus . . : 1040

humeralis, Rana 1039

hydroides, Bitia 1045

Hydrus caerulescens 1167

PAGE

enhydrus 1159

Hydrophiidae 1163

Hydrophis brookei 1046

caerulescens 1166, 1167

cyanocinctus 1046

fasciatus atriceps 1166

holbrookei 1166

klossi 1046

mamillaris 1164

ornatus ornatus 1046

schistosus 1163

torquatus agaardi 1046

torquatus diadema 1046

Hydrostentor pantherinus 1050

Hydrus granulatus 1134

Hylidae 1039

Hyla annectens 1039

chalconota 1064

erythraea 1064

leucomystax 1068

Ichthyophis glutinosus 1047

monochrus 1047

imbricata, Eretmochelys 1040

imprissa, Testudo 1041

inas, Natrix 1151

indica, Chitra 1041

indicum, Hinulia 1113

indicum, Lygosoma 1113

indicus indicus, Sphenomorphus, 1113

inornata, Microhyla 1040

inornatus, Simotes 1138

intermedins, Cyrtodactylus 1081

intermedins, Gymnodactylus . . 1081

intestinalis, Maticora 1045

isodactylus, Riopa 1043

jagori, Enhydris 1045

javanicus, Acrochordus 1043

javanicus, Xenodennus 1043

jerdoni siamensis, Kerilia 1045

joynsoni, Oligodon 1044

Kalophrynus pleurostigma 1040

Kaloula mediolineata 1040

pulchra 1072

pulchra pulchra 1072

kanburiensis, Trimeresurus ... 1169

kaouthia, Naja naja 1162

Kerilia jerdoni siamensis 1045

klossi, Hydrophis 1046

kochangae, Rana 1039

Contribution to Herpetology of Thailand

1185

PAGE

kohtaoensis, Leiolopisma 1119

Kolpophis annandalei 1045

koratensis, Riopa 1043

korros, Coluber 1136

korros, Ptyas 1136

kuhli, Ptychozoon 1041

kuhli, Rana 1057

Lacerta gecko 1083

laevis martensi, Oxyglossus . 1049

laevis martensii Phrynoglossus . 1049

laevis, Pareas 1043

Lapemis hardwickii 1163

laticeps, Rana 1039

Leiolopisma 1119

doriae 1119

eunice 1119, 1121, 1125

kohtaoensis 1119

melanostictum 1119, 1125

pranensis 1119

reevesi 1125

reevesi reevesi 1119, 1121

rupicola 1119

siamensis 1119, 1123

tavesae 1119

vittigerum vittigerum 1119

Lepidochelys olivacea 1040

lepidogaster, Goniocephalus . . . 1042

Leptobrachium hasselti 1077

leptoglossa, Rana 1061

leucomystax, Hyla 1068

leucomystax leucomystax,

Rhacophorus 1068

leucomystax leucomystax,

Polypedates 1068

lima, Ooeidozyga 1048

limnocharis, Rana 1051

limnoeharis limnocharis, Rana. . 1051

lineatus, Typhlops 1043

Liolepis belliana belliana 1100

lionatus, Ptychozoon 1041

Lissonota maculata 1116

longicaudata, Mabuya . . 1109, 1113

longipes, Megophrys 1077

luctuosa, Rana 1040

Lycodon capucinus 1136

fasciatus 1044

laoensis 1044

subcinctus 1044

Lygosoma dawsoni 1102

indicum 1113

quadrupes 1042

rupicola 1119

Mabuya allapallensis 1102

brevis 1 102

longicaudata 1109, 1113

macularia 1102

macularia malcolmi . . 1103, 1107

macularia postnasalis . . 1103, 1105

macularia quadrifasciata . 1103

madaraszi 1102

multifasciata multifasciata . 1107

praesigne 1042

rugifera 1042

macrodactylus, Rana 1039

macrodon blythi, Rana 1053

Macropisthodon flaviceps 1045

rhodomelas 1045

macrodon, Rana 1053

macrognatha macrognatha,

Rana 1054

macrops macrops,

Pseudoxenodon 1149

macrops, Tropidonotus 1149

macrotis, Bufo 1039

macularia, Mabuya 1102

macularius, Euprepis 1102

macularius, Pareas 1126

maculata, Lissonota 1116

maculatus, Draco 1083

1084, 1085, 1089

maculatus, Dracunculus 1089

maculatus, Sphenomorphus ... 1113

maculiceps, Callophis 1045

madaraszi, Mabuya 1102

magnapustulosus, Micrixalus 1066

major, Draco 1091

malaccanus, Pareas 1043

malcolmi, Microhyla 1070

mamillaris, Hydrophis 1166

margaritophorus, Pareas 1126

martensii, Phrynoglossus 1049

martensii, Phrynoglossus laevis, 1049

mediolineata, Kaloula 1040

megacephalum, Platysternon 1079

Megalophrys montana aceras . . 1079

Megophrys aceras 1077, 1079

carinense 1077

feae 1077

1186

The University Science Bulletin

PAGE

hasselti hasselti 1077

longipes 1077

nasuta 1077

montana aceras 1079

monticola 1077

pelodytoides 1077

melanopogon, Draco, 1084, 1086, 1093

melanosticum, Leiolopisma, 1119, 1125

melanostictus, Bufo 1076

Micrixalus magnapustulosus . . . 1066

tenasserimensis 1040

Microcephalophis gracilis

gracilis 1045

Microhyla annectens 1040

annamensis 1040

berdmorei 1070

butleri 1040

fowled 1070

heymonsi 1040

inornata 1040

malcolmi 1070

ornata 1040

pulchra 1040

microlepis, Calotes 1096, 1097

microlepis, Draco 1083, 1086

microlepis, Tropidophorus ... 1043

miopus, Rana 1039

modesta, Natrix 1151

molossus, Glyphoglossus 1072

molurus bivittatus, Python 1043

Monitor nebulosus 1101

monochrous, Ichthyophis 1038

monticola, Megophrys 1077

mortenseni, Rana 1039

mouhotii, Cyclemys 1040

mucosus, Coluber 1135

mncosus, Ptyas 1135

miilleri, Typhlops diardi 1125

multifasciata, Mabuya 1107

multifasciatus, Scincus 1107

nmltimaeulata, Boiga 1142

multimaculata, Dipsas 1142

mutilatus, Peropus 1082

mystaceus, Calotes 1097, 1100

mycterizans, Dryophis 1148

Naja naja kaouthia 1162

siamensis 1162

naja kaouthia, Naja 1162

nasuta, Megophrys 1077

PAGE

nasutus, Dryophis 1148

Natricinae 1149

Natrix 1150

deschauenseei 1151

groundwateri 1151

helleri 1152

inas 1151

modesta 1151

percarinata 1151

piscator piscator 1151

piscator flavipunctata 1151

stolata 1153

trianguligera 1151

nebulosus, Monitor 1101

nebulosus, Varanus 1101

nebulosus, Varanus bengalensis, 1101

Nectophryne hosii 1039

nicobariensis, Rana 1039

nigriappendiculata, Draco .... 1093

nigrocincta, Rhabdophis. . 1151, 1155

nigrocinctus, Tropidonotus 1155

nigropalmatus, Rhacophorus . . 1040

nigrovittata, Rana 1061

Notochelys platynota 1041

novae-guinea, Dibamus 1043

nuchalis, Plagiopholis 1044

oldhami, Cyrtodactylus 1041

Oligodon barroni 1044

cinereus 1138

cyclurus 1136

joynsoni 1044

quadrilineatus 1140

purpurascens 1044

taeniata 1044

violaceous 1138

olivacea, Dasia 1112

olivacea, Lepidochelys 1040

olivaceus, Euprepis 1112

Ooeidozyga lima 1048

ornata, Chrysopelea 1 146

ornata, Microhyla 1040

ornatus ornatus, Hydrophis .... 1046

ornatus, Coluber 1146

Ophioscincus anguinoides 1043

roulei 1043

gyldenstolpei 1043

Opisthotropis 1158

atra 1158

spenceri 1158

Contribution to Herpetology of Thailand

1187

PAGE

oxycephalum, Gonyosoma 1044

Oxyglossis lima 1048

Oxyglossis laevis martensi .... 1049

palustris, Crocodylus 1081

pantherina, Rana tigerina 1050

pantherinus, Hyclrostentor .... 1050

paradisi, Chrysopelea 1044

Pareas carinatus 1128

hamptoni 1043

laevis 1043

macularius 1126

malaccanus 1043

margaritophorus 1043, 1126

Parihelicops annamensis 1158

boonsongi 1156

parvus, Bufo 1039

peguensis, Cyrtodactylus 1041

Pelamus platurus 1045

Pelochelys bibroni 1041

Pelobatidae 1077

pelodytoides, Megophrys 1077

percarinata, Natrix 1151

Peropus mutilatus 1082

Philautus asper 1040

bimaculatus 1040

horridus 1040

petersi 1040

pictus 1040

vittatus 1040

Phrynoglossus laevis martensii . . 1049

Physignathus cocineinus 1101

pictus, Dendrophis 1142

pileata, Rana 1039

piscator flavipunctata, Natrix. . 1151

piscator piscator, Natrix 1151

Plagiopholis nuchalis 1044

platurus, Cosymbotus 1082

platurus, Stellio 1082

Platysternidae 1079

Platysternon megacephalum . . . 1079

Plestiodon quadrilineatum .... 1110

pleurostigma, Kalophrynus .... 1040

plumbea, Enhydris 1159

plumbea, Homalopsis 1159

Polypedates leucomystax

leucomystax 1068

popeorum, Trimeresurus 1169

1171, 1174);

porosus, Crocodylus 1081

porphyracea porphyracea,

Elaphe 1044

praesigne, Mabuya 1042

pranensis, Leiolopisma 1119

prominanus, Rbacophorus .... 1040

prasinus, Dryophis 1147

Psammodynastes pulverulentus, 1146
Psammophis condenarius

indochinensis 1044

Pseudorhabdion longiceps 1044

Pseudoxenodon macrops

macrops 1149

Ptyas mucosus 1135

korros 1136

Ptychozoon lionatum 1041

kuhli 1041

pulchellus, Cyrtodactylus 1081

pulchra, Callula 1072

pulchra, Kaloula 1072

pulchra, Microhyla 1040

pulverulentus, Psammodynastes, 1146

pulverulentus, Dryophis 1148

punctatus, Draco 1083, 1086

puniceus, Trimeresurus 1046

purpurascens, Oligodon 1044

purpureomaculatus andersoni,

Trimeresurus 1046

purpureomaculatus purpureo-
maculatus, Trimeresurus . . . 1169

Python reticularis 1043

molurus bivittatus 1043

quadrilineatum, Plestiodon .... 1110

quadrilineatus, Eumeces 1110

quadrilineatus, Oligodon 1140

quadrilineatus, Simotes 1140

quadrivirgatus, Eumeces 1110

quadrivittatum, Siaphos 1042

quadrupes, Lygosoma 1042

quinquefasciatus, Draco . 1083, 1084

radiata, Elaphe 1134

Rana aenea 1039

alticola 1039

cancrivora cancrivora 1039

cancrivora raja 1039

chalconota 1064

cubitalis 1058

cyanophlyctis 1039

doriae 1039

erythraea 1064

1188

The University Science Bulletin

PAGE

glandulosa 1040

haasei 1040

hascheana 1039

humeralis 1039

koehangae 1039

kuhli 1057

laticeps 1039

limnocharis limnocharis .... 1051

luctuosa 1040

maerodactyla 1039

macrodon 1053

macrodon blythi 1053

macrognatha maerognatha . . 1054

miopus 1039

mortenseni 1039

nicobariensis 1039

nigrovittata 1061

pileata 1039

scutigera 1039

signata 1039

tasanae 1039

tigerina pantherina 1050

tigrina pantherina 1050

Ranidae , 1048

Reptilia 1079

reevesi reevesi,

Leiolopisma 1119, 1121

reticulatus, Python 1043

Rhabdophis chrysarga . . 1151, 1156

nigricincta 1151, 1155

stolata 1151, 1153

subminiata 1151

subminiata helleri ... 1151, 1152

subminiata subminiata 1045

Rhacophoridae 1068

Rhacophorus bimaculatus 1040

colletti 1040

leucomystax leucomystax . . 1068

nigropalmatus 1040

prominanus 1040

rhodomelas, Macropisthodon 1045

rhodostoma, Agkistrodon 1168

rhodostoma, Trigonocephalus . 1168

rhynchops, Cerberus 1045

Riopa bowringi - 1043

herberti 1043

koratense 1043

isodactyla 1043

robinsoni, Tropidophorus 1043

PAGE

roulei, Ophioscincus 1043

rubeseens, Dipsas 1145

rubescens, Dryophiops 1145

rudicollis, Varanus 1101

rufus rufus, Cylindrophis 1043

rugifera, Mabuya 1042

rupicola, Leiolopisma 1119

rupicola, Lygosoma 1119

russelli siamensis, Vipera 1046

salvator salvator, Varanus 1101

Sauria scaphiodontophis 1132

sehistosa, Enhydrina 1163

schistosus, Hydrophis 1163

schneideri, Typhlops 1125

Scincidae 1042

Scineus multifasciatus 1107

scriptus, Liopeltis 1044

siamensis, Cnemaspis 1041

siamensis, Crocodilus 1081

siamensis, Crocodylus 1081

siamensis, Eumeces 1109

siamensis, Kerilia jerdoni 1045

siamensis, Leiolopisma. 1119, 1123

siamensis, Naja 1162

siamensis, Typhlops 1125

Sibynophinae 1130

Sibynophis collaris triangularis, 1130

gemminatus 1043

grahami 1134

sinensis 1134

Siebenrockiella erassicollis ... 1041

Simotes fasciolatus 1136

inornatus 1138

quadrilineatus 1140

smithi 1136

taeniatus 1140

sinensis, Sibynophis 1134

smithi, Enhydris 1045

smithi, Gekko 1041

smithi, Simotes 1136

spenceri, Opisthotropis 1158

spengleri, Geoemyda 1040

spinosa, Geoemyda 1040

Sphenomorphus indicus 1113

indicus indicus 1113

maculatus 1116

zebratus 1115

stejnegeri, Trimeresurus 1169

Stellio platurus 1082

Contribution to Herpetology of Thailand

1189

PAGE

stolata, Natrix 1153

stolata, Rhabdophis 1151, 1153

subcinctus, Lycodon 1044

subminiata helleri, Natrix 1151

subminiata subminiata, Natrix. . 1151

subminiata belleri, Rhabdophis, 1152

subminiata, Rhabdophis 1151

subocularis, Ahaetulla 1044

subplana, Dogiana 1041

subtrijuga, Damonia 1041

taeniatus, Oligodon 1044

taeniatus, Simotes 1140

taeniopterus, Draco 1083, 1086

taeniura, Elaphe 1044

Takydromus sexlineatus 1042

tasanae, Rana 1039

tavesae, Leiolopisma 1119

tentaculatum, Herpeton 1045

tenasserimensis, Micrixalus 1040

Testudines 1079

Testudinidae 1040

Testudo elongata 1041

emys 1041

impressa 1041

thai, Tropidophorus 1043

Thalassophis anomalus 1045

Thalassophina viperina 1163

Thalassophis viperina 1163

tigerina pantherina, Rana 1050

tigrina pantherina, Rana 1050

tigrina lateralis, Rhabdophis . . 1151

trianguligera, Natrix 1151

triangularis, Sibynophis collaris, 1130

Triglyphodon cyaneum 1143

Trigonoeephalus rhodostoma . 1168

trijuga edeniana, Geoemyda . 1041

Trimeresurus 1168

albolabris 1169

gramineus 1173

kanburiensis 1169

popeorum 1169, 1171, 1174

purpureomaeulatus

purpureomaculatus 1169

stejnegeri 1169

Tropidonotus chrysargus 1156

macrops 1149

nigrocinctus 1155

Tropidophorus berdmorei 1043

PAGE

microlepis 1043

robinsoni 1043

thai 1043

Tylototriton verrucosus 1038

Typhlopidae 1125

Typhlops albieeps 1043

braminus 1043

diardi 1125

diardi mulleri 1125

floweri 1043

lineatus 1043

mulleri 1125

schneideri 1125

siamensis 1125

typus typus,

Hemiphyllodactylus 1041

unieolor, Xenopeltis 1043

Uromystax belliana 1100

Varanidae 1101

Varanus bengalensis nebulosus, 1101

dumerilii dumerilii 1101

flaveseens 1101

nebulosus 1101

rudicollis 1101

salvator salvator 1101

vermiformis, Calamaria 1044

verrucosus, Tylototriton 1038

versicolor, Agama 1098

versicolor, Calotes 1097, 1098

violaceus, Oligodon 1138

Vipera russelli siamensis 1046

Viperidae 1046

viperina, Thalassophina 1163

viperina, Thalassophis 1163

vittatus, Philautus 1040

vittigerum vittigerum,

Leiolopisma 1119

volans, Draco 1083, 1085

wagleri wagleri, Trimeresurus . . 1046

whiteheads Draco 1083

1084, 1085, 1086

xanthozona, Dryophis 1147

Xenoderminae 1043

Xenodermis javanicus 1043

Xenopeltidae 1043

Xenopeltis unieolor 1043

Zaocys carinatus 1044

zebratus, Sphenomorphus 1115

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 14

Report on a Collection of Amphibians and Reptiles from
Harbel, Republic of Liberia

BY

Edward H. Taylor * and Dora Weyer **

The collection of Liberian reptiles and amphibians reported
herein has been brought together by Mrs. Dora Weyer, the junior
author, while residing at The Firestone Rubber Plantations at
Harbel, Liberia. Her husband, Dr. Albert E. Weyer, director of
sanitation, has assisted her to some extent in this work.

The collection contains a number of rarities and certain species
are regarded as new. Field notes appearing in quotation marks are
those of the junior author, sent with the collection.

"All of these specimens come from the main Firestone plantation
save one from the Cavalla plantation — very similar ecologically but
southeast of Harbel on the Cavalla River.

"Ecologically Harbel is in the heaviest rain-forest in Africa. The
soil is a gravelly laterite, with many outcroppings of rocks. The
large Du and Farmington rivers curve back and forth through
the area and there is a multitude of small rivers and creeks tributary
to them fed by water from innumerable swamps. This is an area
of low hills rising from the coastal sand plain which edges the
plantation seaward and lying between each hill and the next is
usually a swamp! In fact, the survey map shows 25 percent of the
entire area is swamp.

"Lastly, of interest from an ecological viewpoint, with the excep-
tion of certain experimental plots, the river edges, and the cutover
swamp lands, the area is covered with dense rubber forest. In the
old rubber forest with trees reaching 75 to 90 feet in height, and
where most of the collecting was done, the ground cover is pretty

* Department of Zoology, Kansas University, Lawrence, Kansas.
* * Firestone Plantations, Harbel, Liberia.

(1191)
6—8050

1192 The University Science Bulletin

well limited to grass and ferns which is cut over three or four
times a year to keep it low. In the young rubber the ground is
densely covered with a kudzulike legume used here for a cover
crop; this dies out once the trees get big enough to shade it.

"Ten miles beyond the plantation borders at Kakala the annual
rainfall is only 80 inches while farther on at Ganta it drops to 60
inches. Here at Harbel the annual rainfall runs from 150 to 200
inches.

"While there are no extensive serpent cults in Liberia such as have
been reported elsewhere in Africa, there are 'snake charmers' or
persons who 'play' with snakes.

"The tribes near Harbel have their 'snake men' who handle or
'play' with the snakes. Incidently, they do not necessarily remove
the venom from the poisonous snakes with which they play. I had
assumed that they did remove the poison. Recently two 'snake men'
were brought into the hospital dead, and the physician, my husband,
questioned the people who brought them in. They came from a
village near the plantation — a Kepelle village — and both of the de-
ceased were full-fledged initiates. Presumably each 'snake-man'
keeps his own poisonous snake either in a basket, in his hut, or loose
in the 'bush' behind his hut. When the time comes that he wishes
to show his prowess, or more likely force tribute from someone, he
goes to the bush and calls his snake. When the snake comes to him
he is supposed to ask it whether it wishes to 'play' in the village.
Only if the snake agrees may he handle it safely before the people —
and this is not done often.

"However in the case in question, the two men together were
playing with the snake, a cobra, in the village when suddenly it
bit one. The other man instantly took it from the hand of the bitten
man, and the snake bit him also. Both died after a short time.

"When questioned by my husband the witnesses answered that
they did not think the second man realized that the snake had actu-
ally bitten his associate, but had thought that the snake was simply
playing, and he merely wanted to handle the snake himself.

"When asked why they thought the men's medicine had not
worked they carefully explained two possibilities:

"(1) The snake may not have 'agreed' to play in the village and
they took it in spite of this, or (2) the men may not have fully per-
formed the initial rites or may have broken the prescribed taboos,
one of which is that they must not have had sexual intercourse for
three days previously.

Amphibians and Reptiles from Liberia 1193

"All this of course is a deadly serious matter to these people. An
examination of the snake showed that it retained the fangs. The
poison sac was intact and still contained some poison."

List* of Amphibians and Reptiles Known to Occur in the Re-
public of Liberia
Amphibia
Gymnophionians

Geotrypes seraphini occidentalis Parker
*Geotrypes angeli Parker
Salientia

*Xenopus tropicalis Gray

Bufo regularis regularis Reuss

Bufo regularis maculatus Hallowell

Bufo togoensis Ahl

Bufo eamerunensis camerunensis Parker

Heinisus marmoratus ( Peters )

Rana albolabris albolabris Hallowell

Rana albolabris parkeriana Loveridge

Rana mascareniensis mascareniensis Dumeril and Bibron

Rana bibroni Hallowell

Rana occipitalis Giinther

Rana oxyrhyncbus oxyrbyncbus Smith

Rana oxyrhynchus gribinguiensis Angel

Rana longirostris Peters
*Rana maccarthyensis Andersson

Rana crassipes alleni ( Barbour and Loveridge )

Rana subsigillata A. Dumeril

Phrynobatrachus giorgii De Witte

Phrynobatrachus perpalinatus Boulenger

Phrynobatrachus francisci Boulenger

Phrynobatrachus natalensis (Smith)

Phrynobatrachus latifrons Ahl

Phrynobatrachus phcatus (Giinther)

Phrynobatrachus alleni Parker

Phrynobatrachus ogoensis brongersmai Parker

Phrynobatrachus liberiensis Barbour and Loveridge

Arthroleptis adolfi-friederici Nieden

Arthroleptis calcaratus ( Peters )

Arthroleptis poecilonotus Peters

Arthroleptis taeniatus Boulenger

Arthroleptis werneri Nieden

Arthroleptis gutterosus Chabanaud

Cardioglossa decorata Barbour and Loveridge

Cardioglossa liberiensis Barbour and Loveridge

Leptopelis viridis (Giinther)

Leptopelis bequaerti (Loveridge)

Leptopelis tessmanni Nieden

* Species occurring in the collection at hand are preceded by an asterisk.

1194 The University Science Bulletin

Peptopelis bocagii (Boulenger)
Hylambates hyloides Boulenger
Hylambates cochranae Loveridge
Hylambates leonardi Boulenger
Megalixalus immaculatus Boulenger
Megalixalus fulvovittatus Cope
Megalixalus fornasinii ( Bianconi )
Megalixalus platyceps Boulenger
Hyperolius concolor (Hallowell)
Hyperolius puncticulatus Pfeffer
Hyperolius platyceps Boulenger
Hyperolius festivus Barbour and Loveridge
Hyperolius pleurotaenides (Boulenger)
Hyperolius ocellatus Giinther
Hyperolius fuscigula Bocage
Hyperolius fusciventris Peters
Hyperolius marmoratus Bapp
Hyperolius picturatus Peters
Hyperolius admetzi Ahl

Reptiles
Crocodiles

Crocodylus niloticus Laurenti **

Crocodylus cataphractus Cuvier

Osteolaemus tetraspis Cope
Turtles

Eretmochelys imbricata imbricata (Linnaeus)

Chelonia mydas mydas (Linnaeus)

Dennochelys coriacea coriacea (Linnaeus)

Lepidochelys olivacea olivacea ( Eschseholtz )

Kinixys homeana Bell
*Kinixys erosa (Schweigger)

Pelusios adansonii (Schweigger)

Pelusios f subniger (Lacepede)

Amyda triunguis Forskal
Amphisbaenians

Amphisbaena liberiensis Boulenger

Amphisbaena cynisca leucura Dumeril and Bibron
Saurians

Hemidactylus mabouia ( Moreau de Jonnes)

Hemidactylus muriceus Peters

Hemidactylus fasciatus Gray

Lygodactylus thomensis Peters

Lygodactylus gutturalis ( Bocage )
*Agama agama africana (Hallowell)

Varanus ( Polydaedalus ) niloticus niloticus ( Linnaeus )

** The name of an author associated with a scientific name may be regarded as being
in the genitive case; hence it would appear that Laurenti is the proper form if one inflects
the name as is done in the title of certain books.

t Pelusios derbianus Gray and Pelusios nigricans castaneus Hewitt are placed in the
synonymy of P. subniger by Mertens and Wermuth 1955. These names have been recog-
nized previously as representing distinct species.

Amphibians and Reptiles from Liberia 1195

*Lacerta echinata echinata Cope
*Mabuya maculilabris maculilabris Gray
*Mabuya blandingii Hallowell

Mabuya bensoni (Peters)

Riopa durum durum Cope
*Riopa femandi ( Burton )
Chamaeleons

*Chamaeleon senegalensis Daudin

Chamaeleon gracilis gracilis Hallowell
Serpents

*Typhlops leprosus sp. nov.
*Typhlops punctatus punctatus Leach

Typhlops manni Loveridge

Python sebae Gmelin
*Calabaria reinhardti (Schlegel)
*Neusterophis variegatus (Peters)
*Natrix anoscopus anoscopus (Cope)
*Natrix firestonei sp. nov.

Lycophidion fasciatum Giinther
*Bothropthalmus lineatus lineatus Peters

Boaedon olivaceus ( Dumeril )
*Boaedon hneatus virgatus (Hallowell)
*Boaedon lineatus lineatus Dumeril, Bibron, and Dumenl
*Hormonotus modestus Dumeril, Bibron, and Dumeril

Chlorophis irregularis (Leach)
*Chlorophis heterodermis (Hallowell)

Phylothamnus nitidus (Giinther)

Gastropyxis smaragdina (Schlegel)

Mehelya guirali (Mocquard)
*Mehelya poensis (Smith)

Coronella coronata ( Schlegel )

Hapsidophrys lineata Fischer

Grayia smythii (Leach)

Rhamnophis aethiopissa aethiopissa Giinther

Thrasops flavigularis (Hallowell)

Dasypeltis scaber Linnaeus

Dasypeltis macrops Boulenger

Crotaphopeltis hotamboeia hotamboeia Laurenti

Crotaphopeltis duchesnii guineensis (Chabanand)

Boiga pulverulenta (Fischer)
*Boiga blandingii (Hallowell)
*Psammophis sibilans phillipsi Hallowell
*Thelotornis kirtlandii kirtlandii (Hallowell)

Miodon acanthias (Reinhardt)

Apparallactus lineatus Peters
*Apparallactus modestus (Giinther)

Dendroaspis viridis ( Hallowell )

Dendroaspis jamesonii (Traill)

Naja nigricollis Reinhardt

Naja goeldii Boulenger

1196 The University Science Bulletin

*Naja melanoleuca Hallowell

*Atractaspis corpulenta corpulenta ( Hallowell )

Atractaspis irregularis Reinhardt
*Causus rhombeatus ( Lichtenstein )

Causus lichtensteini (Jan)
*Bitis gabonica Dumeril, Bibron, and D.imeril
*Bitis nasicornis (Shaw)

Atheris chlorechis ( Schlegel )

Report on the Collection
Geotrypetes angeli Parker

Geotrypetes angeli Parker, Zool. Meded, Leiden, vol. 19, 1936, pp. 10U-101
( type locality Labe, French Guinea, Africa ) ; Loveridge, Proc. New England
Zool. Club. vol. 17, May 20, 1938, p. 65, ( Sanoquelle, near Ganta, Liberia).

Although this species is described in Parker's paper on "The Am-
phibians from Liberia and the Gold Coast," it would seem that it
is known from neither territory. It is presumed that Loveridge's
record represents the first report of the species for Liberia.

Two specimens are at hand, EHT-HMS Nos. 31927, 31928 from
Harbel, Liberia. They yield the following data: (the heads of
both, and the tail of one, have been somewhat injured. The count
here given of the grooves and teeth, if not actual, are very close to
the actual number. )

No. 31927. Eye distinct; snout elongate, projecting considerably
beyond lower jaw; distance between eye and mouth contained
nearly three times in distance between eye and nostril; tentacle be-
low nostril nearly midway between nostril and lip; a total of 31
maxillary-premaxillary teeth, the posterior smallest; about 34 vom-
eropalatine teeth, the posterior much the smallest; dentary series 28;
splenial series 18. In No. 31928 the maxillary-premaxillary teeth
are 29; vomeropalatine 34; dentary series 24, splenial series 20.

The general color (in preservative) is lavender-purplish to violet.
Most of the grooves are grayish white, and the head is somewhat
lighter, perhaps somewhat olive in life.

Measurements in mm. (of 31927, 31928 respectively): Total
length 279, 202; head length, 12.2, 9.2; distance between eyes, 4, 3.3;
eye to tip of snout, 4.15, 4; eye to nostril 3.5, 3; eye to lip, 1.25, 1.05;
greatest diameter of body, 10, 5.

The hidden scales begin somewhere on the anterior two fifths of
the body. At first they are minute and usually to be discovered at
the upper end of the primary grooves. At the beginning of the sec-
ondary grooves they are of larger size and cover much of the space

Amphibians and Reptiles from Liberia 1197

between the primary and secondary grooves; the scales anteriorly
are much longer than wide, farther back they may be nearly as
wide as long. The greatest measurement is always transverse.

Xenopus tropicalis (Gray)

Silurana tropicalis Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 14, 1864, p. 315;
and Proc. Zool. Soc. London, 1864, p. 458. (type locality, Lagos).

Xenopus tropicalis Parker, Proc. Zool. Soc. London, April 16, 1936, p. 156 (dis-
tinguishing characters); Loveridge, Proc. U. S. Nat. Mus., vol. 91, 1941, p.
124, (Gibi and Bromley, Liberia).

A young specimen, EHT-HMS No. 31948 (snout to vent 29 mm.)
is from Harbel, Liberia. The head is much more depressed than
the figures presented by Parker (Joe. dr.) show but this may be
due to the youth of the specimen.

Rami maccartlujensis Andersson

Rana mac earthy ensis Andersson, Arkiv. Zool., vol. 29A, no. 16, 1937, p. 9, figs.
3-4 (type locality, McCarthy Island, Gambia); Loveridge, Proc. U. S. Nat.
Mus. vol. 91, 1941, p. 136 (specimens from Gibi and Bellyella, Liberia).

One specimen, EHT-HMS No. 31947, Harbel, Liberia. Loveridge
had certain specimens in which the legs were reported as being
longer than those in the type. In this specimen the tibiotarsal artic-
ulation reaches the nostril. Both inner and outer metatarsal tu-
bercles are present, the latter connected by a row of low tubercles
with the posterior subarticular tubercle of the fourth toe; vocal sacs
external, opening through a slit on each side of the throat; the vocal
sacs are black and the slits are parallel to each other extending
posteriorly toward the "lower insertion of forearm"; three or four
longitudinal folds on each side of dorsum, the longest extending
more than half the length of body. The tympanum is longer than
high, its greatest diameter equals four fifths or more of the length
of the eye-opening. Three large palmar tubercles, the outer elon-
gate; the posterior subarticular tubercles of fingers very large.

There are two longitudinal series of white dots more or less con-
nected on front and back of thigh, and a broad median light stripe
present.

Kinixys erosa (Schweigger)

Testudo erosa Schweigger, Prodr. Monog. Chelon., 1814, p. 52.
Kinixys erosa Schmidt, Bull. Amer. Mus. Nat. Hist., vol. 39, 1919, pp. 403-406,
Map-3, pi. XI, fig. 1. ( bibliography ) .

A single specimen EHT-HMS No. 31938 is from Harbel, Liberia.
The specimen is young, the greatest over-all length of the carapace
being only 145 mm. While the hinge of the carapace is scarcely
evident to the eye, the carapace is closely pressed against the plas-

1198 The University Science Bulletin

tron behind. The denticulation is strongly marked along the flaring
anterior edge of the carapace, some scutes bearing two or more
denticulate projections. The lateral marginals have single serra-
tions, the posterior ones have a scalloped appearance.
There are four claws on the hind foot, five on the fore foot.

Agama agama africana ( Hallowell )

Tropidolepis Africanus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1844,

p. 171 (type locality, Liberia).
Agama agama africana Loveridge, Proc. U. S. Nat. Mus., vol. 91, no. 3128,

1941, p. 116 (Bellyella, Bendaja).

A single young and two adult male specimens, EHT-HMS No.
31926, 31943, and 31944, from Harbel, Liberia, are in the collection.
The number of preanal pores ("callose preanal scales") situated on
the border of the vent are 12, 12, and 11, respectively.

Lacerta echinata echinata Cope

Lacerta (Zootaca) echinata Cope, Proc. Acad. Nat. Sci. Philadelphia, 1862,

pp. 189-190 (type locality West Africa [probably Liberia, fide Loveridge

loc. tit.]).
Lacerta echinata Barbour and Loveridge, in Strong Contr. Dept. Trop. Med.

and Inst. Trop. Biol. Medicine No. 5, vol. 2, 1930, p. 776. (Plantation 3,

Du River ) .
Lacerta echinata echinata Loveridge, Proc. U. S. Nat. Museum, vol. 91, No.

3128, 1941, pp. 116-117 (Harbel).

A single specimen, EHT-HMS No. 31924, has the following char-
acters :

Part of rostral visible above as long as the median suture between
the anterior nasals; frontonasal longer than wide, touching the ante-
rior loreal laterally; prefrontals longer than wide, their common su-
ture less than half their greatest length, touching both loreals, the
anterior supraocular, and the anterior supraciliary; frontal approxi-
mately one-third longer than wide; frontoparietals nearly as large
as prefrontals but with a longer median suture, touching two supra-
oculars; parietals longer than wide, each with an irregular longitudi-
nal ridge or keel die outer posterior part of the scales depressed; a
well-defined postinterparietal; nostril surrounded by rostral (nar-
rowly), first labial, posterior and anterior nasals; five (four) supra-
labials precede the large subocular; latter followed by a large, —
and one or two very small, supralabial scales; seven or eight infra-
labials; a large mental scale with a labial border much greater than
that of the rostral; three pairs of much enlarged chinshields the
anterior pair forming a median suture, second pair narrowly sepa-
rated anteriorly, third pair separated by ten or more rows of small

Amphibians and Reptiles from Liberia 1199

scales; third chinshield followed behind by two scales. Three supra-
oculars, the third one-third or one-fourth times size of the first and
second; a row of 10 supraciliaries, the first and last largest; about 12
rows of keeled temporal scales in preauricular area; tympanum oval,
much higher than wide; scales on neck and body small, strongly
keeled, the keels in nuchal region parallel to body, those on side of
body running up and back, the keels diagonally placed on the indi-
vidual scales.

Venter covered by six longitudinal rows of large scutes in about
32 transverse rows. The median rows are slightly the smallest, the
rows adjoining these largest, the outer series somewhat inter-
mediate, and in turn bordered by a row of scales somewhat larger
than the lateral body scales; scales on underside of neck small with
a distinct collar of larger scales, which may be preceded by one or
two enlarged scales. Anal scale large, bordered on sides and pos-
teriorly by two rows of scales; 10-11 femoral pores.

Tail above and below with sharply keeled scales, those near the
base — between the tenth and twentieth whorls — with longer mu-
crones extending from the posterior part of the keels; a row of five
or six enlarged keeled scales on outer face of upper arm more or less
continuous with a series of ten on front face of forearm; a similar
series on front face of femur and on tibia, the series being largely
ventral; other scales on limbs variable in size; subdigital scales not
strongly keeled.

Color ( in preservative ) : Top and sides of head nearly uniform
blackish; body generally dark grayish-black with transverse gray-
lines crossing neck and extending down to near the ventral surface;
on body the transverse markings are less distinct, rarely continuous,
often consisting of four or more tiny gray spots; tail dark with ex-
tremely narrow vertical lines of white, the upper ones often alter-
nating with those low on sides of the tail; chin and large scutes of
belly lighter, but with considerable pigment, becoming blackish on
the two outer rows of scutes.

Measurements in mm.: Snout to vent, 76; tail 203; total length.
279.

Remarks: The original description by Cope varies somewhat from
the description given here but this may be due, at least in part, to
a different terminology. The color, as given by Cope is: "Above
bluish-green with about fifteen blackish cross bands; those upon the
nape and rump are narrow, the others broad, dark bordered. Be-
neath yellowish. Head shaded with yellowish."

1200 The University Science Bulletin

Whether this pattern is essentially different or has been modified
in preservation cannot be stated at this time.

We have followed Loveridge in using the trinomial, although
there may be some doubt that Lacerta langi Schmidt is actually a
subspecies of this form.

Mabuya blandingii (Hallowell)

Euprepis Blandingii Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1844, p. 58

(type locality, Liberia).
Mabouia raddonii Gray, Catalogue of the specimens of lizards in the collection

of the British Museum, 1845, p. 11 (type locality, West Africa); Boulenger,

Catalogue of the lizards in the British Museum, 2nd ed., vol. 3, pp. 165-166,

pi. 10, fig. 1 (includes blandingii as a synonym).
Mabuya blandingii Loveridge, Proc. U. S. Nat. Mus. vol. 91, no. 3128, 1941,

p. 117. (Gibi, Bromley, Bendaja, Harbel).

There are several mabuyas in the collection (EHT-HMS Nos.
31930, 31931, 31934, 31937) which are in bad condition (having
been dried). It is thus practically impossible to ascertain relative
lengths of limbs or to discern clearly original markings and colora-
tion. It would appear that more than a single species is indicated.
Some of these certainly belong to blandingii, however, all are in-
cluded here. Loveridge ( loc. cit. ) has indicated that he believes the
variation is considerable, and places all twelve specimens, examined
by him, in this species.

All of our specimens have a lateral white line. Two are charac-
terized by having the frontal touching only the two anterior supra-
oculars, with the prefrontals widely separated by the junction of
the frontal and the frontonasal. There are 30 scalerows about the
body. The body scales are 3-keeled, or the outer parts of the scale
may have also a tubercle. The scales in the median nuchal region
may have four normal keels and the two nuchal scales are not or
but dimly keeled. Two other specimens have the frontal touching
only the second and third supraoculars, the body scales are 3-keeled,
but the nuchal scales may have five or seven keels and the nuchals
12 or 13 distinct keels. There are 30 scalerows about the body. An-
other specimen has the frontal touching the first, second, and third
supraoculars, and there are 30 scalerows present.

Mabuya maculilabris macidilabris (Gray)

Euprepis maculilabris Gray, Catalogue of the specimens of lizards in the col-
lection of the British Museum. 1845, p. 114.

Mabuia maculilabris Boulenger, Catalogue of the lizards in the British museum
(Natural History). 2nd ed., vol. 3, 1887, pp. 164-165, pi. 9, fig. 2.

EHT-HMS No. 31933 in the collection from Harbel, Liberia, has
the following characters: internasals injured, presumably minutely

Amphibians and Reptiles from Liberia 1201

separated; frontonasal broader than long; prefrontals forming a
median suture, in contact with first and second supraoculars; frontal
slightly longer than its distance from the tip of the snout, shorter
than the median combined length of parietals; two frontoparietals;
an interparietal enclosed by the parietals, which are wider than
long; a pair of nuchals (nearly smooth, only back edge suggests
keeling); nostril almost directly above suture between rostral and
first labial; a distinct postnasal, in contact with first labial (or also
very narrowly with second on one side); two loreals the posterior
largest, the anterior barely touching a labial on one side; two pre-
suboculars; a transparent eyelid disc; six or seven supraciliaries;
four supraoculars, frontal touching only second and third; seven
supralabials, the subocular supralabial, the fifth, nearly double
width of two preceding; eight infralabials; two primary, three
secondary temporals; four postoculars.

Arms and legs well developed, the toes reaching the elbow of
adpressed arm; subdigital lamellae without distinct keels or spines;
33 scalerows around neck; 32 around middle of body, most of the
body scales having five distinct keels, some of those on neck behind
parietals with seven or eight; ear bordered anteriorly by four irreg-
ular denticulations; 12 ventral rows of scales smooth; tail regen-
erated, with 85 subcaudals, the regenerated scales wider than the
original scales, which are also distinctly wider than other adjoining
scales; scales on limbs often tricarinate; anal scales scarcely, or not,
larger than the scales which precede them.

Color in preservative: Above brown or olive brown; a distinct
brown band from snout through eye to groin covering three or
three and a half scalerows; a whitish line on supralabials involving
lower part of ear continued to a point somewhat past arm insertion;
no white line on side below brown band: tail generally olive; chin,
venter, and underside of tail whitish.

Measurements in mm: Snout to vent, 74; tail, 127; head length,
18; head width, 13; arm, 22; leg, 32; axilla to groin, 35.

A second specimen EHT-HMS No. 31932 agrees with the pre-
ceding specimen in having the prefrontals in contact, the inter-
parietal enclosed by the parietals, the supranasals minutely
separated, and die frontal touching the second and third supra-
oculars. There are 31 scalerows around the middle of the body;
a lateral brown stripe is present on the side and there is no lateral
white stripe bordering it below. The labial markings in the two
specimens are nearly identical.

1202 The University Science Bulletin

Riopa fernandi (Burton)

Tiliqua fernandi Burton, Proc. Zool. Soc. 1836, p. 62 (type locality, Fernando

Po); Peters Monatsb, Akad. Wiss. Berlin, 1874, p. 372.
Plestiodon harlani Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 2, 1845,

p. 170.
Euprepis striata Hallowell, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, June

1854, p. 98 (type locality, Liberia); Trans. Amer. Philos. Soc, ser. 2, vol. 11,

1857, p. 74, pi. 3, fig. 1.
Euprepis harlani Hallowell, Trans. Amer. Philos. Soc, ser. 3, vol. 11, 1857, p. 75,

pi. 3, fig. 2.
Lygosoma fernandi Boulenger, Catalogue of the Lizards in the British Museum,

2nd Ed. vol. 3, 1887, pp. 304-305.

A specimen, EHT-HMS No. 31942, is from Centersite in River
Area, Harbel, Liberia, Dec. 20, 1950; Dr. A. E. Weyer collector.

The following characters obtain: rostral presenting a curved
convex posterior edge, length of part visible above considerably
greater than length of the common suture between the internasals;
paired frontonasal, both scales broadly in contact with the frontal,
in contact laterally with the anterior loreal; prefrontals small, sep-
arated from each other by a distance nearly equal to the width of
one of them; in contact laterally with both loreals, posteriorly touch-
ing the anterior supraocular and anterior supraciliary; frontal
widened anteriorly, narrowing posteriorly almost to a point, more
than a third longer than its distance from the tip of snout, equal
to or greater than the combined length of the parietals; four supra-
oculars, two bordering the frontal; frontoparietals form a common
suture of less than half their length, larger than the interparietal,
which with a scale posterior to it, separates the parietals; latter
scales much broader than long. Nine supraciliaries, first and last
largest; nostril in a divided nasal; no postnasal anterior loreal
narrower than posterior and slightly higher; a preocular superim-
posed above the anterior of two presuboculars; a row of seven or
eight suboculars separating labials from the semitransparent scaled
lower eyelid; seven supralabials the fifth and sixth below eye; two
primary temporals bordered behind by four secondary temporals;
a pair of somewhat enlarged scales border parietals behind; ear
distinct, oval tympanum deeply sunk, bordered anteriorly within
depression by two scales with rounded edges; seven infralabials;
mental with a larger labial edge than rostral; undivided postmental
touching two labials laterally; first chinshields in contact, second
pair separated by a scale; third pair reduced each separated from
its fellow by five scales.

Limbs pentadactyle, rather short, when adpressed they fail to
meet by a distance equal to width of two scalerows; subdigital

Amphibians and Reptiles from Liberia 1203

lamellae on fingers simple, short, with a suggestion of a median
keel or ridge, 10 under fourth finger; palm covered with smooth
rounded, but somewhat raised scales; toes short, the fourth with 16
lamellae bearing irregular longitudinal keels or ridges; sole covered
with scales similar to those on palm, the posterior row bordering
a row of enlarged scales; no axillary or inguinal depressions or
pockets; 33 scalerows about narrowest point of neck, 34 about
middle of body; four preanal scales, not or scarcely larger than the
scales preceding them; median subcaudals, 87, not wider than
adjoining scales; scales from parietal to above vent, 56. All dorsal
and lateral scales (except head scales and temporals) with three
well-defined keels, the two outer strongest; also rarely, a lateral
ridge or tubercle can be seen on scales; scales on lower posterior
face of thigh with two tubercles. Venter brownish white, each scale
outlined in light brown. On underside of chin and sides of neck
some scales are edged with dark brown, forming brown lines on
the edges of several rows of scales on the neck. Head nearly
uniform dark brown on dorsum, lighter laterally, a black moon-
shaped spot below and behind eye.

Color: The ten dorsal scalerows woodbrown, each scale bordered
by black leaving the dorsal surface with uniform rows of equal-sized
pentagons; the outer scales of the ten may be lighter brown and
their outer sides come into contact with a series of black lateral
spots or bars; these tend to fuse together on the side of the neck
and above the arm to form an elongate, but irregularly edged spot;
along the side of the body are eight or nine vertical or somewhat
diagonal bars of black, often irregular, sometimes two in contact
above or below; scales between bars brownish but lighter than
those on the back; lateral caudal scales lighter than those on sides
of body with indefinite darker areas dorsally and laterally to the tip;
however, towards the latter third the dark scales predominate and
there appears a series of bluish-white spots. These whitish spots (in
certain lights) can be discerned nearly to the base of the tail but
there is small contrast with the other light scales.

Measurements in mm.: Snout to vent, 104; tail, 132; length of
head, 21; width of head, 16; snout to arm insertion, 32; axilla to
groin, 56; arm, 25; leg, 305.

Remarks: This species may be somewhat aquatic, individuals
having been seen to enter shallow water in the rivers. It seems
extremely doubtful, as has been proposed, that this is a subspecific
form of Riopa durum Cope.

1204 The University Science Bulletin

Chamaeleon senegalensis Daudin

Chamaeleo senegalensis Daudin, Histoire naturelle generale et particuliere

des reptiles, vol. 4, 1802-03, pp. 203-209 (type locality, Senegal).
Chamaeleon senegalensis J. E. Gray, Proc. Zool. Soc. London, 1864, p. 471.

A single young specimen is at hand, EHT-HMS No. 31925.

Typhi ops leprosus sp. nov.

Type: EHT-HMS No. 31895 Firestone Plantation, Harbel, Li-
beria, Africa; 1954, collected by Mrs. Dora Weyer.

Diagnosis: A presumed small species having a length of at least
185 mm. the width in length 23-25 times. A preocular present; no
subocular; four supralabials; nasal divided, the suture arising from
the lower outer edge of rostral; gray- white to yellowish, with ir-
regular transverse markings of black and yellowish; 28 scalerows
around body at middle, 30 scalerows around body 3 cm. back of
snout tip. Scales from mouth to vent 388.

Description of type: Snout prominent, its lower edge with a blunt
transverse keel, not hooked; eye clearly visible, the pupil distinct
covered by ocular and posterior edge of preocular; rostral large its
greatest width 3.4 mm., much less than width of head at level of
eyes (5.6 mm.), its posterior edge truncate rather than rounded;
prefrontal about 3.5 times wider than long, separating the nasals
dorsally; supraoculars minute, touching frontal; latter small, separat-
ing the somewhat larger parietals; upper part of nasal reaching to
level of middle of eye, widened ventrally the nasal suture arising
from lower part of rostral, dividing the scale into two parts; preocu-
lar touching second and third supralabials, separated from the
supraoculars; part of rostral visible below, much wider than long;
ocular touching two labials, the area surrounding eye uniformly
light; four scales border it behind, between the fourth labial and
the parietal; mental not twice as broad as deep; two infralabials;
30 scalerows around neck; 28 around middle of body; 20 in front of
vent; ventral scales from mouth to vent 388; subcaudals, 8 + 1.

Color: Above gray-white to yellowish with top of head and an-
terior part of neck dark; neck and body blotched with irregular
black spots separated by irregular light yellow areas. All the black
scales have a grayish center and these form dim lines of gray
through the black but are interrupted by the yellowish blotches be-
tween the black areas. Those on the median rows are as distinct as
those on the lateral regions. The 15 to 17 ventral rows largely
yellowish, with smaller and fewer black blotches, and the light areas
in each scale larger.

Amphibians and Reptiles from Liberia 1205

"Color in life: Surface translucent so that the body is grayish
flesh. All pigmented areas are black. The spots on the scales, that
give a striated appearance, are gray."

Measurements in mm: Total length, 185; tail, 3.7; width, 3 cm.
from tip of snout, 7.6 width at middle 7.8; width into length, 237 —
25.6 times.

Remarks: The relationship of this form does not seem to be
with the other known local species. It differs in the peculiar colora-
tion and in having the nasal completely divided and the nasal
suture arising from the rostral. This latter character is known in
anchietae a form described from S. W. Africa. This, however, has
30-32 scales round the body, and is pale yellow with grayish brown
blotches. The nasal is semidivided and the preocular touches only
one labial. The length is 119 mm.

Another species that may be related is Typhlops anomalus also
described from Southwest Africa. This has 28 to 30 scales around
the body; however, this species is uniform brown above, the sides
and lower parts yellow. The nasal suture arises from the rostral but
the scale is only partially divided.

Typhlops manni Loveridge, Typhlops leucostictus Boulenger, and
Typhlops punctatus punctatus (Leach) also are known to occur in
Liberia.

It is possible that this form has been described and is to be found
among the synonyms of punctatus. We believe, however, that the
origin of the nasal suture is sufficiently significant to have been
recorded, if it were present in any of the forms described. The num-
ber of ventral scales is 34 more in punctatus from Harbel.

Typhlops punctatus punctatus (Leach)

Acontias punctatus Leach, in Bowditch, Mission, Ashantee, 1819, p. 493 (type

locality, Fantee).
Typhlops punctatus Boulenger, Catalogue of the Snakes in the British Museum,

vol. 1, 1893, pp. 42-43 (part.) ("total length 630 mm."); Schmidt, Bull.

Amer. Mus. Nat. Hist., vol. 49, 1923, pp. 45-46; Barbour and Loveridge, in

The African Republic of Liberia and the Belgian Congo, edited bv Richard

P. Strong; Cont. Dept. Trop. Med. Inst. Trop. Biol. Med., no. 5, vol. 2, 1930.

pp. 770-772.
Typhlops punctatus punctatus Loveridge; Proc. New England Zool. Club, vol.

17, May 20, 1938, p. 55 (Ganta, R. L.).
Onychocephalus Liheriemis Hallowell, Proc. Acad. Nat. Sci. Philadelphia,

1848, p. 59 (type locality, Liberia).

A single specimen, EHT-HMS No. 31890 5 has the following
characteristics: head much narrower than body, the snout with a
very obtuse horizontal keel; nostrils inferior not visible from above;
rostral large, the part visible from below much wider than long,

1206 The University Science Bulletin

the part visible above two thirds of the width of the head at eye,
its posterior edge reaching the front level of eye; latter can be
distinguished; nasal narrowed above, much widened below, semi-
divided, the suture arising from the first labial, scarcely reaching
beyond nostril; the preocular present much narrower than the nasal,
somewhat narrower or nearly equal to the ocular, in contact with
the second and third supralabial; supraoculars elongate, diagonal,
not covering any part of eye, separated from each other; prefrontal
three times as wide as long; parietals barely differentiated in size
from surrounding scales; frontal smaller than the scale following
it; four supralabials; mental twice as wide as long, with two infra-
labials on each side, concealed when mouth is closed.

The diameter of body at middle is contained in length 22 times;
diameter of neck (three centimeters behind head), in length, about
34 times; tail broader than long, the upper part rather sharply
deflected downward, the under part slightly turned upward, and
ending in a sharp low spine. There are 28 scalerows around body;
30 around neck a centimeter behind tip of snout; scales from mouth
to vent, (ventral count) 422; subcaudals, 11 -|- 1; vent bordered by
six scarcely differentiated preanal scales.

Measurements in mm.: Total length, 543; tail length, 10; width
of body near middle, 25; three cm. behind tip of snout, 16; width
of tail at vent, 15; width of head at back of ocular, 11.

Color: Above generally blackish brown, each scale largely trans-
parent, and showing a dim whitish transverse mark near its middle.
These light areas become larger on the sides, some being contiguous
and tending to form light lines. The dark brown color forms rows
of dots on upper part of sides but lower they become fewer, and
the four to six ventral scalerows are uniformly yellowish save for
an occasional brown dot. The general impression is a brown snake
with ten or twelve lighter lines very dim medianly, but becoming
well defined laterally.

Remarks: The stomach contents consisted of recently ingested
ant eggs with occasional ants. The mouth contained two ant heads
with the mandibles fastened to the mouth tissues.

This species reaches a length of about 800 mm., and is probably
the largest species of the genus Typhlops.

Loveridge (1938) reports a specimen measuring 795 mm. long,
4 inches in circumference and one and one-fourth inches in di-
ameter, from Ganta, Liberia.

Amphibians and Reptiles from Liberia 1207

Calabaria reinhardti Schlegel

Ertjx reinhardtii Schlegel, Bijdr. tot Dierk., vol. 1, 1848, p. 2, pi. (type locality,
Gold Coast).

Calabaria fusca Gray, Proc. Zool. Soc. London, 1858, p. 154, pi. 14 (type
locality, Old Calabar, Southern Nigeria, W. Africa).

Calabaria reinlmrd:i Boulenger, Catalogue of the snakes in the British Museum,
vol. 1, 1893, p. 92, ("Liberia to the Congo"); Proc. Zool. Soc. London,
1919, pts. 3-4, Feb. 1920, p. 274; Stull, Proc. Boston Soc. Nat. His., vol. 40,
no. 8, p. 396; Barbour and Loveridge, Cont. Dept. Trop. Med. Inst. Trop.
Biol. Med., no. 5, 1930, p. 772 (Paiata, St. Paul's River, Liberia).

Two specimens of this burrowing boid serpent, EHT-HMS Nos.
31887 £ —31888 $ from Harbel, Liberia, are at hand. The charac-
ters of No. 31887 follow: the rostral is about as wide as high the
part visible above about as long as wide; this scale followed by three
paired scales wider than long, the first two in contact mesially, the
third separated by a small scale; two supraoculars, frontal and
parietals considerably broken; nostril in a single nasal, not or only
partially divided, followed by a rather large loreal (a small scale
segmented from the third median pair forms a second superim-
posed "loreal" on one side); one preocular, two postoculars (one
only on right side); two or three anterior temporals; supralabials,
7-8, the third and fourth bordering orbit; infralabials, 9-10, the
first three touch the small anterior chinshields, the first pair in
contact mesially; second chinshields not distinguishable from the
chin scales, of which there are seven rows between the chinshields
and the first ventral. Scales smooth, with formula, 31 head, 25 neck,
29 middle, 23 (24) anterior to vent; ventrals 221, anal single, sub-
caudals 1 pair, -f- 19 single -f 1 single terminal. The tail is blunt.

Color: Above lavender brown with many scattered lighter scales
or groups of scales; entire head and first part of neck blackish; upper
part of latter half of tail black; venter largely cream-white with
some small black marks.

Measurements in mm., of Nos. 31887 and 31888 respectively:
total length, 822, 728; tail, 62, 56; head width, 17, 14; width of neck,
19, 15; greatest width of body, 30, 24; width of tail ( at 15 mm. from
tip), 21, 16.

Remarks: "Very common here, usually seen crossing road of the
house areas at night or in the early morning especially at the be-
ginning of the dry season. One was found dead on the cement
air-strip curled in a typical manner and only recently dead. Since it
appeared uninjured we surmised it was unable to burrow into the
cement and the hot sun had killed it.

1208 The University Science Bulletin

"We kept one alive for a while. It was very gentle and my
young daughter often carried it about. It always curled in a ball
with the head carefully tucked in the center, but the tail was left
poking out as though it was the head, and could fool anybody.
When it was put down it remained motionless for ten minutes or so,
then would cautiously uncurl and very slowly move away to escape.
Obviously it was incapable of fast movement. The muscles are
so strong that it was very difficult to unroll it by force. It never
hissed, threatened, or thrashed about.

"There are two color phases in life represented by the two speci-
mens. ( 1 ) Dorsal color dark brown with individual scales or small
groups of scales medium brown or tan; ventral surface with cream
mottling, spotted with medium brown. (2) dark brownish purple,
the markings orange or orange shaded purple. Particularly toward
the posterior part, the orange is very bright and striking both dor-
sally and ventrally.

"The amount of light color on the tail varies a great deal in the
species, many specimens I have seen showing none, others much."

If the preserved specimens are removed from alcohol into the
air for as short a period as five minutes, the entire ventral surface
changes to a uniform paper-white color and certain of the scales
adjoining them also assume this color!

The limb remnant is terminated by a sharp protruding claw.

Variation: The smaller specimen has 226 ventrals, an undivided
anal, and the subcaudals as follows : one divided pair followed by 24
unpaired scales.

There is a small intercalated scale between the second and third
infralabials bordering lip, that is absent in the smaller specimen.
The posterior infralabials in both appear to be segmented from
the adjoining scales. The smaller female specimen has two post-
oculars on each side, and has no trace of the limb externally.

Neusterophis variegatus (Peters)

Mizodon variegatus Monatsb. Akad. Wiss. Berlin, 1861, p. 358. (type locality,

Pel, Gold Coast, Africa)
Neusterophis variegatus Loveridge, Proc. U. S. Nat. Mus. vol. 91, no. 3128,

1941, p. 120. (Gibi, Bromley, Bendaja, in Liberia)

A single specimen, EHT-HMS No. 31918 has the following
characters all included in the known variation: length of eye
equal to distance between eye and tip of snout; scalerows, 15, 15, 15;
ventrals, 130, subcaudals (divided), 76; anal divided; scales smooth,
lacking apical pits; temporals 1 + 2; preoculars, 2-1; postoculars,

Amphibians and Reptiles erom Liberia 1209

3, supralabials, 8, the fourth and fifth bordering orbit; nasal di-
vided; loreal higher than wide; infralabials, 8-9, the last small,
four touching the anterior chinshields, which are much shorter than
the second pair. There are 24 maxillary teeth (counting missing
teeth).

On middle scalerows dark brown, each scale with indefinite
lighter flecks; on each side a row of whitish dots, none as large
as a single scale, each separated longitudinally from nearest dot by
about two scale-lengths; lower scalerows marked like those on
dorsum; median half of venter yellow with the outer fourth of
each ventral bearing a regular quadrangular dark spot, usually a
little darker than body color; median subcaudal area similar to
median ventral area; labials yellowish, the sutures edged with
black.

Natrix anoscopus anoscopus Cope

Tropidonotus anoscopus Cope, Proc. Acad. Nat. Sci. Philadelphia 1861, p. 299;

(type locality, Cuba, ex errore).
Natrix anoscopus Cope, Proc. U. S. Nat. Mus., vol. 14, 1892, p. 673.
Natrix aniscopus aniscopus Loveridge, Proc. U. S. Nat. Mus., 1941, pp. 118-

110 (Gibi, Bendaja, Harbel, Paiata, R. L.).
Tropidonotus ferox Giinther, Ann. Mag. Nat. Hist., ser. 3, vol. 12, p. 355, pi. 6,

fig. F (Fernando Po. ).
Helicops gendrii Boulenger, Ann. Mag. Nat. Hist., ser. 3, vol. — , 1910, p. 512

(French Guinea).
Natrix aniscopus gendrii Loveridge, Proc. U. S. Nat. Mus., 1941, pp. 118-119.

Loveridge has recognized two subspecies, aniscopus aniscopus
and aniscopus gendrii but suggests that the form on Fernando Po
described as ferox could not be given subspecific rank. In the col-
lection at hand there is a Natrix that agrees in very considerable
detail with Gunther's figure of ferox but differs from that form in
having the lateral markings absent, the color being a dull grayish-
brown, without lateral black spots. There are no lines of spots
on the vemer which is orange-tan, each scale edged anteriorly with
a blackish line.

It would appear probable that this is a subspecific form of ferox
[= anoscopus] but it must be very doubtful that it represents the
typical form anoscopus anoscopus, a species having, presumably, an
erroneous type locality "Cuba."

The specimen at hand is EHT-HMS No. 31893 $ , from Harbel,
Liberia. It has the following characteristics: the part of rostral
visible above more than a narrow transverse line; a pair of small
internasals only minutely longer than wide; prefrontals wider than
long, a little shorter than internasals; frontal short, scarcely longer
than wide, slightly less than its distance to tip of snout, distinctly

1210 , The University Science Bulletin

shorter than parietals; nostril in a single nasal, an entrant suture
from first labial partially divides the scale but does not reach nostril;
loreal longer than high; two preoculars, two postoculars, three or
four suboculars; supraoculars slender; temporals, 1 + 2 + 4; supra-
labials, 9-9, none entering orbit; infralabials, 10-10, five bordering
first pair of chinshields, which are smaller, but equally as long as
the posterior pair. Scale formula, head 35, 23, 25, 25, 20; all scales
strongly keeled with apical pits; ventrals, 141; anal, divided; sub-
caudals, 68. The ventral preceding the anal pair is divided; a
small intercalated scale also present between this and the anal pair.

Color: "In life entire dorsal surface dull grayish-brown; no black
lateral markings; there are a few small lateroventral black flecks
low on the side near ventrals; ventral color orange-tan each scale
bordered anteriorly with a dark transverse line; subcaudals gray."

The top of the head is uniform olive to gray brown, the edges of
the upper labials being somewhat darker; infralabials and the scales
and shields on chin and throat edged with gray; occasionally there
may be a tiny black border on outer edge of ventrals.

Measurements in mm: Total length, 524; tail, 131; width of head,
14; length of head, 20; narrowest neck width, 11.4; length of frontal,
4.5; width of frontal, 3.5.

Remarks: This group of Natrix has a deep groove between the
labials and temporals, which is more or less continued between
labials and the suboculars and loreal. In the groove one finds a
narrow expanse of gray skin when the labials are pulled down;
when seen from above the posterior upper part of labials form a
tiny shelf.

"The specimen was caught while we were tearing down a fish
pond. It was taken with a two and one-half inch Tilapa (a pond
fish) in its mouth."

Natrix firestonei sp. nov.

Type: EHT-HMS No. 31894 Harbel, R. Liberia, Mrs. Dora Weyer
collector, 1956.

Description of type: Rostral twice as broad as high, visible above
as a line; internasals narrowed anteriorly, much longer than wide;
prefrontals shorter and wider than the internasals; nostril in a single
nasal, with a small suture entering the scale from the first labial but
not reaching nostril; loreal longer than high; frontal longer than
wide, longer than its distance from tip of snout, shorter than the
parietals, one preocular not reaching the frontal; three or four sub-
oculars separating all labials from orbit; two postoculars; temporals,

Amphibians and Reptiles from Liberia 1211

1 -)- 2 -j- 3, the anterior elongate, others scalelike; supralabials, 9,
separated from temporals by a noticable groove; infralabials, 10-9,
five scales touching the elongate anterior chinshields which are
slightly longer than the second chinshields; latter scales separated
by small scales; two pairs of scales between chinshields and first
ventral.

Scales sharply keeled, those on outer row are smooth; the scale
formula, 28 (head), 23, 23, 20, (the median row lost one centi-
meter anterior to vent). Apical pits paired; ventrals, 143, the last
one separated from the paired anal by two scales not in contact
mesially; subcaudals, 67.

Color in life: Dark brownish-gray dorsally; five rows of black
spots, the median series alternating (usually) with paired lateral
spots that are a little higher than wide; below these on the border
of the ventrals another, more numerous series of smaller spots
varying in size. Head generally dark grayish to blackish; grayish
below with parallel lines of dark brown spots tending to form two
more or less continuous lines near the outer part of ventrals on the
anterior two thirds of body, the area between them gray; the edges
of the ventrals with some brown along their anterior edges; spotting
on tail much less distinct; dark gray in subcaudal region. Scales
on chin and throat more or less outlined in brown or black.

Head relatively slender not strongly differentiated from neck.

Measurements in mm: Total length, 472; tail, 113; width of head,
10; width of neck, 8.2; head length 18.4; length of frontal 4.25; width
of frontal 3.0.

Remarks: Body triangular in cross-section with a rather distinct
median dorsal ridge. The specimen is a female with seven eggs
showing no trace of embryos. This specimen was captured some
distance away from the water. The paired apical pits are relatively
large and distinct over most of body and tail. There are 23 maxil-
lary teeth, the posterior ones only a little stronger and larger than
the anterior, with no diastema present.

Since this and the preceding snake occur in the same locality,
it does not seem fitting to regard one a subspecies of the other.
The scale formula is similar, the chief differences being in the
different shape of the head, which is much wider in anoscopus.
There is a distinct difference in the color pattern; and the somewhat
elevated median line and somewhat triangular body is seemingly
very different from the rounded body of aniscopus; the propor-
tionally larger temporals and parietals are significant also.

1212 The University Science Bulletin

"This specimen was not found in or near water. The snake is
thinner, the head longer and narrower and the snout more rounded
than in our specimen of N. aniscopus aniscopus."

Bothrophthalmus lineatus lineatus Peters

Bothrophthalmus lineatus Schlegel, in Liechtenstein and von Martens, Nomen-
clator Reptilium et Amphibiorum Musei Zoologici Berolinensis, 1856, p. 27
(nomen nudum; type locality, Goldkiiste); Barbour and Loveridge, The
African Republic of Liberia and the Belgian Congo. Cont. Dept. Trop.
Med. and the Inst. Trop. Biol, and Med. no. 5, 1930, p. 784. ( Cambridge,
Harvard Univ. Press).

Elaphis ( Bothra})hthalmus) lineatus Peters, Monatsb. Akad. Wiss. Berlin, 1863,
p. 287 (type locality, "Guinea").

Bothrophthalmus lineatus lineatus Loveridge, Proc. New England Zool. Club,
vol. 17, May 20, 1938, p. 56 (Ganta, Republic Liberia).

This brilliantly marked serpent is represented by a single speci-
men EHT-HMS No. 31891 $ . It has the following scale and
other characters: rostral very narrowly visible above; internasals
small, somewhat elliptic or leaf-shaped; about one third of size of
prefrontals; frontal longer than wide, equal to its distance from tip
of snout, as long as the parietals; inner part of the supraocular fused
to the parietals; nasal divided; a longitudinal trenchlike depres-
sion, having a sharp canthal edge above in loreal region is con-
tinued behind lower posterior part of eye for a short distance; loreal
large elongate, somewhat rectangular on left side, vertically divided
on right side; a preocular, the upper part of which grows forward
above posterior half of loreal;* two postoculars; two anterior tem-
porals, only one, however, in contact with postoculars; two sec-
ondary temporals; seven supralabials the fourth and fifth border
orbit, the upper edge of the fourth bent so as to resemble a lower
preocular; seven infralabials, four touching first pair of chinshields,
which are larger than the second pair; scale formula, 27 (head);
23, 21, 20. Ventrals, 194; subcaudals, 71; anal single; all scales
keeled, the posterior dorsal keels forming continuous ridges.

Color: Body striped in black and red. There are four black
stripes, the two median wider than the intervening red stripes,
the outer stripes of about width equal to the adjoining red stripes;
on the third scalerow there is a series of dashlike marks forming a
more or less continuous line; the head has five lines, two through
eye, a pair of lines passing above the eyes and a short frontal-
occipital line. All stop and are separated from the body lines by a
short diastema; a few dim dark spots on infralabials and on neck;
some pigment present on the posterior subcaudals.

* This must be examined carefully since the anterior part resembles a second ' loreal
and may be mistaken for a separate scale.

Amphibians and Reptiles from Liberia 1213

"A beautiful animal! I saw it immediately after death — the
lateral and dorsal areas between the back stripes were bright
"orangish" brick-red, the dorsal head color darker. The ventral
surface was a lovely iridescent vermillion-pink, the tip of the tail
a darker shade. The head ventrally was white with scattered
orange spots."

Remarks: "This snake though harmless is held by many natives
to be deadly poisonous and by others a snake that foretells death.
All regard it as 'bad-bad' and wanted us to have nothing to do
with it. It apparently holds a place in their superstitions much
like that of the chamaeleon. Dr. Harley in his book, 'Practice of
Native Medicine' describes the special medicines used for the
'red and black snake' by which I think he means B. lineotus lin-
eatus."

The specimen is a female containing eight ovarian eggs.

Boaedon lineotus virgatns Hallowell

Coelopeltis virgata Hallowell, Proc. Acad. Nat. Soc. Philadelphia, (vol. VII),

1854, p. 98 (type locality, Liberia).
Boaedon virgatus Boulenger, Catalogue of the Snakes in the British Museum

(Natural History), vol. 1, 1893, p. 331. (spelled Boodon) (synomymy);

Bogert Bull. Amer. Mus. Nat. Hist., vol. 77, 1940, p. 24, (Ganta, Liberia).

Three specimens EHT-HMS Nos. 31913-31915, all females, yield
the following data respectively: ventrals,213, 211, 218; subcaudals
47, 47, 45; anal single; scalerows, 21, 23, 17; 21, 23, 17; 21, 21, 17;
preoculars, 2-2, 2-2, 1-1; postoculars 2-2; preocular touches frontal,
no, yes, yes; supralabials, 8-8 in all; supralabials border orbit, 4th,
5th, in all; infralabials, 9-9, 8-8, 8-8; total length in mm., 806, 350,
408; tail, 104, 43, 45. In all there are two lines on the side of the
head, and a light mark on the rostral extending down across chin.
The venter is lavender-gray with a line of cream in the middle.
The pitting on the scales is paired throughout save an occasional
scale on the outer row with a single "apical pit." No. 31913 contains
six eggs.

"The color of the entire dorsal surface is glossy black. There are
two sets of yellow lines on the side of head; the lateral surface of
the head between the yellow lines is dark gray. Ventrally, the head
is grayish white with a pinkish wash. This narrows at the neck
into a cream-yellow stripe running the length of the body and
widening to include the entire anal scale, and making a prominent
cream-yellow spot.

"The remainder of the ventral surface and the subcaudal area is
dark glossy gray. A second adult specimen which I have here for

1214 The University Science Bulletin

the hospital collection has the ventral stripe peach-yellow and the
immature which I send had, in life, a pink-orange ventral stripe
heavily shaded with black in the posterior half of the body. All
were black dorsally, not brown with a gray head as it is sometimes
described."

Boaedon lineatus lineatus Dumeril, Bibron, and Dumeril

Boaedon lineatus Dumeril, Bibron, and Dumeril, Erpetologie generale, vol. 7,
pp. 363-364 (type locality, Gold Coast).

We cannot regard this species as subspecifically related to
Boaedon virgatus Hallowell. The two occur together and are sep-
arable on the basis of several characters. For example, markings,
ventral coloration, number of dorsal scalerows, and the character
of the apical pits. One specimen, EHT-HMS No. 31916 5 in the
collection contains a rodent, Rattus rattns * with a total length of
352 mm.

The following characters obtain: rostral narrowly visible above;
internasals subtriangular, about as wide as long, fused together
anteriorly for a short distance; prefrontals about as long as wide;
frontal subtriangular about four-fifths times as long as wide, touch-
ing the preocular, slightly longer than its distance to snout tip,
shorter than the parietals but equal to its distance from their pos-
terior tip; nasal completely divided, the anterior part much wider
than posterior, rather narrowly separated from its fellow from the
opposite side; loreal more than twice as long as high; one large
preocular touching two infralabials; two postoculars, only the lower
touching anterior temporal; temporals, 1 + 2 + 3; supralabials,
8-8, the fourth and fifth bordering orbit; infralabials, 9-9, four
touching the first chinshields, which are larger than second pair;
no scales between either pair of chinshields; eye rather small, pupil
vertical; scalerows, 33 about head, 27 neck, 29 middle of body, 21
anterior to vent; scales of the median rows with single apical pits;
scales of the five outer rows with double terminal pits, except outer-
most, which also have a single pit. Ventrals, 220; subcaudals, 49
(divided); anal single. Maxillary teeth 18, the fourth and fifth
largest, diminishing posteriorly; no diastema.

Measurements in mm.: Total length, 851; tail length, 102; length
of head, 29; width of head, 21.

Color markings: Above lavender to violet with a cream line from

* Identified by Mr. Philip W. Ogilvie.

Amphibians and Reptiles from Liberia 1215

rostral through the upper part of eye to lower tertiary temporal;
a second line, beginning on first supralabial, to eye, and from eye
to last supralabial; the anterior part of the line dim; an arched
light line on rostral, the two ends directed downward across mental
and first infralabial; a light line begins opposits the fourth ventral
and extends to level of the 57th ventral; chin and throat grayish;
venter and subcaudal region white, save for a faint pigmentation
on extreme outer edges of the ventrals.

Bogert (1940) has recognized several subspecific forms in this
species. He regards the typical form, (type locality Gold Goast)
with two lateral white lines as occurring in Tanganyika, Nyasaland,
Belgian Congo, Natal, and Angola. This large series might profitably
be examined for apical scale-pit characters. Since Bogert does not
comment on the color and markings of this series one presumes
that they tend to agree with the type.

My specimen does not agree with the type in color markings so
far as concerns the two lateral body stripes.

Chlorophis hetero dermis (Hallowell)

Chlorophis heterodermis Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1857,
p. 54 (type locality, Gaboon).

Three specimens of this snake are at hand: EHT-HMS Nos.
31920, 31921, 31922, all females. The following characters obtain
in the three specimens respectively: Total length, in mm. 670, 541,
527; tail, 172, 142, 138; ventrals, 152, 149, 149; subcaudals, 81, 79, 79;
anal single in all; supralabials, 9-9, the 4th, 5th, 6th, entering orbit
in all; infralabials, 9-9, 9-9, 10-0, the first five touching the first pair
of chinshields.

The scale formula is 15, 15, 11 in all. The scales are smooth but
in No. 31920 pits are present on the anterior part of the neck. Some
scales have 3 apical pits, others two, others one only or none. The
apical pits on No. 31921 are more pronounced and extend back on
neck for several inches.

In 31922 the epidermis of the scales is lost and apical pits are
dim or absent. No. 31920 is uniformly black above in preservative,
the side of the snout and a postorbital labial spot, white; chin and
anterior part of throat yellowish to ivory-white, the yellow and
black interdigitating to some extent on the sides of the neck. The
ventrals are keeled laterally but the subcaudals lack keels. The
maxillary teeth are 35-34 in No. 31920.

1216 The University Science Bulletin

MeheJya poensis (A. Smith)

Heterolepis poensis A. Smith, 111. Zool. South African Rept, 1847, footnote to
pi. LV (type locality, Fernando Po).

The single specimen EHT-HMS No. 318S4 $ has the following
characters: rostral visible above as a narrow band; internasals
wider than long; prefrontals twice area of the internasals, longer
than wide, touching loreal laterally; frontal small, subtriangular,
the front edge curving somewhat, distinctly shorter than its distance
to the end of the snout, much shorter than the parietals, double the
width of the supraoculars; parietals angulate posteriorly; nostril
between two nasals; loreal longer than high; one preocular, two
postoculars; temporals, 1 + 2 + 3; supralabials, 8-8, the third and
fourth bordering orbit; infralabials, 8-8, the first five touching the
first chinshields, which are distinctly larger than second pair; median
scalerow enlarged, bicarinate, the scales somewhat hexagonal; other
scales with single keels; scale formula, 19 (head), 15, 15, 15; ven-
trals, 240; tail broken, partly missing; anal single.

Color in preservative: Uniform lavender brown above on sides
and on outer edges of ventrals. Venter and subcaudal region uni-
form white; edges of supralabials and infralabials white.

Approximate total length, 857 (tail broken); snout to vent, 691.

Boiga blandingii (Hallowell)

Dipsas blandingii Hallowell, Proc. Acad. Nat Sci. Philadelphia, 1844, p. 170;
idem, ibid, 1854, p. 100 (type locality Liberia).

Dipsadomorphus blandingii Boulenger, Catalogue of the Snakes in the British
Museum (Natural History) vol. Ill, 1896, pp. 77-78 (synonymy).

Boiga blandingi Cope, Proc. Acad. Nat. Sci. Philadelphia 1860, 264; Barbour
and Loveridge, in Contributions from the Department of Tropical Medicine
and the Institute for Tropical Biology and Medicine No. V. Harvard Col-
lege Press, Cambridge, 1930, p. 773 (Paiata, St. Paul's River); Loveridge.
Proc. U. S. Nat. Mus.; vol. 91, p. 122. (Bromley); Proc. New England
Zool. Club, vol. 17, May 20, 1938, p. 59 (Ganta, R. L.).

A large specimen EHT-HMS No. 31886 $ Harbel, Liberia, has
the following scale characters: rostral scarcely visible above, wider
than high; internasals little more than half the area of the pre-
frontals; frontal short, as wide as long, equalling its distance from
snout tip, somewhat shorter than the parietals; eye large, prominent;
nasal divided, the posterior part the larger with a depression; a
loreal; two preoculars, the upper not touching the frontal; two post-
oculars; two anterior and two secondary temporals; supralabials
nine, the fourth, fifth, and sixth bordering orbit; the seventh and
eighth touching the lower postocular; infralabials, 13-13, four or
five touching first chinshields, which are nearly same size as second

Amphibians and Reptiles from Liberia 1217

pair; a pair of scales between second chinshields and first ventral.
Scale formula, 34 (head), 29 (neck), 23, 23, 17, 15, the median
row enlarged, more than double the size of scales in adjoining row;
ventrals, 266; anal divided; subcaudals, 125.

The tooth formula is 9 + 2 + 1; 10 + 2 + 1. The anterior teeth
increase from first to fourth then are nearly equal; the two fangs
stand one behind the other, their grooves deep and open the length
of the tooth; a third fang, after a short diastema shows traces of a
groove but is not open except at tip and base. The dental formula
taken from a head of another specimen EHT-HMS No. 31945, is,
10 + 2 + 1, 10 + 2 + 1. In this specimen there is a suggestion of
a groove but no apparent opening capable of conducting venom
can be discerned on the last fang.

Measurements in mm: Total length. 2357 +; tail, 525 +; snout
to vent, 183.

Color in life: "Dorsal ground color grayish tan, the diamond-
shaped markings darker tan, outlined in brown. A dark brown
collar one scale wide around neck. Dorsal head color a darker
grayer tan than body, the mottlings dark brown; supralabials edged
with some dark brown. Ventral surface of head and neck pale
grayish green; five inches behind head this merges into gray with
a greenish cast; at 16 inches behind head it becomes plain tan, con-
tinuing so to the tip of the tail."

Remarks: "This snake was found electrocuted at the hydroelectric
plant with a large black snake. It would appear that the latter
had been chasing it when they came into contact with the electric
wires. The natives all insisted that the black snake eats this
species."

The stomach contains three young birds, one subadult and two
nestlings of Spermestcs cucuUatus cucullatus*

Psammophis sibilans phillipsii (Hallowell)

Coluber Phillipsii Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1844, p. 169
(type locality, Liberia).

Psammaphis sibilans phillipsii Loveridge, Bull. Mus. Comp. Zool. Harvard
College, vol. 87, no. 1, Oct. 1940, pp. 41-44. (monographic review; bib-
liography )

This species is represented by two complete adult specimens
EHT-HMS 31940 5 31941 <j and two heads 31908-31917; also by
four hatchlings Nos. 31909-31912 two associated with the egg. We
have followed Loveridge (loc. cit.) in regarding phillipsii a sub-
species of sibilans. The specimens so far as can be determined

* Identified by Phillip Ogilvie.

1218 The University Science Bulletin

agree with the limits of variation given by Loveridge, all of the
complete specimens having a single anal plate.

There is a considerable difference in color patterns in the four
hatchlings. Three have the labials and many chin scales bearing
black spots. In No. 31912 the lips and chin lack all black spots,
the venter is darker, and the tail is lighter than in the other three
specimens. This one has a combined ventral-subcaudal count
that totals 283. This is considerably higher than the counts in the
young with complete tails, which are 264, 269.

"This snake is, next to Causas rhombeatus, the commonest snake
here by all means. It is seldom seen in the rainy season, but as
soon as the dry season gets under way in late December or January
they are found around every house or open grassy spot.

"They have most interesting breeding habits; all stories told me
by planters agreeing with the one nest I saw myself. In this last
case, Dr. Kessler phoned me to come to his house saying that the
boys had found a 'nest of snakes.' I went immediately and found
the boys of the yard crew waiting for me. They had, in hand,
obviously just killed, eight large snakes of this species, and they ( and
he) claimed that they had found them all entwined together in this
hollowed out spot in the lawn. The spot was about a foot and a
half in diameter when I saw it, about one foot deep, and so badly
cut by the boys' cutlasses I couldn't tell what it must have looked
like before they found it. However, they had 'squashed' beside
it a mess of snake eggs, how many I couldn't tell exactly. I counted,
uncrushed and still in the hole where Dr. Kessler had had them
leave the rest for me to see, 52 eggs. This was obviously a communal
nest. Because of the mess — you can imagine eight yard boys
killing eight big snakes with cutlasses — I did not examine the snakes
for sex, as I later wished I had. How many eggs might eventually
have been laid I don't know either. However, I have had several
planters tell me of finding similar nests on their own lawns —
always the estimate of the numbers of snakes present together is
between eight and ten. The largest head of this species, in the
specimens I sent you, incidentally, came from this 'nest.'

"As to the color, the 'greenness' of the back varies considerably in
individuals, so that sometimes one calls it a "brown snake' and
sometimes a 'green snake,' varying, in other words, from a dull
olive-brown to a bright olive-green, and sometimes a distinct
gray-green with no tint of olive. Ventrally the green is always
striking, sometimes a clear, light-green, sometimes bluish green,

Amphibians and Reptiles from Liberia 1219

sometimes, yellow-green. There is also eonsiderable variation in
the amount of orange. In all specimens there is some orange along
the side of the head and on the inferior labials. In one of the
specimens I sent you, you will see orange over the entire nose and
mouth area with only the chinshields and the gulars white, the
orange extending over the rostral, nasals, loreals and preoculars.
In some specimens this orange area of the jaws is spotted with
black, in others not. In some the orange continues down the first
ventral plates also, and in the one you have, it runs the length of
the body on the outer scalerow and above the upper (outer) edges
of the ventrals, the anal area and the ventral side of the tail being
(mite orange. In all specimens I have seen there is some yellow
or orange on the under part of the tail, in some the orange being
very bright everywhere but on the vertebral row dorsally. There
is also considerable variation as you see in your specimens on the
black markings dorsally."

Thelotornis kirtlandii kirtlandii (Hallowell)

Leptophis kirtlandii Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1844, p. 62,
idem, ibid, 1854, p. 100. (type locality, Liberia).

Thelotornis kirtlandii Barbour and Loveridge, Cont. Dept. Trop. Med. No. V,
vol. II, 1930, p. 773. (Gbanga, Liberia).

Thelotornis kirtlandii kirtlandii Loveridge, Bull. Mus. Comp. Zool. Harvard
College, vol. 95, no. 2, Dec. 1944, pp. 149-154 (Edina and Gbanga, Li-
beria) (literature listing).

A single specimen (EHT-HMS 31885 $ ) has the following
characters (due to an old head injury certain characters cannot be
determined): head wedge-shaped; rostral much broader than high;
eye large; internasals as long as wide, touching one loreal; pre-
frontals longer than internasals, bordering two loreals laterally;
nostril in a single elongate nasal; two loreals, the second twice the
length of first; one large preocular, not touching frontal; three post-
oculars, the lower elongate; temporals, 1 -f 2; a pair of postparietals
separated by a scale; supralabials, 8, the fourth and fifth bordering
orbit; infralabial, 9-10, the first four touch the first pair of chin-
shields, which are shorter than the second pair; scale formula, 19-19,
15, the scales weakly keeled, elongate. Ventrals 178; anal divided;
subcaudals 141.

Color: The head is uniform lavender (probably green in life)
the labial areas somewhat lighter; neck indefinitely barred with
grayish blotches growing less distinct posteriorly; most of body
light lavender with dark flecks indistinctly outlining spots.

1220 The University Science Bulletin

Aparallactus modestus Giinther

Elapops modestus Giinther, Ann. Mag. Nat. Hist., ser. 3, vol. 4, 1859, p. Ill,

pi. 4, fig. c (type locality "West Africa"); Zool. Record, 1864, p. 152.

(figured by Jan as Elapops petersii); Boulenger, Proc. Zool. Soc. London,

Feb. 20, 1920, pi. 293.
Pariapsis plumbeatra, Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, pp. 241-

242 (type locality, Liberia).
Elapops plumbeater Cope, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 566

(contrasts plumbeater with modestus).

Our specimen, EHT-HMS No. 31892 $ from Harbel, Liberia, has
the following characteristics:

Rostral well visible on dorsal part of snout, one and one-third
times wider than high; internasals small, less than half the area of
the prefrontals; frontal wider anteriorly, longer than wide, its length
greater than its distance from tip of snout, distinctly shorter than the
parietals; latter slender, pointed behind, fused together at tip;
nasal divided, the nostril chiefly in the very small anterior part; no
loreal; one preocular; two small postoculars; eye diameter less than
its distance from the lip; no anterior, but two posterior temporals, the
upper very large; supralabials seven, the third and fourth border
orbit; fifth much the largest, broadly in contact with the parietal;
infralabials six, three bordering the first chinshields, which are wider
and equally as long as the second pair; upper temporals separated
behind the parietals by three scales; two paired scales between
second chinshields and the first ventral.

Scales smooth, without pits, in 15 scalerows throughout neck and
body, the outer row a little larger than the others; ventrals, 158;
anal divided; subcaudals, 35 in a single row. (Tip of tail missing.)

Color in fixative: Dorsally and laterally bluish gray, each scale
with a more or less distinct dark border; head generally slightly
more ultramarine; supralabials a little lighter near edge of lip;
ventral surface dirty whitish, each scale with a line of grayish pig-
ment on its anterior edge, growing more pronounced posteriorly
where it may spread over the entire scale; subcaudals uniformly
grayish black.

Measurements in mm: Total length, 454; tail (incomplete), 58;
width of head, 9; width of body, 10; head length, 13.

Remarks: Although reported by Cope (1860) from Liberia under
the name Pariapsis plumbeatra (sic) it was not included in the
Barbour and Loveridge list (1930). It would appear that it is a
relatively rare snake, the greater part of its range being east and
south of Liberia.

Amphibians and Reptiles from Liberia 1221

The curiously pointed condition of the parietals is seemingly
caused by their fusion posteriorly with the scale normally partially
separating their posterior tips. The normal suture between the
parietals stops at the point of fusion.

Naja melanoleuca Hallowell

Naja hate melanoleuca Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1857, p. 61
( type locality Gaboon, West Africa ) .

Three specimens, two young, and the head and neck of a half
grown individual, are in the collection. These are EHT-HMS Nos.
31905, 31907, and 31906 respectively. The scale characters of the
young respectively are as follows: ventrals, 210/2, 214; subcaudals,
64, 68; scalerows, 23 (24) neck, 17, 13; 23, 17, 13, the scales smooth
without pits; supralabials, 7-7, 7-7, the fifth highest, the third and
fourth bordering orbit; infralabials, both 8-8; preocular, both 1-1;
postoculars, both 3-3. Total length in mm., 633, 432; tail, 106, 78.

The coloration of the young (No. 31905): above blackish the
top of the head a little less dark, the sides light grayish white or
yellowish white, the labial sutures more or less bordered with
black. At a distance of about 30 millimeters back of parietals and
again at about 65 and 120 mm. many of the black scales have small
yellowish dots near their anterior borders and the yellow of certain
ventral bands of yellow encroaches on the outer scale rows. The
chin and neck for about 57 millimeters is yellowish white; this
followed by a black bar, 21 mm. long (9 ventrals) then follows a
yellowish white band 25 mm. long with a single black ventral in its
middle; the following black band is 29 mm.; beyond this the scales
are gray black for some distance then become intense black to
the end of the tail. No. 31907 shows only two yellow areas on
venter separated by a black band 11 mm. wide. The sides of the
neck in both show a few small black spots. Tail banded with a
minute yellow band very near tip.

"This is a very common snake here, probably third in order in
frequency of occurrence. The young are speckled with white when
alive. I have seen four live ones in my garden at various times —
specimens between one and two and a half feet in length and they
all show this speckling noticeably, so that the first time I thought
it was a different species until it spread its hood. The adults when
alive show no white at all, but when preserved the scales separate a
bit and show a little. Incidentally, the young have always, in my

1222 The University Science Bulletin

experience, been more vicious than the adults, which usually try
to get away although they turn and spread their hood when at-
tacked. But the little ones usually turn and spread the hood when
first disturbed and do not take advantage of a chance to escape.
Also, they do not appear to be fast-moving snakes as many accounts
state. I personally found a large one in my chicken house and
had plenty of time to dodge his attacks and when he slipped through
the fence, I was able to run around the enclosing fence and catch
up and kill him on a downhill run to the rubber trees. He had
not eaten and was not sluggish on that account. In motion they
remind me of the common pilot black snake I grew up with in
Ohio. Incidentally, they love eggs, and are often found around
chicken houses, I presume mainly hunting rats, but certainly several
here to my knowledge have been killed when they had been found
asleep, usually under a setting duck, with three or four eggs inside."

Atractaspis corpulenta corpulenta (Hallowell)

Brachycranion corpulentum Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1854,

pp. 99 (type locality, Liberia).
Atractaspis corpulentus Hallowell, Proc. Acad. Nat. Sci. Philadelphia, 1857,

p. 70 (correction of type locality); Boulenger, Catalogue of the Snakes in

the British Museum (Natural History), vol. 3, 1896, p. 514.
Atractaspis corpulenta corpulenta Laurent, Mem. Inst. Roy. Sci. Nat. Belgique,

ser. 2, fasc. 38, 1950, pp. 40-42 (synonymy).

Description (EHT-HMS No. 31889^ Harbel, Liberia): Rostral
large, part visible above wider than long, the length equal to its
distance from frontal, partially separating the internasals; latter a
little wider than their greatest length; prefrontals shorter but wider
than internasals, their common suture slightly greater than that
between internasals; frontal wider than long, equal in length to its
distance from tip of snout, a little shorter than parietals; supra-
oculars very small; nasal divided, the posterior scale more than
double the size of the anterior; no loreal; a very small preocular
and a small postocular; eye small, separated from the mouth by a
distance equal to a little more than three times its diameter; a very
large anterior temporal; three secondary temporals all less than
half the size of anterior; five supralabials, first small, the third and
fourth border orbit; a small mental; first infralabials in contact
behind mental, but separated by the first chinshields, which border
lip, from the second infralabials; five infralabials; second chinshields
small separated by three rows of scales; the second infralabial very
large; seven scales between first chinshields and first ventral. Snout
wedge-shaped, with a somewhat depressed transverse edge. Scale

Amphibians and Reptiles from Liberia 1223

formula: 28 (head), 23, 21, 17; ventrals, 190, subcaudals, 26-27,
part double, part single as follows, 1 paired -f- 7 single -f- 2 paired
+ 1 single -f- 4 double + 2 single -f- 9 paired -f- 1 terminal (single),
(the last paired scales separated). The terminal subcaudal is a
short conical spine; anal single; scale smooth without apical pits.
A row of median widened scales occupying the dorsal tail surface,
about 12 in all.

A pair of functional fangs on maxillary; other teeth greatly re-
duced.

Color: Blackish brown above, slightly lighter on sides and venter;
posterior part of tail white, the color extending more than half its
length on ventral part, less than half on the dorsum.

Measurements in mm.: Total length, 619; tail, 58; width of head,
20; width of body, 21; head length to back edge of parietal, 16.

Remarks: "I have so far found or been given a total of three
Atractaspis corpulenta, whereas, so far at least, none of the other
Atractaspis species have been seen. My collectors tell me that
another kind, similar to these but different, can be found up-country
near Salala. That region has a much drier climate and more sandy
soil.

"I came across a live specimen lying injured in a ditch beside the
road early in the morning (apparently it had been hit a glancing
blow by a passing car) . . . and observed how it behaved.
It flattened its body making itself appear to be a much heavier snake
than it was, and added to this effect by inflating the neck region
considerably. Then the small head was curved down while the
neck was reared up, giving it a very vicious appearance for so
small a snake.

"All of the specimens are dull black or grey-black. Leeson's book
states that they are a slaty blue."

The type description of Br achy cr anion corpnlentum by Hallowell
shows several differences from the generally accepted idea of this
species. For example, the following are recorded for the type:
scale rows about body 25; internasals and prefrontals united into one
pair of scales; subcaudals single; there are but four supralabials.
Laurent (1950) in his revision has not commented on this discrep-
ancy. I am not aware of this condition occurring in any other
specimen (Boulenger 1906 only suggested it might be an anomaly).
It seems unlikely that there was a mistake in the count of the
number of supralabials. In our specimen the tail is only partially
colored white or cream, not "entirely white." Hallowell mentions

7—8050

1224 The University Science Bulletin

no white on the tail of the type! If these are not anomalies one
suspects the possibility that the specimens referred to this name
by authors subsequent to Boulenger, have applied the name to a
snake different from the type of corptdentum. Our specimen agrees
in most characters with the form redescribed by Boulenger. The
second labials are fused to the chinshields leaving the combined
scale bordering the lip. There are five supralabials but there are
23 scalerows such as one expects on leucura.

Catisus rhombeatus ( Lichtenstein )

Sepedon rhombeatus Lichtenstein, Verz. Doubl. Mus. Berlin, 1823, p. 106

(type locality, Gold Coast)
Causus rhombeatus Boulenger, Proc. Zool. Soc. London, 1919, pts. 3 & 4,

(Feb. 1920), p. 295, text fig. 1; Barbour and Loveridge, in Strong, Cont.

Dept. Trop. Med. Inst. Trop. Biol. Med., No. 5, vol. 2, 1930, p. 774.

There are seven specimens in the collection: EHT-HMS Nos.
31881-31883; 31901-31904. The last four numbers are recently
hatched individuals or (one) still in the egg.

The following data are taken from EHT-HMS No. 31881: Nostril
wider than high, the part visible above forming a sharp angle
separating the nasals and partially wedged between the internasals;
internasals longer than the prefrontals, with a narrow posterior
section that extends behind nasal, and is in contact with the loreal;
prefrontals much wider than long; frontal subhexagonal, the sides
parallel, longer than its distance to the tip of snout, longer than
the parietals; nasal scale with a transverse arm separated from its
fellow by the rostral, the large nostril pierced in the larger posterior
part; the internasal forms a slight overhanging shelf above edge
of nasal; loreal somewhat triangular; a single preocular (on one
side partially sutured); supraoculars nearly as wide anteriorly
as posteriorly; one postocular in contact with an elongate post-
subocular, this scale with two other suboculars form a series that
separates the labials from the orbit; supralabials, 6-6; infralabials,
9-9, four touching first chinshields; the second pair of chinshields
not differentiated. Scales of the eleven median rows with a well-
defined keel that does not extend the whole length of the scale;
scale rows, 19-19-13: temporals, 2 + 3; ventrals, 125; anal single;
subcaudals, 22, the last three undivided.

A large triangular or chevron-shaped black mark reaching to an-
terior edge of frontal; a dark indefinite line from eye to jaw angle;
on neck there are chevron-shaped spots, white throughout most of
body the median spots larger, somewhat rhomboidal with a lateral
indefinite series of spots on the outer scalerows; whitish on chin and

Amphibians and Reptiles from Liberia 1225

throat; most of venter lavender gray-brown the pigment forming
transverse markings on each ventral; area in front of vent much
lighter than elsewhere on venter but with some dark flecks; sub-
caudal area immaculate; tail above with a median black line.

Data on Causus rhomheatus Lichtenstein

Number

Sex

Ventrals

Sub-
caudals

Total
length

Tail

Scale formula

31881
31882
31883

9
$

S

125
125
127

22

22
23

460
397
405

42
41

38

19,
19,
19,

19(18), 12(13)
19, 11
18(19), 13

Variation: The subcaudals for the most part are divided but No.
31881 has three, 31882 four and 31883 five of the terminal sub-
caudals undivided. The suboculars two or three. The infralabials
touching the first chinshields are either four or five.

Remarks: Boulenger gives the range of ventrals as 120-155. Li-
berian specimens have a low number (125-127) while the highest
counts occur much farther to the southeast.

"This is the commonest snake here on the plantation. It is called
the 'yard snake' by the house servants, because they are so often
found under the houses or in the yard close by. The men engaged
in clearing operations report them as numerous in the low cut-over
bush of old native farms. The engineer clearing a site for a new
servants' camp behind the hospital compound complained one noon
that he had killed eleven of them that morning.

"I have read that this species is the greatest cause of death by
snake bite in the Nairobi area, but certainly here, although common,
it does not cause as many deaths as Naja melanolenca, or more
especially, Bitis gabonica. It does not behave viciously, and the
white children playing in the yards often find them and call the
servants to kill them."

Bitis gabonica Dumeril, Bibron and Dumeril

Echidna gabonica Dumeril, Bibron and Dumeril, Erpetologie generale, vol. 7,
pp. 1428-1430, pi. 80 bis (type locality, Cote du Gabon).

Heads from two large specimens (EHT-HMS Nos. 31899-31900)
and two smaller complete specimens are from Harbel, Liberia.
The two latter yield the following data respectively: Ventrals, 129,
131; subcaudals, 32, 28; anal single; supralabials, 14 (15), 14; scale-
rows at middle of body, 36, 38; total length in mm., 395, 748; tail
length, 36, 46.

"This is the snake that everyone here fears — an insolent vicious
animal. They are common, four medium to large ones have been

1226 The University Science Bulletin

found on the lawn at our house since we came here two years ago.
Those of us with small children have to think of them constantly.
They are beautifully colored yet seem to be easily overlooked on
a green lawn. I send a color photograph of a specimen just killed
on my driveway, but unfortunately the colors appear mostly tans
and browns. In life, the reddish tan markings of preserved speci-
mens, often, though not always, have a purplish-rose cast, whereas
the areas that are gray in preservative are a rich powder-blue. The
tan flush under the jaw is blood-colored in life, in fact on my first
specimen seen, I thought the jaw was smeared with blood.

"Our soil here is laterite — a reddish soil and the young specimen
I send was just that color. The varied markings of course were
present but the general coloration was that of the soil, and since
the soil is gravelly the markings correspond well to the shadows
caused by the small gravel, and the snake was difficult to discern."

Bitis nasicornis (Shaw)

Coluber nasicornis Shaw, Nat. Misc. vol. 3, pi. 94 (type locality, "interior of
Africa" ) .

Two specimens, EHT-HMS Nos. 31896 $ - 31897 $ yield the fol-
lowing data respectively: Ventrals, 122, 127/2; subcaudals, 27, 28;
anals, single; supralabials, 15 (16), 15 (16); scalerows at middle of
body, 35, 33; total length in mm., 530, 676; tail, 71-94.

"This species is also common along and in the waterways, but
although I have been given a number of smaller specimens I have
never yet seen one that approaches in size the medium-sized Gaboon
viper. I presume they eat fish, at least sometimes, at any rate,
since two of my specimens have been taken from the natives' fish
traps, where they had drowned.

"This species is even more brilliant than gabonicus. The dark
triangle on the head, however, is always black or dark brown,
not reddish as it is sometimes depicted. I doubt that Bitis arietans
occurs here in the rain forest."

Amphibians and Reptiles from Liberia 1227

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1228 The University Science Bulletin

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Hemidactylus formosus described ) .
LovERiDGE, Arthur

1936. African reptiles and amphibians in the Field Museum of Natural
History. Field Mus. Nat. Hist., Zool. Ser., vol. 22, 1936, pp. 1-111.

Amphibians and Reptiles from Liberia 1229

1938. On a collection of reptiles and amphibians from Liberia. Proc.

New England Zool. Club, vol. 17, 1938, pp. 49-75.
1941. Report on the Smithsonian-Firestone Expeditions collection of rep-
tiles and amphibians from Liberia. Proc. U. S. Nat. Mus., vol. 91,

no. 3128, 1941, pp. 113-140, 1 map (fig. 34).
Mkhtens, Robert

1933. Die Scincidengattung Cophoscincus Valliant. Zool. Anz., Band

102, Heft 7, 8, Apr. 15, 1933, pp. 188-190.
1938. Herpetologische Ergebnisse einer Reise nach Kamerun. Abh.

Senckb. Naturf. Ges., no. 442, 1938, pp. 1-52, 10 pis.
Mertens, Robert, and Heinz, Wermuth

1955. Die rezenten Schildkroten Krokodile und Briickenechsen. Zool.

Jahrb., Band 83, Heft 5, 1955, pp. 323-440.
Hampton, Wilder

1927. The caecilian genera Uraeotyphlus and Geotrypetes. Ann. Mag.

Nat. Hist., ser. 6, vol. 15, 1927, pp. 328-
Parker, H. W.

1931. Some new and rare frogs from West Africa. Ann. Mag. Nat. Hist.,

ser. 10, vol. 7, 1931, pp. 492-498.
1936. Amphibians from Liberia and the Gold Coast. Zool. Meded. vol.

19, 1936, pp. 87-102.
1936a. The amphibians of the Mamfe Division, Cameroons, ( 1 ) Zoo-
geography and systematics. Proc. Zool. Soc. London, 1936, pp.

135-163, figs. 1-8, pi. 1.
1936b. Dr. Karl Jordan's expedition to southwest Africa and Angola.

Herpetological collections. Nov. Zool., vol. 40, 1936, pp. 115-146,

2 figs.
Peters, W.

1876. Eine zweite Mittheilung iiber die von Hrn. Professor Dr. R.

Bucholz in Westafrica gesammelten Amphibien. Monatsb. Akad.

Wiss. Berlin, 1876, pp. 117-122, 1 pi.
Stull, Olive Griffith

1935. A checklist of the family Boidae. Proc. Boston Soc. Nat. Hist.,

vol. 40, no. 8, May, 1935, pp. 387-408.

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 15

The Tetranychidae of Mexico (Acarina)

BY

Robert E. Beer and D. S. Lang

Abstract. Mites of the family Tetranychidae known to occur in Mexico are
included in this report. Twenty-five species distributed in seven genera are
included, among which are ten previously named species reported for the first
time from Mexico and eight species new to science. The eight new species are
assigned to genera as follows: Aplonobia, two new species; Neotetranychus,
four new species; Eotetranychus, one new species; Oligonychus, one new
species.

Portions of this study were made possible through grants from
the Greater University Fund, University of Kansas, which provided
funds to finance surveys in Mexico during 1955 and 1956. The re-
port on this project has been divided into sections, the first section,
here presented, being restricted to mites of the family Tetrany-
chidae.

It is undoubtedly true that the additional records of known
species of tetranychid mites plus the new species described from
collections made in Mexico in connection with this study represent
a very small increment of the total number of species of mites in
this family that occur in Mexico. The acarine fauna of Mexico has
been virtually ignored by most collectors, or if not ignored then
collections have been retired to some inaccessible recess where they
will remain until renewed interest in this group of anthropods de-
velops. It is hoped that this paper will contribute in a small way
to the arousal of this interest in Mexican tetranychid mites and will
stimulate field workers in Mexico to collect more assiduously the
mites of this and other acarine families. It will be efforts of this
sort that will ultimately provide the types of fundamental informa-

Contribution number 954 from the Department of Entomology of the University of
Kansas.

( 1231 )

1232 The University Science Bulletin

tion necessary for the solution of complex problems in speciation,
distribution and the like that will result in a better understanding
of the relationships existing in the world fauna within the order
Acarina. The importance of tetranychid mites to agriculture in-
creases the urgency for discovering these relationships, for it is
probably true that many forms as yet undescribed pose serious
threats to areas of intensified and specialized crop production.

Family Tetranychldae Donnadieu

The family Tetranychidae can be readily recognized by having
the basal segments of the chelicerae fused to form a mandibular
plate or stylophore. The stylophore therefore embraces the two re-
curved bases of the movable styletlike chelicerae which are long
and slender appendages adapted for piercing. The fourth segment
of the palpus bears a prominent claw situated dorsally at the distal
end of the segment. The palpal tarsus, displaced to a ventral posi-
tion by the tibial claw, provides a chelate arrangement of these
terminal elements of the appendage often referred to as the "thumb-
claw" process. There are not more than sixteen pairs of dorsal body
setae. Tarsus I usually has two pairs of duplex setae and tarsus II
bears a single pair.

Immediately following is a key to the genera of the family
Tetranychidae now definitely known to be represented by species
in Mexico. Since excellent keys are now available in the compre-
hensive work of Pritchard and Baker (1955), the present authors
feel that this simplification of the key, which excludes several
genera that may subsequently be collected in Mexico, is justifiable.

Key to Genera Known to Occur in Mexico

1. Empodium bearing tenent hairs; duplex setae at extreme end of tarsus;

female with three pairs of anal setae, male with five pairs of anal

setae 2

Empodium lacking tenent hairs; female with two pairs of anal setae,
male with four pairs 3

2. Four pairs of dorsal propodosomal setae present; true claws long and

well developed Bryobia

Three pairs of dorsal propodosomal setae present; true claws absent,

Aplonobia

3. Tarsus I empodium rudimentary; less than two pairs of duplex setae

present Euteiranychm

Tarsus I empodium well developed; two pairs of duplex setae present . 4

4. Two pairs of para-anal setae present; peritreme usually ends in a simple

bulb 5

One pair of para-anal setae present; peritreme usually ends in a
chambered hook 6

Tetranychidae of Mexico (Acarina) 1233

5. Empodium divided on distal two thirds into three pairs of hairs, except

leg I and II of male Eotetranychus

Empodium not divided on at least the proximal one half,

Neotetranycli r/A

6. Empodium with distal end divided into three pairs of hairs, Tetranychus
Distal end of empodium clawlike not divided, empodial claw sub-
tended proximoventrally by numerous hairs OUgonychus

Genus Bryobia Koch
The genus Bryobia is distinctive in possessing four pairs of dorsal
propodosomal setae. The true claws are well developed with lateral
tenent hairs and they are usually retrorse on the distal end. The
peritreme usually ends in an elongated chamber or an anastomosing
enlargement.

Bryobia baked (McGregor)

The type specimens of this species were collected from Mt. Popo-
catapetl near Mexico (City) at 12,000 feet elevation from the trunk
of a pine tree. A single female was collected in Texas by R. K.
Fletcher on wheat.

Bryobia praetiosa Koch

Reported records of Bryobia praetiosa indicate that it is widely
distributed throughout the world. The following collections are
from Mexico: 9 miles east of Huezotzingo, Puebla, June 27, 1955,
R. E. Beer and D. S. Lang, under rocks; 6 miles south of Avila
Camacho, Puebla, June 29, 1955, R. E. Beer and D. S. Lang, under
rocks; 7 miles northwest of Leon, Guanajuato, August 19. 1954,
W. T. Atyeo, under rocks.

Genus Aplonobia Womersley

The genus Aplonobia has the true claws reduced to slender pads
bearing a pair of tenent hairs distally and the empodium consists of
a slender pad with ventrally directed tenent hairs. There are only
three pairs of dorsal propodosomal setae present. The termination
of the peritreme is variable. The inner sacral setae are in line with
the dorsocentral setae and resemble a fourth pair of dorsocentrals.

Aplonobia verrucosa sp. nov.
(Figs. 2, 3, 4, 5)
Female: Terminal sensillum of palpus longer than claw, slender,
spinelike; dorsal fusiform setae slender, half as long as sensillum.
Stylophore about as long as broad, emarginate on posterior border,
striations longitudinal; peritreme strongly elbowed, ending in en-
larged branched leaflike chamber. Tarsus I with three ventral tactile
setae proximal to duplex setae, proximal member of duplex setae

1234 The University Science Bulletin

minute; tibia I with four tactile and three sensory setae. Tarsus II
with three ventral tactile setae proximal to duplex setae; tibia II
with four tactile and two sensory setae. First and fourth pairs of
legs longer than legs II and III, legs of uniform thickness to end of
tarsus, abruptly angled at union of empodium; empodium on slender
pedicel. First pair of dorsal propodosomal setae half as long as
other setae, slightly petiolate, plumose; remainder of dorsal setae
uniformly tapering to point, much longer than interval separating
them, set on strong tubercles. Humeral setae absent. Integumen-
tary striae transverse on propodosoma, longitudinal in areas of
hysterosomal and sacral setae. Body thrown into numerous folds.
Length of body 420^, including rostrum 505pi; greatest width of
body 340>.

Male: Similar to female. First pair of legs longer than other
legs. Aedeagus long, gently undulating, leveling slightly dorsad,
broadest at base, distal end abruptly tapering to rounded tip.
Length of body 412[;., including rostrum 500pt..

In life the females are green with yellowish legs and males are
yellow with legs pink or orange near their apices. Although quite
abundant on the host plants in the area, they were scattered on both
surfaces of the leaves indicating an absence of the gregarious habit.
Webbing was absent. Mites were present in equal numbers on old
and new foliage. No eggs were seen though they were probably
abundant but concealed beneath the dense pubescence of the under
leaf surfaces. Several males were assembled around most of the
molting, quiescent females.

Holotype: Male, 10 miles south of Chila, Oaxaca, Mexico, July 6,
1956, R. E. Beer, on Lippia graveolens.

Allotype: Female, same data as holotype.

Paratypes: Several males and females with same data as holotype.

Location of types: Holotype, allotype and several paratypes of
both sexes in the Snow Entomological Museum, University of Kan-
sas. Four males and four females (paratypes) deposited in the
United States National Museum.

Aplonobia dyschima sp. nov.
(Figs. 3, 6)
Female: Terminal sensillum of palpus thin, setalike. Stylophore
with longitudinal striae; peritremes protruding anteriorly, elbowed,
enlarged anastomosing chambers at distal ends. Duplex setae on
tarsus I adjacent, proximal member short, less than one fourth as
long as distal member; four plumose tactile setae on dorsal surface,

Tetranychidae of Mexico (Acarina) 1235

eight tactile and one sensory setae on ventral surface proximal to
duplex setae; tibia I with six plumose tactile setae and one simple
sensory seta on dorsal surface. Five plumose tactile setae on ventral
surface. Tarsus II with proximal member of duplex setae plumose,
one third as long as distal member, four plumose tactile setae and
three sensory setae proximal to duplex setae; tibia II with seven
plumose tactile setae. Empodium of all legs longer than pad of
true claw. First pair of dorsal propodosomal setae lanceolate, longer
than other dorsal setae which are spatulate. Striations fine, trans-
verse; surface of body thrown into folds. Length of body 495[;.,
including rostrum 555[x; greatest width 410[jl.

Male: Unkown.

Holotype: Female, General Teran, Nuevo Leon, Mexico, July 19,
1954, W. T. Atyeo, on Acacia sp.

Paratypes: Several females, same data as holotype.

Location of types: Holotype and several paratypes in Snow
Entomological Museum, University of Kansas. Two paratypes de-
posited in the United States National Museum.

Aplonobia dyschima resembles Monoceronychus corynetes in
having subspatulate dorsal setae and in the general body shape
but can be separated by the length of the front legs. M. corynetes
has the front pair of legs one and one-half times as long as the
body while the legs of the first pair of A. dyschima are approxi-
mately as long as the body. Further separation is by the duplex-
setae on tarsus I, A. dyschima has the proximal member plumose
and one-third as long as the distal member.

Pritchard and Baker ( 1955 ) state that M. corynetes is intermedi-
ate between Aplonobia and Monoceronychus due to the round body
and in lacking a propodosomal projection, but these authors in a
recent consultation have indicated that a transferral of M, corynetes
to the genus Aplonobia is forthcoming in a future publication.
Aplonobia is similar to M. corynetes in the position of the inner
sacral setae, general body shape and in the degree of the pro-
podosomal modification. However, because of the submedian
position of the inner sacrals, the absence of an anterior dorsal
projection of the propodosoma and the presence of a single rather
than two dorsal shields serve to separate both M. corynetes and A.
dyschima from the typical Monoceronychus. In Pritchard and
Baker's key A. dyschima will run to A. myops but differs from this
species in dorsal setation and other characters. The future dis-
covery of additional species of Aplonobia and Monoceronychus may
provide a better understanding and delineation of the genera.

1236 The University Science Bulletin

Genus Eutetranychus Banks
The genus Eutetranychus can be recognized by lacking the two
pairs of duplex setae on tarsus I and II. A single pair of associated
setae, resembling duplex setae, is often present. The empodium
is greatly reduced and appears absent and the true claws are re-
duced to short pads with a pair of distally directed tenent hairs.

Eutetranychus banksi (McGregor)
This mite is apparently widely distributed, being reported from
North, Central and South America, Europe, Africa and Asia. Its
host associations are numerous with no significant plant relation-
ships indicated. The following collections from Mexico are here
recorded: El Salto Falls, San Luis Potosi, Mexico (Citrus sp. );
6 miles west of Teziutlan, Puebla, Mexico ( orange ) ; 16 miles north
of Juchitan, Oaxaca, Mexico (Vatairea Lundellii). In life, females
are brownish and males reddish-brown. They inhabit the upper
surfaces of the leaves. No webbing is produced.

Genus Neotetranychus Tragardh
The genus Neotetranychus is distinctive in having the single em-
podial claw well developed and undivided on at least the proximal
half. There are two pairs of para-anal setae present and the peri-
treme usually ends in a simple bulb.

As an aid to the identification of the known species of the genus,
a complete key to species of Neotetranychus follows. This key
includes several species not now known to occur in Mexico.

Key to the Species of Neotetranychus

1. Body with dorsal setae longer than longitudinal interval separating

adjacent setae 2

Body with dorsal setae shorter than longitudinal interval separating
adjacent setae 5

2. Dorsal setae set on tubercles 3

Dorsal setae not set on tubercles 4

3. Tibia II with seven tactile setae ( Europe ) rubi

Tibia II with six tactile setae siccus

4. Empodium with three pairs of hairs at distal end undulatus

Empodium simply clawlike hamus

5. Empodium with three pairs of hairs at distal end virginiensis

Empodium simply clawlike 6

6. Dorsocentral hysterosomal setae slender, shorter than the longitudinal

interval between adjacent setae hakea

Dorsocentral hysterosomal setae not slender 7

7. Dorsal setae leaflike flabellosetus

Dorsal setae oblong spatulate hispidosetus

Tetranychidae of Mexico (Acarina) 1237

Neotetrcmychus hispidosetus sp. nov.
(Figs. 1, 3, 4)

Female: Terminal sensillum of palpus two and one-half times as
long as broad; fusiform setae slender, longer than sensillum. Stylo-
phore about as broad as long, strongly emarginate on posterior
border, striations longitudinal; peritreme straight distally ending in
simple bulb. First and third pairs of dorsal propodosomal setae
longer than interval separating adjacent setae, second pair much
shorter, broader and rounder at tip; inner sacral setae small, spatu-
late, outer sacral setae twice as long, swollen at tip. Integumentary
striae beadlike, irregular, longitudinal in area of dorsal propo-
dosomal setae, transverse in area of dorsocentral hysterosomal setae,
transverse in area of sacral setae. Tarsus I with duplex setae ad-
jacent, proximal member small, four tactile and three sensory setae
proximal to duplex setae; tibia I with seven tactile and two sensory
setae. Tarsus II with two tactile and one sensory seta proximal to
duplex setae; tibia II with four tactile and two sensory setae.
Length of body 480[j., including rostrum 560[x; greatest width 316[x.

Male: Similar to female. Terminal sensillum of palpus minute;
dorsal fusiform setae slender, long. Sacral setae all similar in
length and shape. Leg I approximately as long as body; tarsus I
with four tactile and three sensory setae proximal to duplex setae;
tibia I with nine tactile and two sensory setae. Tibia II with five
tactile setae. Aedeagus gently curves dorsad, ending in compact
enlargement, ventral angles approximately equal. Length of body
390;;., including rostrum 430|x.

Holotype: Male, 17 miles north of Tehuitzingo, Puebla, Mexico,
July 17, 1955, R. E. Beer and D. S. Lang on Erythrina sp.

Allotype: Female, same data as holotype.

Paratypes: Several males and females with same data as holotype.

Location of types: Holotype, allotype and several paratypes of
both sexes located in the Snow Entomological Museum, University
of Kansas. Five males and one female (paratypes) deposited in
the United States National Museum.

This species is pale yellow and slightly speckled with brown in
life. The types were collected from a very large population of
mites that was present in equal numbers on both surfaces of the
leaves of the host plant (coral tree). Eggs were pale yellow and
spheroid, webbing sparce. Damage to the host plant was severe.

1238 The University Science Bulletin

Neotetranychus hamus sp. nov.
(Figs. 1, 4)

Female: Terminal sensillum of palpus large, three times as long
as broad; dorsal fusiform setae slender, oblong. Stylophore one
and one-half times as long as broad, strongly emarginate on pos-
terior border; peritreme straight distally ending in simple bulb.
Leg I slightly longer than other legs; tarsus I with four tactile setae
proximal to duplex setae, proximal member of duplex setae half
as long as distal member; tibia I with eight tactile and two sensory
setae. Tarsus II with proximal member of duplex setae half as
long as distal member; tibia II with six tactile setae. Empodium
of all legs similar. Dorsal setae of body petiolate, pubescent,
slightly expanded near base, tapering to point, about as long as
longitudinal interval separating them. Integumentary striae trans-
verse. Length of body 300tx, including rostrum 370ix; greatest width
of body 228pL

Male: Similar to female. Terminal sensillum of palpus minute,
dorsal fusiform setae long, slender. Tarsus I with three tactile
and two sensory setae proximal to duplex setae; proximal member of
duplex setae half as long as distal member; tibia I with seven tactile
and two sensory setae. Tibia II with six tactile setae. Aedeagus
bends dorsad, corniform, one and one-half times as long as broad,
tapering abruptly from base to pointed apex. Length of body
290pi, including rostrum 340pi.

Holotype: Male, 5 miles north of Tepic, Nayarit, Mexico, July 26,
1956, R. E. Beer.

Allotype: Female, same data as holotype.

Paratypes: Several males and females with same data as holotype.

Location of types: Holotype, allotype and several paratypes of
both sexes deposited in the Snow Entomological Museum, Univer-
sity of Kansas. Two males and two females (paratypes) deposited
in the United States National Museum.

This species differs from N. siccus in length of members of duplex
setae, dorsal setation not being set on tubercles, and terminal sen-
sillum of palpus being broad. Aedeagus is most similar to N. rubi
but is not as slender nor as pointed and large; not sickle-shaped as in
N. rubi.

In life this mite is elongate and greenish-yellow in color. Eggs
are pale yellowish and spheroid. Webbing was not apparent in live
colonies. The identity of the host plant is not known but field notes
indicate that it was a woody shrub or tree with trifoliate leaves
(Leguminosae?). Damage to the host was moderately severe.

Tetranychidae of Mexico (Acabina) 1239

Neotetranychus flabellosetus sp. nov.
(Figs. 1, 3, 4)

Female: Terminal sensillum of palpus two and one-half times as
long as broad, uniform diameter on proximal two thirds, tapers to
angulate tip on distal one third. Stylophore twice as long as
broad, emarginate on posterior border, broadly rounded on anterior
margin; peritreme ends in a simple bulb. Tarsus I with duplex
setae adjacent, proximal member of duplex setae small, less than
one-fourth as long as distal member, five tactile setae proximal to
duplex setae; tibia I with seven tactile and two sensory setae.
Tarsus II with proximal member of duplex setae one-third as long
as distal member, three tactile and one sensory setae proximal to
duplex setae; tibia II with six tactile setae. Empodium of all
legs simply clawlike. First pair of dorsal propodosomal setae over
twice as long as other propodosomal setae, clunal setae similar
to first pair of propodosomal setae, remaining dorsal setae short,
spatulate, plumose. Integumentary striae transverse except for
longitudinal striae in area of sacral setae. Length of body 450pi, in-
cluding rostrum 505^; greatest width of body 324jx.

Male: Similar to female. Tarsus I with six tactile setae proximal
to duplex setae; tibia I with seven tactile and four sensory setae.
Aedeagus broad at base, curving slightly dorsad, terminating on
distal end in an angulated enlargement, dorsal and ventral angula-
tions approximately equal. Length of body 300[j., including rostrum
38(V.

Holotype: Male, 16 miles north of Juchitan, Oaxaca, Mexico,
July 13, 1955, R. E. Beer and D. S. Lang, on Cassia bacillaris.

Allotype: Female, same data as holotype.

Paratype: One female, same data as holotype.

Location of types: Holotype, allotype and paratype in the Snow
Entomological Museum, University of Kansas.

The male of Neotetranychus flabellosetus resembles N. virginien-
sis and N. hispidosetiis in the appearance of the aedeagus, N. virgin-
iensis can be recognized by having the empodium divided into three
pairs of hairs. N. hispidosetiis can be separated from N. flabello-
setus by the dorsal setation, N. flabellosetus has short, spatulate setae
on the dorsum while JV. hispidosetiis has oblong, spatulate setae.

Neotetranychus undulatus sp. nov.

(Figs. 1, 4)

Female: Terminal sensillum of palpus two and one-half times as

long as broad; dorsal fusiform setae broad, ovate. Stylophore

broadly rounded, approximately two times as long as broad, emar-

1240 The University Science Bulletin

ginate on posterior margin, with longitudinal striations. Peritreme
straight proximally, bent distally to form an elongated chamber.
Tarsus I with five tactile setae proximal to duplex setae, proximal
member of duplex setae small, one-fourth as long as distal member;
tibia I wtih seven tactile and two sensory setae. Tarsus II with
proximal member of duplex setae one-fourth as long as distal mem-
ber, three tactile and one sensory setae proximal to duplex setae;
tibia II with six tactile setae. Dorsal setae of body slender, about
as long as longitudinal interval separating them. Integumentary
striae transverse except for longitudinal striations in area of sacral
setae. Length of body 560^, including rostrum 600[a; greatest width
of body 380>.

Male: Similar to female. Integumentary striae transverse. Tar-
sus I with proximal member of duplex setae one-half as long as
distal member, three tactile setae and two sensory setae proximal
to duplex setae; tibia I with nine tactile and two sensory setae.
Tibia II with seven tactile and two sensory setae. Aedeagus long
and slender, strongly undulate, tapering to rounded tip. Length of
body 420pi., including rostrum 540[x.

Holotype: Male, two miles west of Antiguo Morelos, Tamaulipas,
Mexico, August 3, 1955, R. E. Beer and D. S. Lang, on Beaucormea
stricta.

Allotype: Female same data as holotype.

Paratypes: Two females, same data as holotype.

Location of types: Holotype, allotype and one paratype in the
Snow Entomological Museum, University of Kansas. One para-
type deposited in the United States National Museum.

In life, mites of this species are pale yellow in color. They as-
semble in small colonies on the under sides of the slender leaves
of the host plant. Eggs are pale yellow and spheroid and webbing
is sparse. (Author's note: A large collection taken in the type
locality in 1956 was lost in transit. )

Genus Eotetranychus Oudemans

The genus Eotetranychus possesses two pairs of para-anal setae.
The duplex setae on tarsus I are adjacent and the legs tend to
terminate abruptly. The empodium consists of three pairs of dis-
tally directed hairs and the peritreme usually ends in a simple bulb.

Tetranychidae of Mexico (Acarina) 1241

Eotetranychus oistus sp. nov.
(Figs. 2, 4)

Female: Terminal sensillum of palpus twice as long as broad,
dorsal fusiform setae slender, about as long as terminal sensillum.
Stylophore one and one-half times as long as broad, emarginate on
posterior border, abruptly rounded on anterior border; peritreme
straight distally ending in simple bulb. Tarsus I with four tactile
setae proximal to duplex setae, proximal member of duplex setae
one-fourth as long as distal member; tibia I with six tactile and four
sensory setae. Tarsus II with four tactile setae proximal to duplex
setae; tibia II with eight tactile setae. Dorsal body setae gently
tapering to a point, longer than interval separating adjacent setae;
dorsal striations transverse. Length of body 390^ including rostrum
450[x; greatest width of body 250(jl

Male: Similar to female. Terminal sensillum of palpus minute,
appearing absent. Tarsus I with four tactile and two sensory setae
proximal to duplex setae. Tarsus II with three tactile and one short
sensory setae proximal to duplex setae. All legs short and stubby.
Aedeagus small, twice as long as broad, dorsal surface with inden-
tation, tapering abruptly to a point, ventral surface uniformly taper-
ing to tip. Length of body 300[a, including rostrum 380[x.

Holotype: Male, Atlixco, Puebla, Mexico, July 19, 1955, R. E.
Beer and D. S. Lang, on Ipomoea arborescens.

Allotype: Female, same data as holotype.

Paratypes: Several males and females with same data as holotype.

Location of types: Holotype, allotype and several paratypes of
both sexes located in the Snow Entomological Museum, University
of Kansas. Two males and one female (paratypes) in the United
States National Museum.

Eotetranychus oistus belongs to the Tiliarum Group of Pritchard
and Baker. It resembles E. malvestris but can be separated easily
by differences in the aedeagi. Tight colonies were present along
the mid-vein of the leaf on the under surface. The mites were
yellow in life, eggs spheroid and webbing sparce. Damage to the
leaves of the host plant was severe.

Eotetranychus deflexus (McGregor)
This species has heretofore been reported only from California
and Oregon with the only recorded host, snowberry ( Oregon ) . The
following collections are now recorded: 19 miles north of Huachi-

1242 The University Science Bulletin

nango, Puebla, Mexico (blackberry); 12 miles west of Huachinango,
Hidalgo, Mexico (Quercus crassifolia) . Mites are yellowish in life
and form dense colonies on the under surfaces of the leaves of the
host plant. Webbing is dense, eggs pale and spheroid. Damage to
blackberry was severe but only negligible on Quercus.

Eotetranychus carpini (Oudemans)

Pritchard and Baker have indicated subspeciation in this species
as an answer to variability in certain morphological characters that
is correlated with distribution. According to their proposed group-
ings, the following collections based upon the morphological dis-
tinctions they suggest, are recorded for E. carpini carpini'. Mexico
(city), D. F. Mexico, (cottonwood and sycamore); 19 miles north
of Huachinango, Puebla, Mexico (alder); 4 miles south of Teziut-
lan, Puebla, Mexico (alder). The subspecies carpini has hereto-
fore been regarded as restricted to the Old World, with borealis
occurring in western North America. This Mexican record for
carpini therefore represents a significant distribution record, in ex-
tending carpini into the Western Hemisphere. Mites from the
above Mexican collections were in each case inflicting serious
damage upon the host plants. Colonies are formed on the under
sides of leaves. Eggs are pale and spheroid and webbing abundant.
Live mites are yellow in color. Previous host records from Canada,
Europe and the United States indicate a restriction to deciduous
trees, shrubs and brambles which is compatible with observations
on the Mexican collections here recorded.

Eotetranychus steganus Pritchard and Baker

This species has previously been known from a single area,
Florida, and its host associations restricted to a single plant group,
palmetto. A second collection is here recorded as follows: El Salto
Falls, San Luis Potosi, Mexico (Sabal mexicana).

Eotetranychus ecclisis Pritchard and Baker

This species is known from a single host species, Artemisia mexi-
cana, and at the present time the known distribution is restricted
to the type collection data as follows: Mexico (city) — Cuernavaca
Highway, kilometer 67, January 22, 1941, E. W. Baker.

Eotetranychus perplexus (McGregor)

This species is recorded from California and Idaho on Cerco-
carpus, from California on Salix, from Washington on Primus emar-
ginatus, and from British Columbia, California and Washington on
Purshia, The following distribution and host record is added at

Tetranychidae of Mexico (Acarina) 1243

this time: 17 miles north of Tehuitzingo, Puebla, Mexico, July 18,
1955, R. E. Beer and D. S. Lang, on Cryptocarpa procera. In life
these mites were small and yellowish. They assembled in small,
tight colonies on the lower surfaces of the leaves. Eggs were pale
yellowish and webbing abundant.

In the Mexican collection the male aedeagus deviates slightly
from the configuration ordinarily associated with the species, in
being bent more strongly ventrad just before its apex. However,
this difference is not considered to represent more than that which
would be expected in comparing specimens varying in appearance
because of orientation on the slide.

Genus Oligonychus Berlese

The genus Oligonychus has the empodium divided into several
hairs proximoventrally and hooklike on the distal end. A single
pair of para-anal setae is present and the dorsal setae are not set on
tubercles.

Oligonychus flexuosus sp. nov.
(Figs. 2, 3, 4)

Male: Terminal sensillum of palpus pronounced, more than twice
as long as broad; dorsal fusiform setae nearly as long as sensillum.
Stylophore broadly rounded anteriorly, slightly emarginate pos-
teriorly, with longitudinal striae. Peritreme straight distally, ending
in a simple bulb. Tibia I with nine tactile and one sensory setae;
tarsus I with four tactile setae proximal to duplex setae; duplex
setae widely spaced, proximal member short; empodium I split to
form two stout curved claws. Tibia II with five tactile setae;
tarsus II with four tactile setae proximal to duplex setae; tibia II
with seven tactile setae. Dorsum of body with slender tapering
setae, longer than the interval separating adjacent setae; dorsal
striae transverse in area of histerosomals, longitudinal posterior to
sacral setae. Four pairs of genitoanal setae present. Aedeagus
broad at basal fifth, abruptly tapering to a uniformly slender shaft
which curves dorsad; distal fifth broadly curving posteriorly.
Length of body 305^, including rostrum 334pi.

Female: Similar to male. Terminal sensillum of palpus shorter
and broader than male, dorsal fusiform setae slightly longer than
sensillum. Empodium I divided to form two claws, three pairs of
empodial hairs below the empodial claws. Two pairs of anal setae
present. Length of body 362[A, including rostrum 411[x; greatest
width of body 234^.

1244 The University Science Bulletin

Holotype: Male, 10 miles south of Chiapa, Chiapas, Mexico,
July 12, 1955, R. E. Beer and D. S. Lang, on wild grass (Paspalum
humboldianum) .

Allotype: Female, same data as holotype.

Paratypes: Several males and females of both sexes, same data as
holotype.

Location of types: Holotype, allotype and several paratypes in
the Snow Entomological Museum, University of Kansas. Two
males and two females deposited in the United States National
Museum.

Oligonychus flexuosus belongs to the Pratensis Group of Pritchard
and Baker. Members of this group can be recognized by empodium
I of the male which is split into two claws, aedeagus bending
dorsad, tibia I bearing nine tactile setae and tarsus I bearing four
tactile setae. The peritreme ends in a simple bulb. The females of
this group cannot be differentiated as to separate species. The male
of O. flexuosus is readily recognized by the extremely long, slender
aedeagus. Other members of the group have the distal end of
the aedeagus sigmoid or with a compact enlargement, while the end
of the aedeagus in O. flexuosus is simple. The aedeagus of O. mc-
gregori resembles O. flexuosus in having a slender prolongation but
differs in having a strong barb at the base. In addition these species
can be separated by the peritremes, O. mcgregori having a retrorse
peritreme. Empodium I of O. mcgregori consists of three pairs of
empodial hairs below the slender empodial claw, while O. flexuosus
has empodium I divided into two claws.

In life both sexes were yellowish. The colonies were in tight
clusters on the lower surface of the leaves. The leaves showed a
yellow to brown discoloration in clearly defined patches embracing
the area occupied by the colony. The eggs were pale and spheroid
without a central stipe.

Oligonychus aceris (Shimer)
This species has been recorded from Indiana, Kansas, New Jersey,
New York, North Carolina, Washington, and District of Columbia
in the United States, with all recorded collections from maple. To
this distribution and host record is added the following: El Salto
Falls, San Luis Potosi, Mexico, August 1, 1955, R. E. Beer and
D. S. Lang, on sycamore. Observations of the mites at the time
the Mexican collections were made revealed that they assembled in
small colonies on the upper surface of the leaf. Feeding caused a
russet discoloration on the infested area. Adult mites were reddish-

Tetranychidae of Mexico (Acarina) 1245

brown with reddish shoulders and capitulum and pinkish legs.
Eggs were pale yellow, circular in outline, flattened and had a cen-
tral stipe. Webbing was present but not profuse.

Oligonychus propetes Pritchard and Baker
All previously recorded collections of this species were taken from
oak in the eastern United States at the following locations: North
Carolina, Virginia, District of Columbia. A collection from Mexico
is here added to the known distribution records and may be identi-
fied by the following data: 6 miles north of Jacala, Hidalgo, Mexico,
July 30, 1955, R. E. Beer and D. S. Lang, on Quercus sp. In life
small colonies of the yellow mites were present on the under sides
of the pubescent leaves and caused a yellowish discoloration of the
dark, glossy, green upper surfaces of the leaves. Eggs were pale
yellow.

Oligonychus stickneyi (McGregor)
Specimens of this species are recorded from Arizona, California,
and Florida in the United States and from Morelos in Mexico. The
U. S. collections were taken from various species of grass and from
corn and maize. The Mexican collection was taken on maize. The
following collection is here added to the Mexican distribution rec-
ord: Mexico (city), D. F. Mexico, June 22, 1956, R. E. Beer, on
grass.

Oligonychus mcgregori (Baker and Pritchard)

This species has been known previously only by the type speci-
mens which were taken from cotton at Chinandega, Nicaragua. It
is here recorded for the first time from Mexico, this collection identi-
fied with the following data: 16 miles north of Juchitan, Oaxaca,
Mexico, July 3, 1955, R. E. Beer and D. S. Lang, on Vatairea
Lundellii. In life the mites are yellowish with brown spots, the
dorsum is flattened. They cluster on the upper surfaces of the
glossy leaves of the host plant along the mid-vein. Eggs are yel-
lowish, circular in outline and flattened. Webbing is sparce. The
mites were quite abundant and damage to the host conspicuous.

Oligonychus viridis (Banks)
This species is recorded from pecan in Florida, Georgia and
Louisiana. Collections here reported from Mexico would indicate
that it is generally distributed throughout the length of this coun-
try. The following localities and host records are added to the
published information on O. viridis: El Salto Falls, San Luis Potosi,
Mexico (avocado); Mexico (city), D. F. Mexico, and Teziutlan,

1246 The University Science Bulletin

Puebla, Mexico (camellia); 4 miles west of San Cristobal de las
Casas, Chiapas, Mexico (Chimaphyla maculata). Adult females
in life are maroon or brownish with pale legs. They assemble in
small, tight colonies with moderate webbing on both leaf surfaces.
Damage to the host is a russet discoloration of the green leaves.

Genus Tetranychus Dufour

The genus Tetranychus can be recognized by the single pair of
para-anal setae and by having the duplex setae separated on tarsus
I. The legs tend to taper to the empodium which is composed of
three pairs of hairs. The peritreme usually ends in a chambered
hook.

Tetranychus mexicanus (McGregor)

This species has been collected on citrus in the following local-
ities: Laredo, Texas, Quarantine station, host (orange) shipment
originating in Mexico; Monte Alto, Texas ( lemon ) ; Valles, San Luis
Potosi, Mexico; Concordia, Argentina. To these distribution and
host records may be added the following: 20 miles south of
Acayucan, Veracruz, Mexico (wild banana); Tecolutla, Veracruz,
Mexico (coconut palm). The following information concerning
the Acavucan collection from banana was recorded in field notes:
In life, female mites are robust and maroon in color with pink legs
and gnathosoma; males are orange. Eggs are pale yellow and
spheroid, webbing profuse, and dense colonies occur on the under
sides of the leaves. The mites were exceedingly abundant and
damage to the host severe. As regards the Tecolutla collection
field notes indicate that in life, female mites are robust and red in
color.

Tetranychus pacificus (McGregor)

This species is known from California, Idaho and Oregon where
it has been reported from a multitude of hosts including several
deciduous trees and shrubs, herbaceous annuals, melons and the
like. To this distribution and host list may be added the following:
10 miles north of Matias Romero, Oaxaca, Mexico (Asarum sp.);
El Salto Falls, San Luis Potosi, Mexico (Bacconia frutescens).
In both of the Mexican collections, living mites were scattered
on the leaves of the host plants and populations small. It would
seem that the scarcity of individuals would mask a true gregarious
habit if such were the case with this species.

Tetranychidae of Mexico (Acarina) 1247

Tetranychus telarius (Linnaeus)
The following locality and hosts are added to the exceedingly
long lists of similar information for this species: Mexico (city),
D. F. Mexico, (Fraxinus sp. and Convolvulus sp.). Damage to
each of these hosts, ash planted as street trees and wild Convolvulus
weeds was very severe. In the Fraxinus collections the live mites
were the typical yellowish color phase. In the Convolvulus col-
lections the living mites were maroon in color.

Tetranychus dugesii Cano and Alcacio
As suggested by Pritchard and Baker (1955) this species is at
present under questionable assignment to the genus Tetranychus.
Type material is identified with the following information, though
the location of the type specimens is not known: Hacienda Escuela,
Mexico, on Medicago denticulata. This is the only known reference
to this species.

The authors owe a debt of gratitude to several botanists who gave
generously of their time and energy to assist with host identifica-
tions. We wish to express our thanks to Eizi Matuda and Faustino
Miranda of the Instituto Biologico in Mexico (city) and to W. H.
Horr of the University of Kansas.

REFERENCES CITED

Pritchard, A. Earl, and Raker, Edward W.

1955. A Revision of the Spider Mite Family Tetranychidae. Pacific
Coast Entomological Society, Memoirs Series, vol. 2, 472 pp., 1955.

1248 The University Science Bulletin

FIGURE 1

Figures in left column are female tibia and tarsus I. Figures in right column
are female tibia and tarsus II. All magnifications are identical.
Neotetranychus hamus sp. nov.
N. hispidosetus sp. nov.
N. jlahellosetus sp. nov.
IV. undulatus sp. nov.

Tetranychidae of Mexico (Acarina) 1249

FIGURE 1

N. homu*

N. hUpidOMtui

N. flabellosatut

N. undulotut

1250 The University Science Bulletin

FIGURE 2

Figures in left column are female tibia and tarsus I. Figures in right column
are female tibia and tarsus II. All magnifications are identical.
Eotetranychus oistus sp. nov.
Oligonychus flexuosus sp. nov.
Aplonobia verrucosa sp. nov.

Tetranychidae of Mexico (Acarina)

1251

FIGURE 2

E. oistus

0. flexuosus

A. verrucoto

1252 The University Science Bulletin

FIGURE 3

Tibia and tarsus I and II, Aplonobia dyschima sp. nov. Pretarsus of leg I
of male, Oligonychus flexuosus sp. nov. Palpus of Aplonobia verrucosa sp. nov.
Typical dorsocentral setae (identical magnification) of:

Neotetranychus hispidosetus sp. nov.

N. jiabellosetus sp. nov.

Aplonobia dyschima sp. nov.

Tetranychidae of Mexico (Acarina) 1253

FIGURE 3

A dyschimo

0- flexuosus

A. verrucosa

I
f

N hlspidosetus

N. flobellosetus

A. dyschimo

1254 The University Science Bulletin

FIGURE 4

Aedeagi of males (identical magnification):
Neotetranychus hamus sp. nov.
N. flabellosetus sp. nov.
N. hispidosetus sp. nov.
N. undulatiis sp. nov.
Eotetranychus oistus sp. nov.
Oligonychus flexuosiis sp. nov.
Aplonobia verrucosa sp. nov.

Tetranychidae of Mexico (Acarina) 1255

FIGURE 4

N. hamus

N. flobellosetus

N. hispidosetus

N. undulotus

E. oistus

0- f lexuosus

8—8050

A. verrucosa

1256 The University Science Bulletin

FIGURE 5

Adult female, Aplonobia verrucosa sp. nov. (dorsal aspect).

FIGURE 5

Aplonobia verrucosa

1258 The University Science Bulletin

FIGURE 6

Adult female, Aplonobia dyschima sp. nov. (dorsal aspect).

Tetranychidae of Mexico (Acarina)

1259

FIGURE 6

Aplonobio dytchimo

THE UNIVERSITY OF KANSAS

SGIENGE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 16

A Revision of the Bees of the Subgenus Epinomia in the
New World ( Hymenoptera-Halictidae ) *

BY

Earle A. Cross

Abstract. This is the first of two papers dealing with the systematics and
geographical variation of the New World members of the subgenus E7)inomia
Ashmead. It contains a revision of the subgenus and includes information on
distribution, geographical variation, and flower preferences of its species.
About 1500 specimens were examined.

Indications of the relationships between the subgenera of Nomia and a
subgeneric key are presented.

Four species are recognized, one of which (N. nevadensis) is further di-
vided into five subspecies. A more detailed study of the geographical varia-
tion of this species will be published elsewhere. Two species, N. boharti and
N. micheneri, and one subspecies, N. nevadensis stellata, are described as new.

TABLE OF CONTENTS

PAGE

Introduction 1261

Acknowledgments 1262

Systematic Treatment

The Genus Norriia 1263

Key to the American Subgenera of Nomia 1265

The Subgenus Epinomia 1266

Key to the American Species of Epinomia 1268

Literature Cited 1296

INTRODUCTION

The purpose of this paper is to reclassify the species of the sub-
genus Epinomia and to give a brief survey of their variation. The
subgenus is a small one, consisting of only four species, one of which
(N. nevadensis) is divided here into five subspecies. From an

* Contribution number 957 from the Department of Entomology, University of Kansas.

(1261)

1262 The University Science Bulletin

evolutionary and systematic standpoint, however, the group is in-
teresting since two of its species exhibit strikingly different forms.
A more detailed study of the geographical variation within one
species, N. nevadensis, will be published elsewhere.

Descriptions of new forms are based upon the type specimens,
variation encountered within the type series being indicated in
parentheses. The redescriptions of previously described forms are
based upon series rather than single specimens. The author has
followed the terminology utilized by Michener, 1944, for all de-
scriptive purposes. The various processes of the apical margin of
the clypeus referred to in the descriptions are obscure when viewed
in facial aspect and are best seen from beneath.

Measurements of width and length of face were made as follows:

1. Greatest possible width of face, including eyes.

2. Greatest possible length of face, taken along sagittal plane.
The former was chosen for statistical treatment because it is meas-
urable with a greater degree of accuracy than the other and be-
cause it is believed to be a good index of general size.

The first segment of the labial palpus was considered to consist
only of the sclerotized portion of that segment, the conjunctiva be-
tween palpus and palpiger being omitted from measurements. The
length of the wing was measured from the center of the subalar
area to the wing tip. No wing measurements were made of spec-
imens with spread or curled wings. The ratio of eye width to genal
width was determined with the head in perfect profile, the greatest
possible measurement between anterior eye margin and posterior
genal margin and between anterior eye margin and posterior eye
margin being used.

ACKNOWLEDGMENTS

The author gratefully acknowledges his indebtedness to Dr.
C. D. Michener, University of Kansas, for his guidance and criticism
during the preparation of this paper, and to Dr. R. R. Sokal, Uni-
versity of Kansas, for his guidance in the statistical portions of the
work.

Thanks are also due to the following persons who generously
lent specimens from their own collections or from collections in
their care: R. H. Beamer and W. E. LaBerge, University of Kan-
sas, Lawrence; G. E. Bohart, U. S. Legume Seed Research Labora-
tory, Logan, Utah; G. D. Butler, University of Arizona, Tucson,
Arizona; E. A. Chapin, U. S. National Museum, Washington, D. C.;
E. F. Cook, University of Minnesota, St. Paul, Minnesota; R. R.

Bees of the Subgenus Epinomia 1263

Dreisbach, Midland, Michigan; H. J. Grant, Jr., Academy of Nat-
ural Sciences of Philadelphia; P. D. Hurd, University of California,
Berkeley, California; M. T. James, Washington State College, Pull-
man, Washington; G. F. Knowlton, Utah State Agricultural Col-
lege, Logan, Utah; K. V. Krombein, U. S. Dept. Agriculture, Wash-
ington, D. C; I. LaBivers, University of Nevada, Beno, Nevada;

A. T. McClay, University of California, Davis, California; T. B.
Mitchell, North Carolina State College, Baleigh, North Carolina;

B. E. Pfadt, University of Wyoming, Laramie, Wyoming; L. W.
Quate, University of Nebraska, Lincoln, Nebraska; H. J. Bhinehard,
Texas Agricultural and Mechanical College, College Station, Texas;
E. S. Boss, California Academy of Sciences, San Francisco, Califor-
nia; H. H. Boss, Illinois Natural History Survey, Urbana, Illinois;
H. F. Schwarz, American Museum of Natural History, New York,
New York; H. A. Scullen, Oregon State College, Corvallis, Oregon;
L. H. Shinners, Southern Methodist University, Dallas, Texas; B.
B. Snelling, Turlock, California; P. H. Timberlake, Citrus Experi-
ment Station, Biverside, California.

I am further indebted to Dr. H. J. Grant, Jr., Academy of Nat-
ural Sciences of Philadelphia, for comparing specimens with the
type of IV. nevadensis Cresson, to Drs. E. G. Linsley, University of
California, Berkeley, and G. D. Butler, University of Arizona, for
unpublished information concerning the nesting habits of N. ne-
vadensis, and to Dr. Ira LaBivers and Mr. Ted Frantz, University
of Nevada, for use of an unpublished manuscript concerning the
ecology of a desert locality at Fairbanks Springs, Nevada.

Thanks are also due to my wife, Dorthy S. Cross, for typing the
manuscript and to Mrs. Sylvia Jost, University of Kansas, for inking
the drawings.

The Genus Nomia in the New World

The genus Nomia Latreille, 1805, (type species Andrena cur-
vipes Fabricius) is a large (Dalla Torre, 1896, lists about 130
species), diverse group, widespread in distribution, with the bulk
of its species in Africa, Asia and Australia. It is represented in
the New World by approximately twenty species which may be
divided among four subgenera: Epinomia Ashmead, Acunomia
Cockerell, Dieunomia Cockerell, and Paranomina Michener.* These

* Michener, 1944, pointed out that Puranomia Friese is preoccupied (Conrad, 1860)
and proposed Paranomina as a substitute, retaining N. chalybeata Smith as the type.
Curvinomia Michener, 1944, named in the same paper (type N. californicnsis Michener),
is a synonym of Paranomina (new synonymy) and has line precedence over it. However,
it is felt that Paranomina should take preference over Curvinomia because the former is
based upon the original type of Paranomia (See Copenhagen Decisions on Zoological
Nomenclature, 1953, page 67). Moreover, the status of N. californicnsis is in doubt.

In placing certain American species in the same subgenus as N. chah/bcata, I follow
Cockerell, 1930.

1264 The University Science Bulletin

subgenera comprise two diverse groups, those with bright integu-
mental bands on the metasoma and those with bands of hair on the
metasoma. The first group embraces Parano7nina and Acunomia.
The latter is restricted to the Americas while the former is found
in Asia (and Australia?) as well. The second group contains
Epinomia and Dieunomia. Again, the latter is found only in the
New World while the former also has close relatives in Asia and
Australia. It seems probable, therefore, that invasions of Para-
nomina- and of Epino?nia-\ike bees reached this continent from
Asia independently, probably before mid-Tertiary time, since they
do not now occur north of about 50 degrees north latitude and
hence must have reached the Western Hemisphere when the Bering
Strait area was relatively warm. These two stocks are probably
ancestral, respectively, to Acunomia and Dieunomia.

The four New World subgenera are distributed in a similar man-
ner, and range from southern Washington through Minnesota and
Virginia, southward to Florida, central Mexico and Baja California.1
One subgenus (Acunomia) is established on the Bahamas (IV.
wickhami)2, and on Cuba (N. robinsoni). No records of Nomia
are known from Central or South America except N. (Hoplonomia)
expulsa Cockerell (Cockerell, 1930) from French Guiana, and N.
(Dieunomia) heteropoda (Say) from Brazil (see Michener in
Musebeck, et al, 1951, p. 1129). Both records are apparently in-
correct. The distribution of these subgenera thus leaves little, if
any, clue as to their origin and phylogeny. Acunomia is the most
widespread, Dieunomia the least.

Acunomia and Paranomina, although easily separable, are mor-
phologically similar. The relationship between Epinomia and
Dieunomia is not as easily perceived. Certain Dieunomia, e. g. N.
heteropoda (Say), differ greatly from the Epinomia, and certain
authors have regarded Dieunomia as a distinct genus. Examina-
tion of the various species of Dieunomia, however, shows a con-
tinuous gradient of morphological divergence from the Epinomia
group, commencing with Nomia (Dieunomia) bolliana Cockerell,
whose female shows a striking resemblance to the female of N.
(Epinomia) trianguHfera Vachal.

Further evidence of the close relationship between Dieunomia
and Epinomia has been found in the nest structure. Nests of N.
(Epinomia) trianguHfera, N. (Epinomia) nevadensis, and N. (Die-

1. Provancher's N. compacta (1889) from Quebec may be a small Epinomia, but be-
cause its description is ambiguous, and since its type locality is so far from the known
range of any other Epinomia, it is not included in this paper, and is presumed to belong
to some other genus.

2. I have not seen specimens of N. wickhami Ashmead, but infer from Cockerell's de-
scription (1910), that it, like N. robinsoni, belongs in Acunomia.

Bees of the Subgenus Epinomta 1265

unomia) heteropoda are similar to one another and strikingly dif-
ferent from those of N. (Acunomia) melanderi Cockerell (Cross
and Bohart, in press).

Key to the American Subgenera of Nomia

1. Metasomal terga without hair bands, with apical pale green, bluish,

yellow, or coppery integumental bands 2

Metasomal terga with apical hair bands (often worn off), without

integumental bands 5

2. ( 1 ) Males 3

Females 4

3. ( 2 ) Ultimate segment of antenna acuminate or at least acutely pointed,

usually narrower than those preceding; basal vein of forewing
only slightly arcuate; fifth metasomal sternum with median
subapical processes contiguous or nearly so, arising from me-
dian longitudinal line Acunomia

Ultimate segment of antenna blunt, always as wide as or wider
than penultimate segment; basal vein of forewing slightly to
distinctly arcuate; fifth metasomal sternum with median sub-
apical processes distinctly separated from the median longi-
tudinal line (except in N. fedorensis which resembles Acunomia
in this respect) Paranomina

4. (2) Middle tibial spur finely and almost evenly serrate along entire

margin; basal vein of forewing slightly to distinctly arcuate,

Paranomina

Middle tibial spur with but a few, uneven, coarse teeth along

apical one third, basal two thirds finely and evenly serrate;

basal vein of forewing only slightly arcuate Acunomia

5. (1) Males ' 6

Females 7

6. (5) Labial palpus with segments two to four together shorter than,

rarely as long as, the first segment alone. Apical antennal seg-
ment distinctly wider than penultimate segment, flattened and
expanded, sometimes disclike; tegula somewhat rectangular
in shape, sinuate along lateral margin; length 10-23 mm.,

Dieunomia
Labial palpus with segments two to four together longer than first
segment alone. Apical antennal segment not wider than penul-
timate segment, somewhat flattened but obliquely rounded
distally, not expanded and disclike; tegula as described for
Dieunomia or distinctly rounded posterolateral^; length 7-15
mm Epinomia

7. (5) Labial palpus with segments two to four together rarely as long

as first segment alone; pubescence of metasomal venter black,
brown, deep orange, or reddish; tegula as in males; length

10-23 mm Dieunomia

Labial palpus with segments two to four together longer than first
segment alone; pubescence of metasomal venter white, grey-
ish, buffy, or light golden yellow; tegula as for males; length
8-15 mm Epinomia

1266 The University Science Bulletin

Using the above key, the names of the American species, as listed
by Michener, in Musebeck et al (1951), may be placed as follows:

Epinomia Dieunomia

boharti sp. nov. apacha Cresson

micheneri sp. nov. bolliana Cockerell

nevadensis Cresson heteropoda (Say)

triangulifera Vachal mesillae (Cockerell)

xerophila ( Cockerell )

Paranamina Acunomia

californiensis Michener melanderi Cockerell

fedorensis Cockerell nortoni Cresson

foxii Dalla Torre robinsoni Cresson

maneei Cockerell wickhami Ashmead

mesillensis Cockerell
parksi Cockerell
tetrazonata Cockerell
universitatis Cockerell
uvaldensis Cockerell
zabriskii Cockerell and Blair

Subgenus Epinomia Ashmead

Epinomia Ashmead, 1899, Trans. Amer. Ent. Soc, vol. 26, p. 88. Type, (N.
persimilis Cockerell )= Nomia triangulifera Vachal, monobasic and original
designation.

Diagnosis: This subgenus may be separated from its closest rela-
tive, Dieunomia, by the lengths of the segments of the labial palpi
as indicated in the key. The apical antennal segment of male
Dieunomia is distinctly wider than the penultimate segment, while
in male Epinomia the apical segment is the same width as the
penultimate. Females of Epinomia may be separated from those
of Dieunomia by the color of the hairs of the abdominal venter. In
Epinomia, these are white, greyish, buffy, or golden yellow (some-
what rufous in Mexican specimens of N. (Epinomia) nevadensis
arizonensis, while in Dieunomia they are black, brown, deep orange,
or rufous. N. bolliana Cockerell, the smallest of the species of
Dieunomia, is perhaps the only species of that subgenus that can
be confused readily with species of Epinomia. In addition to the
above-mentioned characters, N. bolliana may be known from N.
triangulifera by the shape of the horizontal rugose base of the
propodeum, which is distinctly crescentic, and by the lack of stria-
tions on the anterior lateral margins of the prothorax, from N. bo-
harti by the lack of impunctate transverse ridges on the first two
metasomal terga, and from other species of Epinomia by the shape
of the tegulae, which are rectangular and sinuate along their lateral
margins.

Bees of the Subgenus Epinomia 1267

Description: Male: Length 7-15 mm.; color variable, ranging
from wholly black to wholly fulvous or fulvo-ferruginous except for
portions of the head and mesosoma; wings yellowish-hyaline to
darkly fuliginous.

Pubescence of entire body pale greyish white to light ochraceous-
buff, color usually uniform on any one specimen; face, posterior
margin and lobes of pronotum, posterior margin of metanotum,
axilla, and propodeal spiracles all densely clothed with downy,
plumose hairs; metasomal terga one to five with distinct apical bands
of appressed hair, that of sixth tergum indistinct.

Segments two to four together of labial palpus longer than the
first alone; apical antennal segment somewhat flattened, obliquely
rounded, or almost truncate, not expanded or wider than the pre-
ceding segment; face appearing round or elliptical, eyes subparallel
or slightly convergent below, inner eye margins never strongly
curved, not strongly convergent above; tegula rectangular, sinuate
on lateral margin, or rounded posterolaterally, thus appearing oval;
basal vein of forewing only slightly arcuate; hind basitarsus usually
shorter than, not distinctly longer than hind tibia; hind tibia ex-
panded distally except in N. boharti where the dilation is very
slight; inner apical angle of hind tibia produced, forming a distinct
tooth or nodule, a median tooth along inner edge present or absent;
hind femur never greatly swollen, not wider than distance between
lateral ocelli; hind trochanter with or without a ventral apical
nodule; middle tibia and femur unmodified; dorsal base of pro-
podeum with a horizontal, strongly rugose area which may be either
crescent-shaped or distinctly truncate posteriorly; anterior face of
the first metasomal tergum with a deep, longitudinal excavation;
fourth metasomal sternum with a pair of distinct, arcuate sutures,
meeting posteriorly to form a median triangular area, or sutures
becoming mere impressions except posteriorly, or absent entirely;
fifth metasomal sternum with a pair of pubescent subapical nodules
and a pair of arcuate carinae converging posteriorly to form a
median triangular area; apex of sixth metasomal sternum recurved
and with a deep median cleft; seventh metasomal sternum ribbon-
like; eighth metasomal sternum produced to a long acuminate proc-
ess posteriorly; gonostyli converging evenly, not abruptly bent
mesad, modified setae on ventral portions of apices in the form of a
few thick bristles; gonostyli each with a posteriorly directed ventral
process.

Female: Length 8-15 mm., form more robust than male; downy
plumose pubescence as in male except that of face, which is greatly

1268 The University Science Bulletin

reduced; hair bands distinct on metasomal terga one to four, fifth
tergum bearing densely appressed pubescence of various shades
from seal brown to bright rufous, sometimes also with an indistinct
hair band anteriorly; color of pubescence white to rufous, often
varied on any one specimen; pubescence of metasomal venter long,
white to golden yellow, becoming rufous in some Mexican speci-
mens of N. nevadensis arizonensis, face wider than in male; other
structural characters as given in description of male.

Habits: Information concerning the habits of Epinomia is scanty,
and much remains to be done in this respect. Bohart ( 1952, also
in litt. ) , Cockerell ( 1934 ) , Cross and Bohart ( in press ) , Linsley
and MacSwain (in litt.), Pierce (1904), Rau (1929), and Snelling
(in litt.) have studied JV. triangulifera and N. nevadensis.

These bees nest gregariously, often in large densely populated
sites, but insofar as known they are true solitary bees, each female
digging and provisioning one or more nests in a season, after which
she dies, leaving her progeny to repeat the cycle the ensuing year.
Adults generally appear in middle or late summer, although there
is some evidence of a longer season in areas having summer rains
where it is possible that several generations occur in the same year.

Insofar as known, the females prefer the flowers of various com-
posites. G. Bohart (in litt.) mentions the possible importance of
this group in areas where sunflower seed is grown for oil.

Key to the American Species of the Subgenus Epinomia

1. Anterior lateral portion of prothorax with a series of vertical or

nearly vertical striations; basal area of propodeum distinctly
truncate posteriorly; tegula somewhat rectangular in shape,
lateral margin distinctly •sinuate; dark species, 11-15.5 mm.

long triangulifera

Anterior lateral margin of prothorax without striations; basal area
of propodeum crescent-shaped or weakly truncate posteriorly;
' tegula as in N. triangulifera or rounded posterolateral^, appear-
ing somewhat oval; species dark or with varying amounts of
fulvous or ferruginous, 7-12.5 mm. long 2

2. ( 1 ) Males 3

Females 5

3. (2) Hind tibia with distinct median excavation and usually with a

tooth basal to it along anterior inner edge (fig. 4, G, H),

nevadensis
Hind tibia without median tooth or excavation along anterior inner
edge 4

4. (3) Hind tibia greatly expanded distally, apex (including tooth) half

as wide as length of segment; tegula rounded posterolaterally,
appearing somewhat oval in shape; clypeal margin with a

Bees oe the Subgenus Epinomia 1269

single, rectangulate median process*, the apex of which is not

sinuate (fig. 3, F, G) micheneri

Hind tibia not greatly expanded distally, apex (including tooth)
not half as wide as length of segment; tegula as described for
triangulifera; clypeal margin with a single median process, the
apex of which is distinctly sinuate ( fig. 3, I ) boharti

5. (2) First two metosomal terga each with a distinct, transverse, sub-

apical, rounded carina that is impunctate and shining; tegula

as described for N. triangulifera boharti

First two metasomal terga without carinae as described above;
tegula rounded posterolaterally, appearing rather oval 6

6. (5) Apical margin of clypeus with a pair of distinct median teeth f

(fig. 3, K) micheneri

Apical margin of clypeus not dentate nevadensis

Nomia (Epinomia) triangulifera Vachal

(Fig. 1; 3A, E, I; 4 A, E)

Nomia triangulifera Vachal, 1897, Miscellanea Entomologica, vol. 5, p. 9.

( male ) .
Nomia persimilis Cockerell, 1898, Denison Univ. Sci. Labs. Bull., vol. 11, pp.

50, 72; and Bull. Univ. New Mexico, vol. 1, pp. 50, 72.

Diagnosis: Although the females of this species superficially re-
semble N. (Dieunomia) bolliana, either sex may be readily sepa-
rated from the latter, as well as from any species of Epinomia, by
the presence of a series of vertical or nearly vertical striations on
the lateral anterior portions of the prothorax. N. triangulifera is
also separable from all other forms of Epinomia except N. nevaden-
sis bakeri by the shape of the horizontal rugose base of the propo-
deum, which is truncate rather than crescent-shaped, and from
all forms of Epinomia except N. boharti by the shape of the tegulae,
which are rectangulate.

Description of male:

Size: Length 12-15.5 mm.; length of forewing 9.0-12.5 mm.;
width of face 2.9-3.2 mm.; length of face 3.0-3.7 mm.

Color: Usually wholly black, two basal metasomal segments occa-
sionally somewhat piceous; legs black to piceous; antenna black
or becoming reddish beneath; tegula translucent-testaceous to dark
brown; wings with a dull orange or brownish tint or subhy aline,
apical one fifth infuscate, sometimes lightly so; veins and stigma
yellow-brown to dark brown.

Pubescence: Pale throughout, that of head, sides and venter of
mesosoma and femora and tibiae of legs white or grayish white;
that of mesonotum greyish or pale buffy white, densest on anterior

* Seen best from beneath,
t Seen from beneath.

1270 The University Science Bulletin

one third; fasciae of metasomal terga one to five white; metasomal
sterna two and three with several thin rows of long, shortly-
plumose hairs and with short, subapical fringes of feathery plumose
setae which are recurved medially, particularly on sternum three,
and which are prolonged laterally to form tufts; sternum four with
a thin apical fringe, also becoming tufted laterally; middle femur
beneath with a few, scattered, short bristles; front femur beneath
evenly and sparsely covered with tiny setae.

Punctation: That of clypeus very fine, subcancellate, becoming
coarser above, that of vertex coarser and sparser, areas about ocelli
often minutely reticulate; punctures of mesoscutum close, coarse,
similar to those of vertex, those of mesoscutellum coarser, variable
in size and distribution, sparsest medially; first four metasomal terga
with very fine and close punctures, the first, and sometimes the
second with two distinct sizes, the coarser usually sparsely and
evenly distributed among the finer; punctures of tergum five about
as fine as the finer punctures of the first four but more sparse. All
punctured surfaces moderately shining.

Structure: Apical margin of clypeus with a wide, slightly pro-
duced truncation (fig. 3, E); first flagellar segment almost as long
as second, segments two to ten approximately equal in length,
about .48 mm. each, apical (11th) segment longer, about .65 mm;
face almost round, ratio of length to width 1:1.00-1:1.08; genal
area about one and one quarter times as wide as eye; labial palpus
with segments two to four together distinctly longer than first free
segment alone; anterior lateral portion of prothorax with a series
of vertical or nearly vertical striations; tegula with posterolateral
angle rather abrupt, appearing somewhat rectangular, shallowly
sinuate along lateral margin; horizontal rugose basal area of pro-
podeuin truncate posteriorly, somewhat polished; posterolateral
angles of propodeum distinctly angulate, becoming sharply carinate
ventrally; first three metasomal terga with shallow transverse basal
constrictions; hind trochanter quadrate, without ventral nodule;
hind femur as described for N. nevadensis; hind tibia exDanded

' IT

evenly distally, not excavated medially, roughly triangular in shape,
anterior inner margin produced to form a large apical tooth ( fig. 4,
E); fourth metasomal sternum with a pair of distinct, arcuate su-
tures which converge posteriorly to form a median triangular area;
tubercles of fifth sternum large, angulate, sinuate along posterior
margin, reticulate, lacking conspicuous pubescence (fig. 4, A); apex
of gonostylus as seen in ventral aspect acuminate-rounded (fig. 3,

Bees of the Subgenus Epinomia 1271

A); length of ventral process of gonostylus about .54 mm.; ratio of
length of ventral process of gonostylus to length of gonostylus about
1:2; subapical ventral margin of gonostylus with four to five dis-
tinct long rather stout bristles; laterodistal process of penis valve
falcate.

Description of Female:

Size: Length 11.5-14 mm.; length of forewing 9.5-11.5 mm.;
width of face 3.2-3.8 mm.; length of face 2.7-3.3 mm.

Color: As described for male except that antennae are some-
times wholly reddish.

Pubescence: That of face, sides and venter of mesosoma white,
that of mesoscutum greyish-white to pale buffy-white; fasciae of
metasomal terga one to four white, often worn on discs; fifth meta-
somal tergum with an apical fringe of thickly appressed, seal brown
hairs, another row of appressed hairs just basad, these often lighter;
apical one third of metasomal sterna two to six with veiy long
hairs, thickest on sterna four and five, greyish white to buffy white
tinged with gold; scopa long, white to buffy brown, hairs along
posterior margin of hind tibia buffy white to brown admixed with
buff; hairs on outer side of middle tibia whitish to buffy brown,
admixed with bristles of seal brown.

Punctation: Face punctured as in male; punctures of mesoscutum
coarse and close, slightly larger than those of vertex, those of meso-
scutellum variable in size but finer and much closer than those of
mesoscutum; metasomal terga punctate throughout, the first two
with fine and very close punctures, those of the third finer and
somewhat sparser, those of the fourth still finer and distinctly more
sparse than those of the third; under surfaces of front and middle
femora as described for male; all punctured surfaces moderately
shining.

Structure: Apical margin of clypeus with truncations as de-
scribed for male, or truncation reduced or absent (fig. 3. J); face
wider than that of male, ratio of length to width 1:10-1:19; genal
area about one and one-fourth times as wide as eye; tranverse basal
constrictions of first three metasomal terga shallow, often nearly
lacking, raised areas behind constrictions also lacking.

Variation: Despite its wide range, this species exhibits compara-
tively little interpopulational variation in the characters studied.
Color variation is very slight, specimens from desert localities in
New Mexico and Utah having at most the two basal terga fuscous
rather than black. Size differences are apparent within populations

1272 The University Science Bulletin

but are more difficult to establish between them. In one instance,
the means of measurements of the facial width, calculated for males
from two widely separated localities (Topaz, Utah and Lawrence,
Kansas) and using sample sizes of 18 and 25 specimens, respec-
tively, differed by only .125 mm. (3.479 mm.-3.354 mm.). A dif-
ference of means test applied to this difference, however, resulted
in a significant "t" of 2.88, (P< .01) indicating that the means of
the two populations were significantly different for the character
tested.

No differences in wing color and punctation between populations
were found, although intrapopulational variation was observed.
A slight difference in the pubescence of the legs was found to exist
between females from the Utah populations and those of the plains
populations, a high percentage of the former exhibiting more brown
hairs on the outer sides of the middle tibiae and along the posterior
margins of the hind tibiae.

Distribution: Central and west central United States, from
southern Kansas and New Mexico, north to central Minnesota and
southern North Dakota, and from central Illinois west to western
Utah (fig. 1).

Type material: The location of the type of triangulifera is not
known to me. A lectotype female of persimilis from Las Cruces,
New Mexico ( Ckll. # 4885 ) is here designated and returned to the
University of Nebraska collection.

Specimens examined: Colorado: Alamosa, Elbert. Illinois:
Meredosia. Iowa: Sergent's Bluff, Sioux City. Kansas: De Soto,
Douglas Co., Eudora, Hutchinson, Larned, Lawrence, Nickerson,
Phillips Co., Syracuse. Minnesota: St. Paul. Missouri: Ama-
zonia, Clay Co., St. Louis. Nebraska: Cedar Bluffs, Lincoln,
Mitchell, Nebraska City, Omaha, West Point, Wymore. New
Mexico: Albuquerque, Las Cruces, Montoya. North Dakota:
Mcleod, Mott. South Dakota: Elk Point. Utah: Lampo, Pen-
rose, Promontory, Topaz.

Total number of specimens examined: 254 males, 148 females.
The majority of these were caught between Aug. 10, and Sept. 15.
Males have been found as early as July 7, and females as late as
Oct. 2. No difference as to flight season between eastern and west-
ern populations is discernible.

Flower Records: Bidens involucrata, 1^,4$; Cleome semdata,
2 J ; Gilia spp., 2 g ; Grindelia squarrosa, 1^,1$; Grindelia, spp.,
2 g , 2 $ ; Helianthus annum, 16 $ , 36 $ ; H. lenticidaris, 5^,7$;
H. maximiliani, 4 ^ ; H. petiolaris, 1 £ ; Helianthus spp., 14 ^ , 4 5 ;

Bees of the Subgenus Epinomia

1273

Medicago sativa, 1 $ ; Polygonum spp., 3 $ ; Rudbeckia triloba,
1 $ , 1 $ ; Silphium perfoliatum, 4 <y , 2 $ ; Soiidago spp., 2 <? ;
Vernonia spp., 3 J1 .

Fig. 1. Map showing the distribution of N. (Epinomia) boharti, N. (£.)
micheneri, and N. (£.) triangulifera.

Nomia (Epinomia) micheneri* sp. nov.

(Figs. 1; 3B, F, G, K; 4B, F)

Diagnosis: This rather large, variably colored species is sympatric
with N. nevadensis arizonensis throughout much of its range and
superficially resembles that form. The males are immediately sepa-
rable from those of the other Epinomia by the margin of the clypeus,
which bears a single rectangulate median process, the apex of which

* This species is named for Dr. Charles D. Michener, University of Kansas.

1274 The University Science Bulletin

is not sinuate. The hind tibia of the male is also distinctive, being
broadly dilated apically and lacking a median excavation on the
anterior inner edge, somewhat resembling that of N. triangulifera.
Unlike triangulifera, however, the large inner tooth is subapical
rather than apical (fig. 4, F).

Both sexes may be distinguished from triangidifera by the absence
of striations on the anterior lateral portions of the prothorax and by
the shape of the horizontal rugose basal area of the propodeum,
which is crescentic. They may also be distinguished from triangu-
lifera, as well as from boharti, by the shape of the tegulae, which are
rather arcuate posterolaterally.

In the northern parts of its range, micheneri is distinctly larger
than nevadensis arizonensis, but in central Mexico the two species
are commonly equal in size (see section on variation). Like neva-
densis, the color of micheneri may range from wholly black to partly
fulvo-ferruginous. In partially red males, however, the first meta-
somal tergum alone is usually fulvo-ferruginous, while such males
of nevadensis usually have the first two metasomal terga colored.
The females are rather difficult to distinguish from those of nevaden-
sis, the most distinct character being a pair of small teeth situated
medially on the apical margin of the clypeus. Also, partially red
females of micheneri usually have only the first two metasomal
terga fulvo-ferruginous, while those of nevadensis generally have
the first three terga of that color. The mesoscutellum of micheneri
often bears a pair of mediolateral impunctate areas which are usu-
ally lacking in nevadensis.

Description of male:

Size: Length: 12.1 mm. (9.0-12.1); length of forewing 11.4 mm.
(9.2-11.8); width of face 2.89 mm. (2.38-2.92).

Color: Head black; mesasoma black (or posterior portion of
mesonotum fulvo-ferruginous); metasomal terga two to five black,
metasomal tergum one and the apex of tergum two fulvo-ferrugi-
nous (metasomal terga wholly black to first two and part of third
fulvo-ferruginous); coxae fuscous (to black); trochanters fuscous
(to black); fore and middle femora piceous basally, becoming
fusco-fulvous distally (or entirely black); posterior femora fulvo-
ferruginous (to black); antennae darkly fuscous (wholly black to
fuscous with reddish beneath); tegulae translucent testaceous (to
dark testaceous); wings yellowish hyaline (to brownish hyaline),
apical one fifth infuscate, nervures and stigma yellow-brown to
dark brown.

Pubescence: Pale throughout, similar to that described for the

Bees of the Subgenus Epinomia 1275

male of N. boharti. Head, sides and venter of thorax, and femora
and tibiae with long greyish-white hairs, those of mesoscutum
thin, buffy wln'te (or greyish white); fasciae of metasomal terga
one to five white (to buffy white); metasomal terga one to five
with several thin rows of long, rather stout, posteriorly directed
hairs just basad of fasciae (not as prominent on tergum one in
some), rest of these terga sparsely and evenly covered with shorter
whitish hairs (or these as long as prefascial hairs); subapical
fringes of short hair on metasomal sterna two and three as described
for N. boharti (more distinct in some micheneri), apical fringe of
sternum four absent (rather dense, long subapical fringe some-
times present); middle femora beneath bare and shining medially,
front femora also bare beneath medially, patches of tiny setae
present posterobasally, apically, and along anterior margin.

Punctation: That of clypeus and paraocular areas close and
coarse, that of vertex coarser and sparser except behind ocelli,
deeper than that of boharti; punctures of mesoscutum only a little
coarser than those of vertex, about as close, those of mesoscutellum
variable in size (to almost uniform in size), densest around peri-
phery and along median scutellar line, sparser mediolaterally (or
having impunctate areas mediolaterally), with a medium polish;
posterior surfaces of propodeum as described for boharti; puncta-
tion of first tergum coarse and rather sparse basally, becoming
finer and closer apically (to evenly and more finely punctate
throughout), second tergum with punctation as in apical portion
of first, that of third finer and closer than that of second, that of
fourth finer and as close as that of third, that of fifth still finer and
closer than that of fourth; middle femora impunctate and polished
beneath, except for a few variable punctures basally; front femora
impunctate beneath medially, but having tiny, shallow punctures
basally and anteroapically; all punctured surfaces with a medium
polish.

Structure: Apical margin of clypeus with a single, narrow, rec-
tangulate process which differs from that described for boharti in
that its apex is truncate rather than emarginate (fig. 3 F, G); an-
tennae with first flagellar segment about five-sixths as long as sec-
ond, segments two to ten equal in length, about .45 mm. (about
.41 mm. in some specimens), apical longer, about .54 mm.; face
wider than long, appearing rather elliptical, ratio of length to width
about 1:1.14; labial palpi as described for boharti; eyes about as
wide as genal areas behind them. Anterior lateral portion of pro-
thorax smooth and shining except for a few scattered punctures;

1276 The University Science Bulletin

tegula with posterolateral angle rather rounded, never abrupt;
horizontal rugose basal area of propodeum crescent-shaped; pos-
terodorsal angles of propodeum rounded, broadly so dorsally.
Transverse basal constrictions of first three metasomal terga as
described for boharti but excavations more shallow. Hind tro-
chanter quadrate, without ventral nodule; hind femur feebly
carinate basally along both anteroventral and posteroventral angles,
carina distinctly more pronounced along latter (in most, antero-
ventral carina subequal to posteroventral one ) ; hind tibia expanded
broadly apically, without median excavation, width at apex, in-
cluding inner tooth, much more than half of length; inner tooth
large, appearing subapical (fig. 4, F); fourth metasomal sternum
without converging sutures (or a pair of sutures or impressed lines
present, these converging posteriorly to form a median triangular
area as described for N. triangiilifera) ; tubercles of fifth sternum
small, nodulate, not pubescent (fig. 4, B); apex of gonostylus as
seen in ventral aspect acuminate (fig. 3, B); ventral process of
gonostylus about .35 mm. long; ratio of length of ventral process
to length of gonostylus 1:2.2; subapical ventral margin of gono-
stylus with two to five small indistinct bristles; laterodistal process
of penis valve tuberculate.

Description of female:

Size: Length, 11.9 mm. (11.0-12.2); length of forewing 10.8 mm.
(9.9-10.8); width of face 3.43 mm. (2.61-3.43); length of face
2.82 mm. (2.82-2.88).

Color: Head and mesosoma black (ferruginous spots on posterior
face of propodeum in one specimen); antennal scape fusco-fulvous
(to fuscous), flagellum black; metasomal terga three to five black,
except for a narrow fusco-fulvous line basad of the fasciae on terga
three and four (or fusco-fulvous line restricted to tergum three or
absent altogether); metasomal terga one and two fulvo-ferruginous;
coxae and trochanters darkly fulvous to fuscous (or black), remain-
ing leg segments fulvous except for basal one fourth of femora,
which are fusco-fulvous (or remaining segments entirely fulvous);
wing color as described for male but slightly darker.

Pubescence: Pale throughout, that of face, sides, and venter of
mesosoma buffy white, that of mesoscutum sparse, pale ochraceous-
buff; fasciae of metasomal terga one to four buffy white (to light
ochraceous-buff ); several thin rows of long stiff posteriorly directed
hairs present just basad of fasciae on terga one to four ( these absent
or indistinct in some Mexican specimens) pale yellow to golden in
color; apical fringe of tergum five rufescent; apical one third of

Bees of the Subgenus Epinomia 1277

metasomal sterna two to five with long, light ochraceous-buff hairs;
discs of terga one to five without feathery-plumose appressed hairs,
but with sparse erect or suberect hairs; scopal hairs long, buffy
white, hairs along posterior margin of hind tibia buffy white (or
tinged with rufous ) ; hairs on outer side of middle tibia buffy white
tinged with rufous.

Punctation: That of clypeus very coarse and close, that of vertex
a little finer and sparser except behind ocelli; punctures of meso-
scutum like those of vertex but very close, those of mesoscutellum
like those of mesoscutum, densest around periphery and along
median mesoscutellar line, leaving a pair of shining, impunctate
lateromedian areas (not well defined in all specimens); metasomal
terga punctate throughout, first two with medium-fine and rather
close punctures, punctures of third finer and as close as those of
second, those of fourth tergum still finer and somewhat closer than
those of third; middle femur beneath with coarse sparse punctures
basally and posteriorly, impunctate medially (or with a few scat-
tered punctures); front femur beneath generously and evenly
spotted with minute punctures, heaviest on basal two thirds.

Structure: Face about as wide as that of male but shorter (ratio
of length to width 1:1.22 [1:1.18-1:1.25] ); apical margin of clypeus
with a pair of small median teeth similar to those described for
boharti but less prominent; genal area about one and one-fourth
times as wide as eye; first three metasomal terga only shallowly
constricted basally, area immediately posterior to constriction -not
prominently raised.

Variation: N. micheneri is a polychromatic species comparable in
color variation to certain forms of N. nevadensis. This variation
has been stated as fully as possble in the description of the species,
and no further discussion will be made here. Its geographical
occurrence in males is shown in a general way in Table I. Because
of the paucity of specimens and lack of reliable climatological data,
no attempt will be made here to correlate climatic factors with color
differences as was done with N. nevadensis, but it seems likely
that a correlation exists as in that species.

Another important variable in micheneri is size. Table I will
suffice to show that in males this character varies rather consistently
in a general north-south direction with the largest specimens being
from the northernmost portions of the range, a notable exception
being the specimens from Jalostotitlan, Jalisco. Because of their
distinct differences in size, it is felt that the specimens examined
possibly represent more than one subspecies and it is for this reason

1278 The University Science Bulletin

that only the northernmost specimens have been designated as types.

The specimens from Jalisco are indicative of the extreme variabil-
ity which may be found in the species. Although reference to Table
I indicates that only one male specimen from San Juan de Los Lagos
was measured, two others were examined. Both were noticeably
smaller than specimens from any other part of the species range,
and in addition were partly red. Six miles northeast of Jalostotitlan,
at a distance of only six miles from the above locality, a sample of
ten very large, wholly black males was taken. Only males from
southern Arizona and northern Chihuahua were larger than these.

The sutures of the fourth sternum of the male are highly variable.
In both micheneri and boharti, these sutures may be absent, feeble
and incomplete, or distinct and complete. They are invariably
present in nevadensis and triangulifera.

Distribution: The species is apparently restricted to the Mexican
Plateau and southern Arizona. Most of the specimens before me
are from the mesquite-grassland or live-oak regions in higher alti-
tudes, ranging up to 7500 feet.

Table I — Facial measurements ( in mm. ) and color variation in males 1 of

Nomia (Epinomia) micheneri

Color 2

Mean,

Mean,

(in percent)

width of

length of

No.

Partly Wholly

face

face

Localities

specimens

red black

±S.E.

±S.E.

So. Arizona 3

•

5

100 000

3,340
±.032

2.860
±.035

Matachic,

Chihuahua

2

000 100

3.351

2.812

So. Chi-

huahua 4

14

85 15

3.079
±.028

2.585
±.025

So.

Durango 5

4

50 50

3.067
±.067

2.515
±.054

San Juan

de los Lagos,

Jalisco

1

100 000

2.862

2.416

Jalosto-

titlan,

Jalisco

10

000 100

3.174
±.026

2.591
±.028

Leon,

Guanajuato

1

000 100

3.162

2.550

1. Females are not included because adequate samples of them were not available
over most of the species range.

2. Because of the small number of specimens examined, a detailed listing of all color
differences is impractical here. Specimens listed in the column headed "partly red" had
at least the basal metasomal tergum fulvo-ferruginous.

3. Includes males from Sonoita (2); 10 miles east of Sonoita (1); 10 miles west of
Bisbee (1); mouth of Carr Canyon. Huachuca Mts. (1).

4. Includes males from Santa Barbara, Chihuahua (7); Ojito, Chihuahua (1); Jimenez,
Chihuahua (4); Canutillo, Durango (2).

5. Includes males from San Lucas (2); San Juan del Rio (1); Yerbanis (1).

Bees of the Subgenus Epinomia 1279

Type material: Holotype male from Sonoita, Arizona, Aug. 9,
1940 (P. H. Timberlake); allotype female from 25 miles east of
Sonoita, Aug. 9, 1924 (E. P. Van Duzee). Eight male and female
paratypes as follows: one male from Sonoita, Aug. 9, 1940 (P. H.
Timberlake), two males from ten miles east of Sonoita, Aug. 9, 1940
(E. S. Ross), one male from ten miles west of Bisbee, Arizona,
Aug. 10, 1940 (E. S. Ross), one male from Carr Canyon, Huachuca
Mts., Arizona, Aug. 10, 1940 (C. D. Michener), one male from near
Ramsay Canyon, Huachuca Mts., Arizona (P. H. Timberlake), one
female from Douglas, Arizona, Aug. 27, 1939 (R. H. Crandall), and
one female from Alpine, Texas, July 7, 1942 (E. C. Van Dyke).

Type repositories: Holotype, allotype, one female and two male
paratypes to the California Academy of Science, two male paratypes
to the collection of Prof. P. H. Timberlake, one male paratype to
the collection of Dr. G. E. Bohart, one male paratype to the Snow
Entomological Museum at the University of Kansas, and one female
paratype to the University of Arizona at Tucson.

Specimens examined: Chihuahua: Matachic, July 7, 1947
(Rockefeller Exped. — Michener), 2 J ; Santa Barbara, July 17-18,
1949 (Rockefeller Exped.— Michener), 3 <? , 1$; July 17, 1947
(Rockefeller Exped.— Cazier), 3^ ; Aug. 17, 1947 (G. M. Bradt),
1 S ; Ojito, 36 kilo, St. Barbara, Sept. 29, 1947 (G. M. Bradt), 1 $ ;
10 miles W. Jimenez* Sept. 11, 1950 (R. F. Smith), 4 $ . Durango:
8 miles S. Canutillo, Aug. 9, 1951 (P. D. Hurd), 2 $ ; San Juan del
Rio, Aug. 7, 1951 (H. E. Evans), \ $ ; San Lucas, Aug. 2, 1947
(Rockefeller Exped. — Michener), 2$; Yerbanis, Aug. 19, 1947
(Rockefeller Exped. — Michener), 1 $ . Zacatecas: Fresnillo, Aug.
15, 1947 (Rockefeller Exped. — Michener), 1 $ . Jalisco: San Juan
de los Lagos, July 27, 1951 (P. D. Hurd), 3 $ ; Jalostotlitlan, July 19,
1954 (Univ. Kans. Mex. Exped.) 10 $ . Guanajuato: 6 miles
N. W. Leon, Aug. 9, 1954 (Univ. Kans. Mex. Exped.), 1 £ .

Flower records: No flower records are available for the females,
but three males have flower labels attached as follows: Yerbanis,
Durango, on Helianthus; Canutillo, Durango, on Guardiola tulo-
carpa; and San Juan de los Lagos, Jalisco, on Eysenhardtia poly-
stachya.

Nomia (Epinomia) boharti* sp. nov.
(Figs. 1; 3D, I, N; 4D, I)

Diagnosis: This rather small black species most nearly resembles
certain black subspecies of N. nevadensis. It is immediately sepa-

* This species is named in honor of Dr. George E. Bohart, U. S. Legume Seed Research
Lab., Logan, Utah, without whose guidance and encouragement this paper would not have
been written, and from whose private collection the type series was taken.

1280 The University Science Bulletin

rable from the latter group, however, by the shape of the tegula,
which has the posterolateral corner abruptly bent in the same man-
ner as N. triangidifera. The males may be distinguished from those
of any other Epinomia by the distinctive shape of the hind tibia and
the shorter antenna. The females are likewise separable from
others of the subgenus by the presence of highly polished, impunc-
tate ridges immediately basad of the fasciae on metasomal terga one
to three ( more distinct on terga one and two ) .

Description of Male:

Size: Length 10 mm. (9.7-10.4); length of forewing 9.5 mm.;
width of face 2.9 mm. (2.82-3.02); length of face 2.55 mm. (2.51-
2.58).

Color: Head, mesosoma, and metasoma black; legs black to
piceous; antenna black, becoming fuscous beneath (or entirely
black); tegula very pale testaceous, subhyaline; wings brownish
hyaline ( hyaline to yellow-brown hyaline ) , veins and stigma yellow
brown to dark brown.

Pubescence: Generally pale and rather sparse, hairs often shortly
plumose; hairs of head, sides and venter of mesosoma, and femora
and tibiae white to greyish white (or buffy white), pubescence of
mesoscutum sparse, greyish white ( to buffy white ) ; fasciae of meta-
somal terga one to five buffy white to ochraceous-buff; terga three
to five with several thin rows of long, rather pale, yellowish hairs
just basad of fasciae, tergal discs otherwise evenly covered with
whitish (to pale ochraceous-buff) suberect hairs, these becoming
much longer laterally; subapical fringes of short hair on the second
and third metasomal sternum barely evident, not recurved; apical
fringe of fourth sternum absent, subapical fringe present; middle
femur beneath clear and shining but for a patch of small setae
posterobasally; front femur bare and shining except for a patch of
fine basal setae and a few coarse, scattered hairs anteriorly.

Punctation: That of clypeus and paraocular areas close and
coarse, that of vertex much sparser and coarser; mesoscutum shining,
punctures as sparse as those of vertex but coarser and deeper, those
of mesoscutellum similar to those of posterior one third of meso-
scutum, densest around periphery and along median scutellar line,
sparser in mediolateral areas; dorsolateral angles of propodeum al-
most impunctate, brightly shining, posterior face of propodeum also
shining, very coarsely and sparsely punctate; punctation of first and
second metasomal terga moderately fine, even and close, third more
finely punctate than the first two, fourth with punctures finer but no
closer than those of third; middle femur beneath impunctate ex-

Bees of the Subgenus Epinomia 1281

cept for a few fine punctures basally and posteriorly, front femur
beneath with only a few tiny basal punctures and larger scattered
punctures along anterior margin; integument in general highly
polished.

Structure: Apical margin of clypeus with a rather narrow, emar-
ginate protuberance ( fig. 3, I ) ; antenna noticeably shorter than in
other male Epino?nia, first flagellar segment about three-fourths as
long as second, segments two to ten equal in length, about .34 mm.
each, apical (11th) segment longer, about .39 mm.; face wider than
long, appearing elliptical, ratio of length to width 1:1.15 (1:1.13-
1:1.17); labial palpus with segments two to four together shorter
than those described for N. triangulifera, but still distinctly longer
than the first segment alone; eye a little wider than genal area be-
hind it; anterior lateral portion of prothorax smooth, shining, nearly
impunctate; tegula with posterolateral angle rather abrupt as de-
scribed for N. triangulifera, lateral margin shallowly sinuate; hori-
zontal rugose basal area of propodeum crescent shaped; dorsolateral
angles of propodeum rounded, broadly so dorsally; transverse basal
excavations of terga one to three deep (that of tergum three shal-
lower than basal two), the areas immediately posterior raised, first
three terga thus appearing corrugated when viewed in profile;
hind trochanter quadrate, without ventral nodule; hind femur with
posteroventral angle sharp (to carinate), anteroventral angle
rounded; hind tibia expanded only slightly apically, width at apex,
including inner tooth, much less than half the length (fig. 4, I);
inner tooth small, barely subapical; anterior inner margin of hind
tibia without a median excavation; fourth metasomal sternum with-
out converging sutures (or a pair of sutures or impressed lines
present, these converging posteriorly to form a median triangular
area as described for N. triangulifera); tubercules of fifth sternum
wide, rounded on posterior margin, with conspicuous bladelike setae
(fig. 4, D); apex of gonostylus as seen in ventral aspect, emarginate,
with a dorsal, medially directed, pubescent, triangular flange, (fig.
3, D); length of ventral process of gonostylus about .4 mm.; ratio
of length of ventral process of gonostylus to length of gonostylus
about one to two; subapical ventral margin of gonostylus with about
eight short, spinose setae; laterodistal process of penis valve tuber-
culate.

Description of female:

Size: Length 9.5 mm.; length of forewing 9.4 mm.; width of
face 2.9 mm.; length of face 2.4 mm.

1282 The University Science Bulletin

Color: That of head and mesosoma as in male, metasoma fuscous
(to black); legs fuscous, antennae tinged with reddish (to black);
tegulae as in male; wings yellow-brown (to brownish-hyaline),
apical one fifth lightly infuscate (to darkly infuscate); veins and
stigma yellow-brown to dark brown.

Pubescence: That of face, sides and venter of mesosoma buffy
white (to grayish white); dense, downy plumose pubescence of
dorsolateral angles and lobes of pronotum, mesonotum and axillae
light ochraceous-buff (darker on Nayarit female); hairs of meso-
scutum sparse, light ochraceous-buff (to ochraceous-buff); fasciae
of metasomal terga one to four light ochraceous-buff, apical fringe
of fifth tergum buffy, with a tinge of rufous; stiff, posteriorly directed
prefascial hairs absent; discs of terga one to five without feathery-
plumose appressed hairs; apical halves of metasomal sterna two to
five densely covered with long, light buff hairs; scopa buffy, hairs
along posterior margin of hind tibia ochraceous to pale rufous;
hairs on outer side of middle tibia buffy, becoming ochraceous
to pale rufous apically.

Punctation: That of clypeus coarse and close, that of vertex a
little coarser and much sparser; punctures of mesocutum about as
coarse as those of vertex but closer, those of mesoscutellum like
those of mesoscutum, densest around periphery and along median
mesoscutellar line, leaving a pair of shining, impunctate latero-
median areas; metasomal terga one to three each with a distinctly
shining, impunctate, transverse ridge just basad of fascia, more
prominent on terga one and two, terga otherwise punctate through-
out; punctures of first metasomal tergum as described for male, that
of second closer and finer than that of first, that of third much
finer ( one half the size of those on terga one and two ) and slightly
closer than that of second; middle femur beneath with a few
coarse punctures basally and posteriorly, front femur beneath
generously spotted with minute shallow punctures.

Structure: Face no wider than that of male, facial length-to-
width ratio also as described for male; apical margin of clypeus
with a pair of median teeth separated by about twice the width of
one of their bases, margin between them sinuate ( fig. 3, N ) ; width
of genal area subequal to that of eye; first three metasomal terga
strongly excavated basally, the first two more so than the third,
areas immediately posterior to excavations distinctly raised, im-
punctate, and with a high polish.

Variation: Both sexes from Nayarit exhibit darker pubescence

Bees of the Subgenus Epinomia 1283

and antennae than those from San Bernardo. Because of the small
sample sizes at hand, it was therefore deemed best to exclude
the former population from the type series. Intrapopulational
variation includes size, shape and size of the protuberances of the
clypeal margin, and, in the male, the presence or absence of the
posteriorly converging sutures of the fourth sternum. These last
two characters are also variable in N. micheneri but remain constant
in N. triangulifera and IV. nevadensis.

Distribution: Apparently restricted to the forested areas of west-
ern Mexico.

Type material: Holotype male, allotype female, and three para-
types, San Bernardo, Sonora, Mexico, Aug. 16, 1935 (collector
unknown ) .

Type repositories: Holotype and allotype to the California Acad-
emy of Sciences, one paratype to the University of California at
Davis, one paratype to the private collection of Dr. G. E. Bohart,
and one paratype to the Snow Entomological Museum at the Uni-
versity of Kansas.

Specimens examined: Nayarit: 5 mi. E. San Bias, Elev. 25 ft.,
July 25, 1953 (Kans. Univ. Exped.), 5 J , 1 ? .

Habits: Nothing is known of the habits of this species. The
Nayarit series was taken on an unidentified yellow composite which
grew in open areas in a tropical forest. Several of the males from
the same series had mud caked on the hind tibiae and presumably
had recently been underground.

Nomia nevadensis Cresson

This widespread polychromatic species occurs throughout the
southern half of the United States and is found as far south as the
state of Jalisco, Mexico. Five easily separable forms are found
within this range, and these have been retained (or designated) as
subspecies to show their close relationship. In a paper yet to be
published, the author has attempted to associate various climatic
factors with the integumental color of this species. There is evidence
that such color is well correlated with relative humidity.

The following characters will facilitate separation of this species
from the remaining Epinomia.

Diagnosis: Both sexes may be separated from specimens of IV.
triangulifera by the absence of striations on the anterior lateral por-
tion of the prothorax. The males are separable from those of the
other Epinomia by the shape of the hind tibia, which is broadly
dilated apically, and which possesses a curved median excavation

1284

The University Science Bulletin

(fig. 4, G, H). Females are distinguished from those of boharti by
the shape of the tegula, which is rather arcuate posterolaterally, and
by the absence of impunctate, shining ridges on the first three meta-
somal terga. They are more difficult to distinguish from those of
micheneri, the most distinct characters being size (in the U. S.
specimens) and the absence of distinct teeth on the apical margin
of the clypeus. Both characters are somewhat variable; the distinc-
tive characters are further discussed under N. micheneri and refer-
ence to the descriptions and to the diagnosis of the latter species is
advised.

Fig. 2. Map showing the distribution of N. (E.) nevadensis.

Nomia nevadensis nevadensis Cresson
(Figs. 2; 3C, H; 4C, H)

Nomia nevadensis Cresson, 1874, Trans. Amer. Ent. Soc., vol. 5, p. 101; also
Cresson, 1875, in Wheeler, Report upon Geographical and Geological Ex-
plorations and Surveys West of the 100th Meridian . . . Vol. 5 (Zool-
ogy), p. 722, PI. XXXIV.

Bees of the Subgenus Epinomia 1285

Description of male: *

Size: Length 10.7 mm. (measured from a specimen from Fairbank
Springs, Nev. ); length of forewing 9.8 mm.; width of face 2.92 mm.
(2.92-3.12); length of face 2.65 mm. (2.65-2.95).

Color: Face black but for apical margin of clypeus, which is
fulvous (face entirely black to face black, clypeus admixture of
fuscous and fulvous areas); mesosoma fuscous anteriorly, posterior
one third becoming fulvous (to mesosoma entirely fulvous but for
black mesoscutum ) ; metasomal terga one to three fulvous, four and
five fuscous (to metasomal terga one to two fulvous, three to five
black); coxae and trochanters fuscous to fulvous, remaining seg-
ments of legs fulvous; antennae piceous above, becoming lighter be-
neath; tegulae translucent-testaceous (to hyaline), wings pale yel-
lowish hyaline, nervures and stigma yellow brown to dark brown.

Pubescence: (The following is based upon specimens taken at
Fairbanks Springs, Nevada, since the original types have been im-
mersed in alcohol and the hair color has doubtless been altered. )

Pubescence pale throughout. Head, sides and venter of mesosoma,
and femora and tibiae with greyish white hairs, that of mesoscutum
moderately dense, thickest anteriorly, greyish white; fasciae of meta-
somal terga one to five buffy white; metasomal terga two to five with
several thin rows of long, rather stout, posteriorly directed hairs
just basad of fasciae, all terga otherwise covered evenly with shorter
thin whitish hairs; subapical fringes of short hair on metasomal sterna
two to three distinct, recurved, most prominent on sternum two;
both apical and subapical fringes of sternum four present, dense, the
subapical densest medially and laterally; middle femur bare and
shining beneath medially ( or with a few short bristles posteromedi-
ally ) ; front femur beneath rather evenly and sparsely covered with
microscopic setae, heaviest basally and along anterior margin.

Punctation: That of clypeus moderately close and coarse, that of
vertex coarser and sparser except behind ocelli; punctures of meso-
scutum coarser than those of vertex (sometimes only slightly so),
and about as close (or mesoscutal punctures closer than those of
vertex ) ; punctures of mesoscutellum of uniform size and distribution,
slightly smaller than those of mesoscutum; posterior surface of pro-
podeum as described for boharti; punctation of first two metasomal
terga moderately close and coarse (punctures of basal half of first
sparser), punctures of third and fourth almost alike, slightly finer

* Except as otherwise noted, this description is based upon a specimen ( lectallotype,
designated below) of the original type series kindly lent to me by Dr. Harold Grant of the
Academy of Natural Sciences of Philadelphia. Statements of variation in parentheses are
based upon other material of the subspecies.

1286 The University Science Bulletin

and closer than those of second, terga otherwise punctate through-
out; middle femur impunctate and polished beneath except for a
few punctures basally; all punctured surfaces with a medium
polish.

Structure: Apical margin of clypeus with a wide, shallow, weakly
emarginate process about seven times as wide as high, resembling
that of triangidifera but more prominent (fig. 3, H); antenna with
first flagellar segment about °/n as long as the second, segments two
to ten equal in length, about .39 mm. each, apical (11th) segment
longer, about .58 mm.; face only slightly wider than long, appearing
almost round, ratio of length to width 1:1.10 (1:1.05-1:1.10); eye
about as wide as genal area behind it; labial palpus as described for
N. boharti; anterior lateral portion of prothorax smooth and shining
except for a few scattered punctures; tegula with posterolateral angle
rather evenly rounded, not abruptly bent; horizontal rugose basal
area of propodeum crescent shaped; posterodorsal angle of pro-
podeum moderately rounded, not as angulate as those described for
boharti; transverse basal constrictions of first three metasomal terga
as described for N. boliarti, but more shallow; hind trochanter quad-
rate, possessing a single conspicuous nodule on its inner apex; hind
femur keeled but not distinctly carinate basally along anteroventral
and posteroventral angles (or keeled along anteroventral angle and
feebly carinate along posteroventral angle); hind tibia broadly ex-
panded apically, width at apex, including distal inner tooth, much
more than half the length of the segment, distal tooth only slightly
subapical; anterior inner margin of hind tibia with a broad, curved,
median excavation whose proximal margin becomes sharply angulate,
hind tibia thus appearing to be two-toothed (fig. 4, G, H); fourth
metasomal sternum always with a pair of distinct sutures which con-
verge apically to form a median triangular area; tubercles of fifth
sternum of moderate size, nodulate, reticulated, pubescent (fig. 4,
C ); apex of gonostylus as seen in ventral aspect tapering and rounded
(fig. 3, C); ventral process of gonostylus long, about .65 mm.; ratio
of length of ventral process of gonostylus to length of gonostylus
1:1.6; subapical ventral margin of gonostylus with two to four rather
indistinct bristles; laterodistal process of penis valve falcate.

Description of female: *

Size: Length 11.3 mm. (measured from a specimen from Fair-
banks Springs, Nevada); length of forewing 9.7 mm. (9.7-9.9);
width of face 3.1 mm. (3.1-3.2); length of face 2.7 mm. (2.7-2.9).

* This description, like that of the male, is based upon a specimen of the original type
series.

Bees of the Subgenus Epinomia 1287

Color: Head fuscous, lightest along clypeal margin (or head
black, clypeus and supraclypeals largely fulvous); antennal scape
fulvous, clavola fuscous, becoming lighter beneath; mesosoma ful-
vous, becoming somewhat fuscous anteriorly (mesosoma entirely
fulvous to mesosoma fulvous but for black mesoscutum); metasoma
entirely fulvous; legs entirely fulvous; tegula translucent-testaceous;
wings pale brownish-hyaline, nervures and stigma yellow-brown to
dark brown.

Pubescence: (Described from Fairbanks Springs specimens as in
male.) Pale throughout; hairs of face, sides and venter of meso-
soma buffy white, those of mesoscutum ochraceous-white, short and
rather dense; fasciae of metasomal terga one to four buffy white to
ochraceous-white, apical fringe of tergum five rufescent; stiff, pos-
teriorly directed hairs just basad of fasciae on terga one to four
colored as fasciae or with a pale yellow tinge, discs of metasomal
terga one to five covered with short, feathery-plumose, appressed
hairs interspersed with suberect short hairs, the former most promi-
nent on terga three and four, present only apically on the first
tergum; apical one third of metasomal sterna two to five with long,
ochraceous-white hairs, those of sterna four and five with a golden
or rufous tinge at their bases; scopa buffy white, hairs along posterior
margin of hind tibia buffy white; hairs on outer side of middle tibia
buffy white or with a tinge of gold.

Punctation: That of clypeus very coarse and rather sparse, that
of vertex finer and closer, especially behind ocelli; punctures of
mesoscutum like those of vertex but closer, those of mesoscutellum
like those of mesoscutum, almost evenly distributed; metasomal
terga punctate throughout, first with moderately fine and rather
close punctures, those of second as close but distinctly finer, those
of third still finer and a little closer than those of second, those
of fourth yet finer and about as close as those of third; middle
femur beneath with sparse, coarse punctures basally and posteriorly,
bare and shining medially and apically; front femur beneath coarsely
and evenly covered with minute punctures.

Structure: Face a little wider than that of male and shorter,
ratio of length to width 1:1.125 (1:1.117-1:1.132); clypeus long,
hiding mandibles when the latter are fully retracted; apical margin
of clypeus with a wide, very shallow, median process, the margin of
which is sinuate (superficially appearing bidentate in some), of this

9—8050

1288 The University Science Bulletin

process indistinct or absent; genal area slightly wider than eye; first
three metasomal terga only shallowly constricted, areas immediately
posterior to constrictions not prominently raised.

Variation: The specimens from eastern Nevada are slightly
smaller than the Fairbanks Springs group and, in general, somewhat
more darkly colored. In these respects, they resemble the nearby
Utah populations, and it is probable that the area of the Nevada-
Utah line is a zone of intergradation between N. nevadensis neva-
densis and N. nevadensis arizonensis. The group from Fairbanks
Springs appears to be isolated by Death Valley and mountains from
populations to the west and south.

Distribution: Known only from Fairbanks Springs, Nye Co.,
Nevada, and from White Pine Co., Nevada.

Type locality: Cresson, in the description of the species, gives
the following information regarding type locality: "8 male, female
specimens, Eastern Nevada, 1872. H. C. Yarrow."

Dr. Yarrow, in his account (Wheeler, 1875) of the 1872 expedi-
tion on which the above specimens were collected, writes: "Lieu-
tenant Hoxie's route was from Fairfield, Utah, making a detour
westward to Fillmore, Utah . . ., and following quite closely
the outward course of Captain Simpson in 1858 and 1859, the
southern limit of the so-called American Desert was crossed, the
extreme western limit reached being Schell Creek Valley, Nevada.
From this point, the direction was south by east to Snake Creek
Valley, due east across Confusion Range, . . . south to the
crossing of the Sevier, a short distance above Deseret City, and
thence to Fillmore." The above narrative leaves no doubt that the
type specimens were collected in either White Pine County, Nevada,
or in the extreme western portion of Millard County, Utah. It
seems most likely that the former is correct, and that the specimens
actually came from the vicinity of what is now McGill, Nevada.

Type repositories: Lectotype female, designated by Cresson
(1916), and lectallotype male, here designated, as well as one male
paratype are in the Academy of Natural Sciences of Philadephia.

Specimens examined: Nevada: Nye County; Fairbanks Springs,
June 21, 1949 (C. B. & R.N. Phillip), 1 <j , 1$; same locality,
June 23, 1951 (I. LaRivers, T. Frantz), 1^,3$.

Bees of the Subgenus Epinomia 1289

Nomia (Epinomia) nevadensis arizonensis Cockerel]

(Figs. 2; 3L)

Nomia arizonensis Cockerell, 1899, Entomologist, vol. 32, p. 128.

Nomia nevadensis arizonensis Cockerell, 1925, Pan-Pac. Ent., vol. 1, p. 179.

Description: As decribed for N. nevadensis nevadensis except as
follows :

Smaller, width of face of male 2.41-3.10 mm. (average approxi-
mately 2.75), of female 2.55-3.13 mm. (average approximately 2.86);
color varying, sometimes wholly black but usually ranging from
head and mesosoma black, metasoma black but for two basal
fulvous segments, to color as described for N. n. nevadensis; wings
of female hyaline-yellow to darkly infuscate; pubescence of meso-
notum, fasciae and sterna of females white to rufescent; appressed
hairs on discs of terga one to five of female absent or very sparse;
face short, clypeus not almost covering folded mandibles, average
ratio of facial length to facial width of male 1:1.17, of female 1:1.25.

This form is easily distinguished from N. n. bakeri by its smaller
size, more coarse and sparse punctation, and by the shape of the
rugose propodeal base, which is narrowly crescent-shaped. More-
over, in areas where the two might be confused, arizonensis is
usually mostly red, whereas bakeri is usually mostly black. N. n.
arizonensis is separated from N. n. stellata by the absence of ap-
pressed tergal pubescence in the female of the former, and by the
sharp proximal tooth of the male hind tibia. The black specimens
are difficult to distinguish from N. n. angelesia, their smaller size
being the best criterion.

Variation: Cockerell (1899, 1910) applied the species name
arizonensis to a series of wholly black specimens labelled "Tucson,
Arizona" or "Arizona." The same author (1925) reclassified the
form as a black subspecies of N. nevadensis. Since forms with vary-
ing amounts of red as well as wholly black forms are recorded from
the same localities over a wide area (and are, in fact usually pre-
dominant over the black forms), there seems to be little reason for
restricting the name to the black specimens alone. It might be sup-
posed that the different color forms are sympatric species, but the
evidence does not support such a conclusion.

The fact that an almost complete series of intergrades exists be-
tween black and red indicates that interbreeding probably occurs.
Moreover, all attempts to separate red from black forms by morpho-

1290 The University Science Bulletin

logical differences, by host plants visited, and by seasons of activity
have been unsuccessful. A difference of means test, using the width
of face measurement was applied to red and black females from
Tucson, Arizona, and Blythe, California. In both localities, results
showed no significant differences between the two color forms
(P>.05) indicating that in each locality, they are members of a
single population on the basis of the character tested. Further
evidence concerning the status of these color forms is found at
their nesting sites. In 1951, Dr. G. E. Bohart and the author found
black and partially red bees nesting together in a site near Delta,
Utah, and this same observation has been reported separately by
Dr. E. G. Linsley (in litt.) for a population at Blythe, California,
and by Dr. G. D. Butler (in litt.) at Roll, Arizona. On the basis of
the above facts, there is little doubt that the color forms represent
normal components of certain populations and that their taxonomic
separation is neither natural nor desirable.

N. nevadensis arizonensis, as herein defined, includes several
geographically recognizable forms. It is felt that no useful purpose
would be served by their taxonomic recognition, since these forms
grade imperceptibly into one another. The details of this geo-
graphic variation will be presented elsewhere.

Distribution: N. n. arizonensis ranges over a large area from
southeastern California to eastern New Mexico and from central
Utah to northern Jalisco. It forms a wide zone of intergradation
with populations of N. n. angelesia in southern California and also
appears to intergrade with N. n. stellata in this region. It has not
been found to intergrade with N. n. bakeri at any point along their
adjoining ranges except in western Texas (see description of N. n.
bakeri). N. n. arizonensis as defined within this paper has often
been mistaken for N. nevadensis nevadensis, with which it inter-
grades along the Utah-Nevada border, but the two are separable on
the basis of the characters described above.

Type locality: "Arizona."

Specimens examined: Arizona: Ajo, Arivaca, Arlington, Ashfork,
Benson, Bill Williams Fork (Mohave Co.), Carr Canyon (Huachuca
Mts. ), Cochise Co., Continental, Dateland, Douglas, Eloy, Fredonia,
Gila Bend, Harshaw, Marinette, McNeal, Oracle, Phoenix, Red
Rock, Roll, Roosevelt Lake, Sabino Canyon, San Francisco Mts.
(Coconino Co.), San Simon (Cochise Co.), Sentinel, Toltec,
Tombstone, Tucson, Winslow, Yuma. New Mexico: Albuquerque,
Bernalillo, Carlsbad, Columbus, Glenwood, Lagnna, La Luz, Las

Bees of the Subgenus Epinomia 1291

Cruces, Malaga, Mesilla Park, Portales, Roswell, Santa Fe, Santa
Rosa. Texas: Alpine, Cornndas (Hudspeth Co.), El Paso, Ft.
Hancock, Marathon, Salt Flat (Hudspeth Co.). Utah: Delta,
Jericho, Lvnndvl. Aguas Calientes: Rincon de Romos. Chihua-
hua: Chihuahua, Jimenez, Parral, Samalayuca, Santa Barbara, Villa
Ahumada. Coahuila: Cabos, Guadalupe. Durango: 8 mi. So.
Canutillo, Durango, Nombre de Dios, San Juan del Rio. Jalisco:
San Juan de los Lagos. Zacatecas: Sain Alto.

Total number of specimens examined: 201 males, 277 females.
The majority of these were taken between July 15 and Aug. 21.
Males have been reported as early as May 6 and females as late
as Sept. 22.

Flower records: Asclepias spp., 1 5 ; Baccharis glutenosa, 6 5 ;
Chrysothamnus spp., 1 5 ; Cleome serrulata, 1 £ ; Eriogonum spp..
1 5 ; Euphorbia spp., 1 $ , 1 5 ; Eysenhordtia pohjstachya, 2 <£ , 1 $ ;
Grindelia squarrosa, 3 $ , 40 9 ; Grindelia spp.. Guard iola tulocarpa.
5 g , 3 5 ; Gutierrezia spp., 1 5 ; Haplopappus spinulosis, 5 9 ;
Helianthus spp., 2 £ ; Hymenothrix wislizeni, 1 5 ; Hymenoxys spp.,
3 5 ; Isocoma heterophylla, 1 $ ; Lygodesmia juncea, 1 J , 2 $ ;
Pedis papposa, 3 J* , 5 $ ; Psilotrophe cooperi, 1 $ ; Solatium eleag-
nifolium, 4 $ ; Solatium spp., 1 g ; Sphaeralcea spp., 1 J ; Verbesina
spp., 2 J ; Wislizenia refracta, 1 5 .

Nomia (Epinomia) nevadensis angelesia Cockerell

(Fig. 2)

No7nia arizonensis angelesia Cockerell, 1910, Proc. U. S. Nat. Mus., vol. 38,

p. 293.
Nomia nevadensis angelesia Cockerell, 1925, Pan-Pac. Ent., vol. 1, p. 179.

Description: As described for N. nevadensis nevadensis except
as follows:

Width of face of male 2.62-3.30 mm. (average approximately
2.99), of female 2.79-3.37 (average 3.10); color wholly black, wings
honey brown, moderately infuscated; pubescence of mesonotum
long, buffy white, that of fasciae and sternal hairs also buffy white:
appressed hairs on discs of terga one to five of female absent
or very sparse; face short, clypeus not almost covering folded
mandibles, average ratio of facial width of male 1:1.17. of female
1:1.23.

This subspecies is separable from the red forms of nevadensis
arizonensis by color and by its larger size and longer mesonotal
pubescence. It is difficult to distinguish from black forms of the
latter in the more southern parts of its range, but is larger than

1292 The University Science Bulletin

arizonensis in the more northern parts of its range. It differs from
N. n. stellata by being larger, by the absence of appressed tergal
pubescence in the female, and by the presence of a sharp median
tooth on the hind tibia of the male.

Distribution: N. n. angelesia is found from southern Oregon to
the northern coast of Baja California, and east to west central
Nevada and south central California. There is a wide zone of
intergradation with arizonensis in the latter area, but no intergrades
with stellata have been found.

Type locality: Los Angeles County, California.

Specimens examined: California: Antioch, Berkeley, Blythe,
Coachella, Davis, Dos Palos, Dulzura, Firebaugh, Hagerman Park,
Helendale (San Bernardino Co.), Hemet, Hot Creek (Mono Co.),
Imperial Co., Indio, Jacumba, Kane Springs (Imperial Co.), Los
Angeles, Mammoth Lakes (Mono Co.), Mendota, Newport Beach
(Orange Co.), Oakley, Palo Verde, Rialto, Rio Vista, Riverside,
Sacramento, San Diego, San Diego Co., San Jacinto, Tulare, Tur-
lock, Vina, Yucca Valley. Nevada: Walker Lake (Mineral Co.).
Oregon: Diamond Lake (Douglas Co.). Baja California: 35
mi. south of Tijuana.

Total number of specimens examined: 205 males, 194 females.
The majority of these specimens were taken between July 10 and
Aug. 28. Throughout the range of the form, however, the flight
season is long (April 20-Oct. 26) and it seems probable that there
is commonly more than one generation per year.

Flower records: Aster spinosa, 3 9. ; Brassica spp., 2 J ; Centro-
madia pungens, 6 £ , 6 9; Chrysothamnus spp., 1 £ ; Compositae
(no further data given), 24 $ ; Croton calif ornicus, 4 J1 ; Cruciferae
(no further data given), 3 g ; Grindelia camporum, 1 5 ; Grindelia
spp., 2 $ , 10 5 ; Gutierrezia sarothrae, 7 $ , 13 9 ; Heterotheca
grandiflora, 3 9. ; Isocoma spp., 2 9. ; Lotus americana, 4 $ ; Medi-
cago sativa, 8 J ; Melilotus alba, 11 J , 2 5 ; Melilotus spp., 11 ^ ,
5 5 ; Pedis papposa, 4 j , 19 5 ; Solidago occidentalis, 1 ^ ; Soli-
dago spp., 3 $ , 22 2 ; Tamarix gallica, 1 $ , 1 $ ; Trichosterna spp.,
1 5 ; Wislizenia refracta, 1 J .

Nomia (Epinomia) nevadensis stellata, subsp. nov.

(Figs. 2; 4G)

Description: As described for N. n. nevadensis except as follows:

Male: Smaller, width of face 2.82 mm. (2.45-2.82); color wholly

black ( to head and mesosoma black, two basal metasomal segments

fulvo-ferruginous ) ; wings hyaline-brown; median excavation of hind

tibia shallow ( fig. 4, G ) , prominence formed by its proximal margin

Bees of the Subgenus Epinomia 1293

rounded rather than angulate (or excavation deeper, prominence
angulate); under surfaces of middle and hind tibiae shallowly un-
dulate (or smooth); face shorter, clypeus not almost covering folded
mandibles, average ratio of facial length to facial width 1:1.16.

Female: Smaller, width of face 2.79 mm. (2.55-2.89); head, meso-
soma, legs, and lateral portions of metasomal terga black, discs of
metasomal terga dull fulvo-ferruginous (or wholly black); wings
hyaline-brown; pubescence of mesonotum and fasciae pale buffy
white; discs of metasomal terga more densely covered with short,
plumose, appressed hairs, most prominent on terga three and four,
stiff, posteriorly directed hairs basad of fasciae inconspicuous; long
hairs of sterna two to five uniformly buffy white; posterior margin
of horizontal rugose base of propodeum carinate (or this carina
lacking); face short, clypeus not almost covering folded mandibles,
average ratio of facial length to facial width approximately 1:1.22.

N. n. stellata differs from N. n. angelesia in being smaller with
much lighter wings. It may be separated from N. n. arizonensis,
as well as from angelesia by the presence of the appressed pubes-
cence on the metasomal terga, the carinate posterior margin of the
propodeal base in the female, and by the absence of the sharp
median tooth on the hind tibia of the male (fig. 4, G).

Variation: Due, perhaps, to its occurrence on a narrow peninsula,
this form shows a wide range of intrapopulational variation. Males
overlap in every key character with those of arizonensis and are only
partially separable from that form. Females are distinct from those
of surrounding subspecies. Moreover, marked differences in color
occur between the sexes, males usually being wholly black while
females often possess one or more red terga.

Since the males can be referred to adjoining forms, the author
has preferred to designate this form as a subspecies, being aware
that later research may prove it to be a distinct species.

Distribution: Middle Baja California north to the southern por-
tions of California. Male specimens from Imperial Co., California,
were examined which are believed to be intergrades between this
form and N. n. arizonensis. No evidence of intergradation with
N. n. angelesia in northwestern Baja California has been found.

Type material: Holotype male, allotype female, ten male and ten
female paratypes from San Pedro, Baja California, Oct. 4, 1941
(E. S. Ross, G. E. Bohart).

Type respositories: Holotype, allotype, four male and four fe-
male paratypes in the California Academy of Sciences. One male
and one female paratype in the Snow Entomological Museum.

1294 The University Science Bulletin

Specimens examined: Baja California: 15 mi. So. San Domingo,
Oct. 4, 1941 (E. S. Ross, G. E. Bohart), 2 $ ■ San Pedro, Oct. 7,
1941 (E. S. Ross, G. E. Bohart), 13 $ , 15 $ , plus type material.

Nomia (Epinomia) nevadensis bakeri Cockerell

(Figs. 2; 3M)

Nomia bakeri Cockerell, 1898, Entomologist, vol. 31, p. 32.

Nomia pattoni Cockerell, 1910, Proc. U. S. Nat. Mus., vol. 38, p. 292.

Nomia nevadensis pattoni, Cockerell and Blair, 1930, Amer. Mus. Novitates,

no. 433, pp. 14, 16.
Nomia bakeri var. rufibasis Cockerell, 1930. In Cockerell and Blair, Amer.

Mus. Novitates, no. 433, p. 14.
Nomia nevadensis bakeri, Cockerell and Blair, 1930, Amer. Mus. Novitates, no.

433, pp. 7, 14, 16.

Description: As described for N. n. nevadensis except as follows:

Male: Width of face 2.41-3.13 mm. ( average approximately 2.99
mm.); color varying, rarely as described for N. n. nevadensis, usually
ranging from wholly black to head and mesosoma black, metasoma
black but for two ferruginous basal segments; apex of hind femur
usually black, remaining segments of leg fulvous; wings honey
brown to moderately infuscated; punctures of vertex finer and
closer than those of clypeus, punctation finer and closer throughout
than that described for N. n. nevadensis; horizontal rugose base of
propodeum widely crescent shaped or appearing truncate pos-
teriorly; face shorter than that described for nevadensis, clypeus
not almost covering folded mandibles, average ratio of facial length
to facial width, 1:1.17.

Female: Width of face 2.75-3.30 mm. (average approximately
3.06 mm. ) ; color as described for male; wings moderately to heavily
infuscated; pubescence of mesonotum white to buffy white; hairs
of metasomal sterna buffy white; appressed hairs on discs of terga
one to five absent, stiff, posteriorly directed hairs basad of fasciae
inconspicuous; punctures of first and second metasomal terga
nearly alike, punctation finer and closer throughout than that
described for N. n. nevadensis; rugose propodeal base as described
above for male; face short, clypeus not almost covering mandibles,
average ratio of facial length to facial width 1:1.28.

This subspecies is easily distinguishable from N. n. arizonensis
by its larger size, finer and closer punctation, and by the widely
crescentic or truncate configeration of the rugose propodeal base.
In areas where the two meet, bakeri is usually wholly or largely
black while arizonensis usually possesses at least two fulvous meta-
somal segments.

Variation: N. n. bakeri is usually readily separable from all other
subspecies of N. nevadensis on the basis of the characters given

Bees of the Subgenus Epinomia 1295

above, and was considered as a distinct species until Cockerell and
Blair (1930) placed it under N. nevadensis because of similarities
in the male genitalia of the two forms.

Despite the differences mentioned in the description, the genitalia
and hind tibiae of the males agree with those of N. n. arizonensis,
which it neighbors to the east. These two characters have been
especially useful in separating the species of Epinomia. Moreover,
neither the size nor color differences between baked and other
subspecies of nevadensis are extraordinary. The largest bakeri are
no larger than northern California specimens of N. n. angelesia,
and in the southern part of its range, fulvous bakeri are found
which closely resemble other red forms of the species.

As stated previously, N. n. bakeri geographically replaces ari-
zonensis to the east. However, the status of these bees where their
ranges meet is obscure and requires further investigation. Cockerell
(1930) thought of bakeri only as a plains bee and suggested that
it might intergrade with arizonensis in southeastern Colorado.
Because specimens from this area were not available for purposes of
the present paper, the truth of this surmise is still untested. A
single typical specimen of bakeri from each of two localities in
western Colorado (Durango and Leadville) indicates, however,
that this form is established west of the Rocky Mountains. Since
typical arizonensis is known from Albuquerque, if intergradation
does take place in this area, it must occur between the latter
locality and Durango. Along the eastern border of New Mexico,
there is evidence suggesting that the two ranges overlap. A single
small male bakeri from Tucumcari, New Mexico, has been examined,
and a single typical male arizonensis is known from Portales, New
Mexico, southwest of Tucumcari.

From Barstow, Ward County, Texas, however, comes a series of
fulvous bees which seem to be intergrades between the two forms.
On the basis of this evidence, bakeri is placed as a subspecies of
nevadensis. It is clear, however, that more study and material is
needed before the status of these forms is settled.

Distribution: Northern Florida to south central Wyoming, the
eastern two thirds of Colorado, and the extreme northern and
eastern portions of New Mexico south to near Barstow, Texas,
thence southeast to the gulf coast near Matamoros, Tamaulipas.

Type locality: "Colorado."

Specimens examined: Alabama: Selma. Colorado: Berkeley,
Boulder, Canfield, Crook, Crowley, Denver, Durango, Ft. Lupton,
La Junta, Lamar, Leadville, Limon, Logan Co., Roggen, Sterling,

1296 The University Science Bulletin

White Rock, Valmont. Florida: Flagler Beach. Kansas: Douglas
Co., Garden City, Hutchinson, Lakin, Newton, Pawnee Co., Pierce-
ville, St. John, Stafford. Louisiana: Tallulah. Mississippi: Hat-
tiesburg. Nebraska: Mitchell. New Mexico: Tucumcari. Texas:
Barstow, Bishop, Conlen, Cotulla, Dalhart, Dilley. Wyoming:
Guernsey, Lingle. Tamaulipas: Matamoros.

Total number of specimens examined: 110 males, 232 females.
The majority of these were taken between July 21 and Aug. 30.
Males have been taken as early as April 17 and females as late as
Oct. 15. There are probably several generations per year in the
southern portions of the range.

Flower Records: Bidens invohicrata, 3 J ; Boltonia asteroides,
2 £ ; Chrysopsis microcephalia, 2 £ ; Chrysopsis spp., 1 $ ; Coreopsis
spp., 3 $ , 1 5 ; Dalea lanata, 2 $ ; Grindelia spp., 5 J1 ; Heterotheca
spp., 2 $ , 42 $ ; Isopappus sp., 2 J , 15 $ ; Melilotus alba, 8^,1$;
Prionopsis spp., 2 $ ; Prosopis spp., 2 5 ; Solidago spp., 3 $ ; Thele-
sperma gracile, 1 J ; Verbesina spp., 1 ^ .

LITERATURE CITED

ASHMEAD, W. H.

1899. Classification of the bees of the superfamily Apoidea. Trans. Amer.
Ent. Soc., vol. 26, pp. 49-100.

BOHART, G. E.

1952. Pollination by native insects. In Insects, Yearbook of Agriculture,
pp. 107-121.

COCKERELL, T. D. A.

1899. Notes on American bees. Entomologist, vol. 32, pp. 128-129.
1910. The North American bees of the genus Norma. Proc. U. S. Nat.

Mus., vol. 38, pp. 289-298.
1925. Some bees collected by Mr. P. H. Timberlake at Riverside, Cali-
fornia. Pan-Pac. Ent., vol. 1, pp. 179-180.
1934. Records of western bees. Amer. Mus. Novitates, no. 697, pp. 1-15.
Cockerell, T. D. A., and B. H. Blair.

1930. Rocky Mountain bees I. Amer. Mus. Novitates, no. 433, pp. 1-19.
Cross, E. A., and G. E. Bohart.

Observations on the biologies of some solitary bees of the genus
Nomia ( Hymenoptera-Halictidae ) . In press.
Dalla Torre, Karl Wilhelm von.

1896. Catalogus Hymenopterorum; hucusque descriptorum systematicus
et synonymicus. Leipzig. Vol. 10, viii -f 643 pp.
Hemming, F.

1953. Copenhagen decisions on zoological nomenclature. International
Trust for Zoological Nomenclature, London, 135 pp.

Latreille, P. A.

1805. Histoire naturelle, generale et particuliere des Crustaces et des In-
sectes, vol. 13. Dufart, Paris, 432 pp., tab. 98-103.

Bees of the Subgenus Epinomia 1297

Michener, Charles D.

1944. Comparative external morphology, phylogeny, and a classification

of the bees ( Hymenoptera ) . Bull. Amer. Mus. Nat. Hist., vol. 82,

art. 6, pp. 151-326, 13 diagrams, 246 figs.

1947. A character analysis of a solitary bee, Hoplitis albifrons (Hymen-

optera-Megachilidae). Evolution, vol. 1, no. 3, pp. 172-185, 8 figs.

Museheck, C. F. W., K. V. Krombein, and H. K. Townes.

1951. Hymenoptera of America north of Mexico. Synoptic catalog.
U. S. D. A. Agricultural Monograph No. 2, 1420 pp.
Pierce, W. Dwight.

1904. Some hypermetamorphic beetles and their hymenopterous hosts.
Nebr. Univ. Studies, vol. 4, pp. 153-189, 2 pi.
Provancher, L'Abbe L.

1889. Additions et corrections au vol. II, de la faune entomologique du
Canada, traitant des Hymenopteres, 475 pp.
Rau, Phil.

1929. The nesting habits of the burrowing bee, Epinomia triangulifera
Vachal. Psyche, vol. 36, pp. 243-248, 1 pi.
Smith, R. F.

1954. The importance of the microenvironment in insect ecology. Jour.
Econ. Ent, vol. 47, no. 2, pp. 205-209.
Yarrow, H. C.

1875. In Wheeler, G. M. Report upon geographical and geological ex-
plorations west of the 100th meridian. Vol. 5, Zoology, pp. 509-
1021, pi. 16-45.

1298 The University Science Bulletin

A-D. Genital capsules of Epi7iomia, ventral aspect.

A. N. triangulifera (Amazonia, Mo.); B. N. micheneri (Can Canyon, Ariz.,
paratype); C. IV. nevadensis nevadensis (Fairbanks Springs, Nev. ); D. N.
boharti (paratype).

E.-N. Clypeal margins of Epinomia as seen from beneath.

E. N. triangulifera, male (Douglas Co., Kans. ); F. IV. micheneri (holo-
type); G. N. micheneri (Carr Canyon, Ariz.), (male paratype); H. N. nevaden-
sis nevadensis, male (Fairbanks Springs, Nev.); I. IV. boharti (male paratype);
J. N. triangulifera, female (Lawrence, Kans.); K. N. micheneri (allotype);
L. N. nevadensis arizonensis, female (Delta, Utah); M. IV. nevadensis bakeri,
female (La Junta, Colo.); N. boharti, female (San Bias, Nayarit).

Bees of the Subgenus Epinomia J 299

Ffg. 3

1300 The University Science Bulletin

A-D. Sterna five of male Epinomia.

A. N. triangulifera (Lawrence, Kans. ); B. N. micheneri (Sta. Barbara,
Chihuahua); C. N. nevadensis nevadensis (Fairbanks Springs,, Nev. ); D. N.
boharti (paratype).

E-I. Right hind tibiae of male Epinomia.

E. N. triangulifera (Promontory, Utah); F. IV. micheneri (holotype); G. N.
nevadensis stellata (paratype); H. N. nevadensis nevadensis (lectotype); I.
N. boharti (paratype).

Bees of the Subgenus Epinomia

1301

Fig. 4

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 17

The Status of the Mexican Lizards of the Genus Mabuya

BY

(

Robert G. Webb *

Abstract. The Mexican lizards of the genus Mabuya are examined criti-
cally. Material consisting of 75 specimens from seven Mexican states was
available. A study of the variation from the various localities was made and
it was found that the populations were for the most part homogeneous and
no subspecific variants were recognized. It was determined that the proper
name for the Mexican species is Mabuya brachypoda Taylor.

INTRODUCTION

The conservatism of Dunn (1936), who considered Mabuya
mabouya mabouya as ranging from Mexico to Ecuador, Rrazil and
the Lesser Antilles has been questioned. Burger (1952) revived
the name alliacea of Cope (1876) and regarded the mabuyas rang-
ing from Mexico to Costa Rica as Mabuya mabouya alliacea. In
addition, Taylor ( 1957 ) resolved Costa Rican material into three
distinct species, Mabuya unimarginata, Mabuya brachypoda, and
Mabuya alliacea. Dunn, himself, (loc. cit. : 545) in his concept
of M. mabouya mabouya, recognized that the populations were
". . . numerous and quite diverse, . . ." This variation in
populations of the mainland coupled with an insular type locality
(restricted to St. Vincent by Smith and Taylor, 1950:156) for M.
mabouya mabouya suggests the presence of mainland populations
worthy of taxonomic recognition.

In view of this variation, Dr. Edward H. Taylor suggested to
the author that a study of the variation exhibited by the genus
Mabuya in Mexico be made in an effort to clarify the taxonomic
relationships of the northern mainland populations. The author
is grateful to Doctor Taylor for permission to study specimens in

* Department of Biology, West Texas State College, Canyon, Texas.

(1303)

1304 The University Science Bulletin '

his private collection and for constructive criticism of the manu-
script.

The present study is based on an examination of 75 specimens
from seven Mexican states which were collected intermittently in
the years 1932 to 1940. All numbers are those of the EHT-HMS
collection (private collection of E. H. Taylor), and are listed by
locality below.

Campeche.—Balchaca], 14435-36, 14438-39, 14450-51, 14457; Tres
Brazos, 14441-45, 14447-48; Encarnacion, 14453-54, 14456 ( 17 speci-
mens ) .

Chiapas. — San Ricardo, 10504; Libertad, 15425; Tuxtla Gutierrez,
15343 (3 specimens).

Guerrero. — 12 mi. S. Chilpancingo, 10453; 7 mi. E. Chilpancingo,
22237-38, 23915; near Chilpancingo, 23885; 1 mi. N. Organos, 10454;
El Limoncito, near Acapulco, 19094-95, 19097-98, 19100-01, 19103-
04, 19106-07; between Chilpancingo and Omilteme, 23882-83; Agua
del Obispo, 10456-57, 23907-08 ( 22 specimens ) .

Michoacdn.— Uruapan, 10460, 10462-63, 10465-66, 10468-69,
10471-72, 10474-75, 10477-78, 10480-81, 10483-84, 10486-87, 10489-
90, 10492-93, 10495-96, 10498-99, 10501-02 (29 specimens).

Morelos. — Puente de Ixtla, 19092 ( 1 specimen ) .

Oaxaca. — Totalapan, 10459 ( 1 specimen ) .

Yucatan. — Progreso, 14432; Merida, 14433 (2 specimens).

VARIATION

Body proportions. — The following measurements were utilized:

Snout-vent length — from the tip of the snout to the anterior mar-
gin of the anus.

Tail length — from the anterior margin of the anus to the tip of
the tail; only measurements of complete tails were taken.

Head length — from tip of snout to anterior margin of ear along
a line parallel to the lateral axis of the head.

Head width — the greatest width in a transverse straight line.

Leg length — from the tip of the longest toe (including the claw)
to the angle with the body (posterior to fore limb and anterior to
hind limb), with the leg extended at right angles to the body.

Axilla-groin distance — a straight line from the posterior margin of
insertion of the fore limb to the anterior margin of the insertion of
of the hind limb. -

Also, ratios from six body proportions were calculated; these were
snout-vent/axilla-groin, snout- vent/f ore limb, snout-vent/hind limb,

Mexican Lizards of Genus Mabuya 1305

snout-vent/head length, snout-vent/head width, and axilla-groin/
hind limb.

In an attempt to determine ontogenetic variation, individuals
were assigned to three arbitrary size groups as follows: 1) 20-40
mm., 2) 41-60 mm., 3) 61-83 mm.

The largest sample available from a single locality is 29 speci-
mens from Uruapan, Michoacan. A consideration of geographic
variation involved a comparison of the Michoacan sample with 22
specimens from Guerrero and 19 from Campeche.

The construction of scattergrams and calculation of mean and ex-
treme ratios revealed little sexual diamorphism, ontogenetic growth,
or geographic variation in body measurements. Smaller individuals
have larger heads. In the 20-40 mm. size-group the head length is
contained in the snout- vent length an average of 4.2 times ( 13 speci-
mens), whereas in the 61-83 mm. size-group it is contained 5.3
times (38 specimens). Similarly, the snout-vent/head width aver-
age ratio for 13 specimens in the 20-40 mm. size-group averages
6.2, whereas the same for 38 specimens in the 61-83 mm. size-group
averages 7.4.

The hind limb is longer in the smaller individuals in relation to
the axilla-groin length. The average axilla-groin/hind leg ratio
in the 20-40 mm. size-group was 1.4, whereas the same in the 61-83
mm. size-group was 1.7. The length of the fore limb in relation to
the axilla-groin length was not considered. It is of interest that no
ontogenetic growth in the hind limb is revealed if a comparison of
the snout-vent/hind limb ratios of the three size-groups is made, as
an average ratio of 3.1 is maintained; this is probably reflected in
the differential growth of the head.

In regard to tail length, seven juveniles ranging in snout-vent
length from 26 to 31 mm. ( sex undetermined ) have an average tail
length/snout-vent length ratio of 1.3 (1.2-1.4). The analysis of
variation in tail length of adults was limited to five specimens (2
males, 3 females) in which the tail length/snout-vent ratios in
males (1.9 and 2.0) are larger than that in females (1.7, 1.7, and
1.7). A male, 64 mm. in snout-vent length, had the longest tail,
( 130 mm. ) ; all females exceeded 64 mm. in snout-vent length. The
meager data indicate that adult males have longer tails in relation
to snout-vent length than do adult females.

Females attain a larger size than males and have an average
snout-vent length of 71.2 mm. (extremes 62-83 mm.) based on 19
specimens, whereas males have an average snout-vent length of

1306

The University Science Bulletin

85-1

80-

75-

70-

hS-

60-

5£

SNOUT- VENT LENGTH

KilDDORSf\L SCALES

2853

2-299

2&SS

■fc5

-60

-5S

■SO

L45

Fig. 1. Variation between sexes in snout-vent length and number of mid-
dorsal scales. The horizontal line indicates the mean; the vertical line, the
observed variation; the white rectangle, four standard deviations; the black
rectangle, four standard errors of the mean.

63.0 mm. ( extremes 56-70 mm. ) based on 28 specimens ( Fig. 1 ) .
Data used are from specimens regarded as sexually mature (see
page 1312).

Scutellation. — The dorsal body scales of all specimens are smooth
with no evidence of keels.

Middorsal scales: A count of the dorsal scales from the mo^t
anterior body scale (excluding the nuchal), in a straight line at
or near the middorsal line as far back as a line even with the
posterior margins of the thighs, when the hind legs are held at
right angles to the body. There is a distinct sexual dimorphism
in the number of middorsal scales ( Fig. 1 ) . Males averaged 53.9
(extremes 49-58), based on 28 individuals, whereas females aver-
aged 57.6 (extremes 55-60), based on 22 individuals.

Mid-body scales: A transverse count of the longitudinal scalerows
at a point one fourth the axilla-groin length anterior to the insertion
of the hind leg. Counts taken at mid-body usually include inter-
calary rows which extend posteriorly from the axillary region.
These scales varied in number from 28 to 32. A count of 28 scale-
rows (51%) was recorded for 29 of 57 specimens; thirty scalerows

Mexican Lizards ok Genus Mabuya 1307

occurred in 24 counts (42%) and 32 scalerows in 4 counts (7%).
The count of 32 scalerows was restricted to females.

Supralabials: The scales bordering the upper edge of the mouth
except at the tip of the snout where the rostral occurs. The largest
posterior scale is regarded as the last supralabial. Of 73 speci-
mens in which both sides could be counted, 51 (70%) had a combi-
nation of 6-6, 15 (20%) had a combination of 6-7, 5 (7%) had a
combination of 5-6, and 2 specimens (3%) had a combination of
7-7. Considering each side separately and based on 148 counts,
6 supralabials occurred in 124 counts (84%), 7 occurred in 19
counts (13%), and 5 occurred in 5 counts (3%). When a count of
6-6 occurs, the fifth supralabial is the largest and lies immediately
beneath the orbit, with one labial posterior and four anterior. In
the case of seven supralabials, the sixth is the largest and beneath
the orbit, and with five supralabials, the fourth is the largest and
beneath the orbit. Five supralabials occurred only in certain
specimens from Guerrero.

Infralabials: The scales bordering the lower edge of the mouth
except for the most anterior mental scale. Of 71 specimens in
which both sides could be counted, 59 specimens (84%) had a
combination of 6-6, 7(9%) had a combination of 6-7, 3(4%) had
a combination of 7-7, and 2(2%) had a combination of 5-6. Con-
sidering each side separately and based on 146 counts, six infra-
labials occurred in 129 counts (89%), 7 occurred in 15 counts (10%),
and 5 occurred in 2 counts (1%). All specimens with five infra-
labials are from Guerrero.

Postmentah A scale following the mental along the midventral
line. The postmental was unpaired in 72 specimens and asymmetri-
cally divided in another.

Nuchals: The enlarged paired scales immediately posterior to
the parietals. Of 72 specimens, all had one pair of nuchals; the
two nuchals were fused medially in one specimen from Yucatan.

Parietals: Paired scales posterior to the frontoparietals and bor-
dering the interparietal on either side. Of 73 specimens, all had
the parietals in contact posteriorly.

Supraoculars: The scales above the orbits. A count of 4-4 oc-
curred in 68 specimens (95%), two specimens had a count of 3-4
(3$), and one specimen had a count of 3-3 (2%). Considering each
side separately and based on 146 counts, four supraoculars occurred
in 141 counts (97%) and three supraoculars occurred in 5 counts
(3%). The most anterior supraocular is the smallest, the one
immediately posterior to it is the largest.

1308 The University Science Bulletin

Prefrontals: Paired scales immediately preceding the frontal.
The prefrontals were separated in 67 specimens (92%) and in con-
tact in six specimens ( 8% ) .

Supranasals: Paired scales immediately preceding the frontonasal
and posterior to the rostral. Of 75 specimens, 55 (73%) had the
snpranasals in contact medially, whereas they were separated in
20 specimens (27%). Of possible geographic interest is the fact
that none of the 28 specimens from Michoacan had the supranasals
separated. Dunn {Joe. cit. :540) reports contact of the supranasals
in only 33% of the specimens he examined from Mexico. Burger's
comments (loc. cit. :187) concerning the variation of this character
in Mexico closely parallels the results of this study.

Chin shields: A paired series of scales extending posteriorly from
the postmental. From a total of 73 specimens, 52 (71%) had a
combination of 2-2, 10 (14%) had a combination of 1-2, 8 (11%)
had a combination of 1-1, 2(3%) had a combination of 2-3, and 1
specimen (1%) had a combination of 3-3.

Coloration. — The following comments pertain to all sizes of both
sexes, there being little or no sexual or ontogenetic variation in
coloration. Dorsally the ground color ranges from a dark olive
to a light burnt-brown and may be uniform or marked with small
dark spots. The venter, throat, and undersurface of the limbs
are light and without markings. The anterior, posterior, and dorsal
surfaces of the limbs have a marbled or reticulate appearance. A
lateral white stripe extends from the angle of the mouth posteriorly
through the lower half of the tympanic cavity and just above the
insertion of the fore limb to terminate just above the insertion of
the hind limb or just posterior to it on the tail. This bilateral white
stripe is about one scale wide and, at mid-body, usually (73%)
overlaps two longitudinal scalerows, the ventral half of the
seventh and dorsal half of the eighth scalerows on the right side,
and the ventral half of the sixth and dorsal half of the seventh
scalerows on the left side (excluding the middorsal scalerow).
Occasionally (27%), the lateral white stripe engages only one scale
row on each side, the seventh on the right and the sixth on the left.
Immediately dorsal to the lateral white stripe is a longitudinal dark
brown band on either side which is two (rarely one and one-half
or two and one-half) scalerows wide; it begins anteriorly on the
lateral head scales and extends posteriorly through the eye, the
upper half of the tympanic cavity and terminates posteriorly at the
posterior insertion of the hind limb. Ventral to the lateral white

Mexican Lizards of Genus Mabuya 1309

stripe on both sides is a dark band varying in width, which merges
with the immaculate venter. Adjacent and dorsal to the lateral
brown band on either side is a narrow light line which may be
indistinct.

Geographic variation and sexual differences were noted in the
extent of markings on the dorsum. In males from Guerrero, Oaxaca
and Morelos, the dorsum (area between lateral dark brown bands)
was uniform without markings (19092, 19098, 10459), or with only
a few small scattered black spots (19095, 22238, 23882). One
specimen, however, (10457) had considerable dark marking. In
males from Michoacan, the dorsum is more consistently marked
with small black marks which in some tend to form short narrow
longitudinal stripes (10465, 10483, 10498); others have a more
or less uniform dorsum (10466, 10492-93). The tendency to form
longitudinal dark stripes by fusion of the small black spots is in-
creased in the more eastern states of Campeche, Chiapas and
Yucatan, and is apparent in three males (14456, 14442, 14445).
Another (14432) has a spotted dorsum but no evident stripes. All
other males have some markings, though sparse. Females have
for the most part a uniform dorsum and do not exhibit as strong
a geographic trend as do males. Guerrero and Michoacan females
may have a uniform dorsum or one with scattered dark marks.
One female from Michoacan (22237) is strongly marked but there
is no tendency for longitudinal stripes. Of 11 females from Chiapas,
all have a uniform dorsum, or nearly so, except three specimens,
one of which has narrow longitudinal stripes (14439), and two
(14435, 15343) which tend toward the formation of longitudinal
stripes.

One of Burger's criteria (loc. cit. :186) for recognizing the popu-
lation occurring from Mexico to Costa Rica is based on the narrow
dorsolateral dark stripe which is two or less scales in width rather
than two and one-half or three scales. The dorsolateral dark stripes
are the same as the lateral bands referred to in this paper, which
are two scales wide.

COMPARISONS

The current appellation for Mexican mabuyas is Mabuya
mabouya alliacea. Taylor ( loc. cit. ) dealing with the Costa Rican
Mabuyas re-elevated alliacea to specific rank but made no mention
that the binomial should be applied to Mexican representatives of
the genus; his redefinition of Mabuya alliacea affords a basis for
comparison with the Mexican population. The presence of an

1310 The University Science Bulletin

azygous scale between the supranasals and the unequal, trans-
verse division of each parietal scale resulting in a separation of
the parietal scales, as depicted by Cope (loc. cit. :pl. 6, fig. 1) in
the illustration accompanying the type description, is regarded as
an anomaly as neither configuration has been encountered by Taylor
or myself. The other characters given by Cope in the type descrip-
tion are either in agreement with or encompassed by the variation
in the data presented by Taylor.

In regard to the enlarged preanal scales described for the types
of alliacea, the author agrees with the comments of Dunn ( loc. cit. :
542 ) who regarded the character of no useful taxonomic significance.
Taylor calls attention to an axillary pit or pocket behind the inser-
tion of the arm. These had been overlooked by previous workers.
In all the Mexican specimens examined they were well developed.

Using, then, the data given by Taylor, selected characters of
Mabuya alliacea from Costa Rica are compared with those ex-
hibited by the specimens from Mexico ( Table 1 ) . The range of
variation in the number of mid-body scales is slightly lower in
alliacea though an average of 28 scales is common to both. Mabuya
alliacea differs from the Mexican specimens in that 1) the sixth
supralabial subtends the orbit, 2) the rostral normally touches
the frontonasal, 3) a dorsolateral dark stripe is present, and 4)

Table 1. — Comparison of Characters of Mabuya alliacea, Mabuya brachypoda

and specimens from Mexico.

alliacea
18

midbody scales 26-29

28 (59%)

supralabial below eye 6th (84%)

supranasals in contact no ( 82% )

dorsolateral dark stripe yes

axillary pit reduced

Mexican

brachypoda
12

specimens

75

28-32
30 (66%)

28-32
28 (51%)

5th (92%)

5th (84%)

yes (50%)

yes (73%)

no

no

well defined

well defined

axillary pits are reduced and not prominent. The characters men-
tioned above from which the Mexican specimens differ from alliacea
are in agreement with those mentioned by Taylor for Mabuya
brachypoda (Table 1).

Mabuya brachypoda was described by Taylor as having limbs
which, when adpressed, fail to overlap by a distance equal to or
greater than the length of the hand. The limbs of the 12 speci-

Mexican Lizards of Genus Mabuya 1311

mens which formed the type series were examined by the author.
The separation of the adpressed limbs in Nos. 31312 and 31313
was about 5 mm. However in No. 34326 the limbs were found
to overlap when adpressed. Doctor Taylor concurs with this
measurement. This latter specimen has the parietals in contact
and the sixth supralabial is below the eye, two characters which
do not occur in the type or other paratypes. This is definitely aber-
rant or through some metathesis of data the locality is incorrect and
the specimen may not belong where placed by Taylor.

The distance between adpressed limbs of the 12 M. brachypoda,
as observed by the author, varied from approximately 10 mm. in the
type specimen to 3-5 mm. Such variation could be due to the
differential bending of the individual limbs, and a distance of 4-5
mm. between adpressed limbs need not be regarded as diagnostic.
This is borne out by the fact that the limbs of No. 36313 do not
overlap when adpressed, yet the sum of the fore and hind limb
measurements exceeds the axilla-groin length by 5.5 mm. In other
words, the length of limbs is not necessarily reflected in the degree
of separation or overlap of adpressed limbs.

Although the limbs of every Mexican specimen were not ad-
pressed, casual observation revealed either slight overlap or sepa-
ration of the toes and fingers. The sum of the limb lengths ex-
ceeded the axilla-groin length in all but nine of the Mexican speci-
mens, and in only four of 12 M. brachypoda. Axilla-groin /hind leg
ratios of 10 specimens exceeding 64 mm. in snout-vent length from
Costa Rica averaged 1.9, whereas that of 38 individuals from Mex-
ico exceeding 61 mm. averaged 1.7. Although slightly shorter hind
limbs in the Costa Rican specimens are suggested, the data are in-
conclusive, and the author, at present, regards the difference in
limb length between the two populations negligible.

There are, therefore, no characters known which will certainly
distinguish the Mexican mabuyas and M. brachypoda as described
by Taylor. It is proposed that the binomial, Mabuya brachypoda,
apply to all the mabuyas occurring from Mexico to Costa Rica.
Pertinent data given in accounts by Mertens (1952:58, table 11,
fig. 59) and Werler and Smith (1952:563, fig. 14) do not reveal any
significant deviations from the variation as now known. The
morphological geographic variation in this species does not seem
to be great and is indicative that the species is a recent immigrant
to Mexico from the south. M. brachypoda is apparently a lowland
form, and in Mexico probably does not occur above an elevation of
3000 feet.

1312 The University Science Bulletin

BREEDING BEHAVIOR

A macroscopic examination of the gonads and associated ducts
of the Mexican specimens was undertaken in an effort to determine
the size at sexual maturity and other aspects of the reproductive
cycle.

Females. — Specimens were regarded as sexually mature if em-
bryos were present in the oviducts. Ten specimens, ranging in
snout-vent length from 62 to 83 mm. contained embryos and were
collected between June 26 (10453) and July 28 (10477); embryo
complements varied from four to six. Ovaries of these individuals
were 4 mm. in length with the largest ovocytes about 1 mm. in diam-
eter. Four specimens collected within the interim July 21-28 and
exceeding 62 mm. in length, did not have embryos in the swollen
and convoluted oviducts. The ovaries measured 4-5 mm. in length
with the largest ovocytes 1 mm. in diameter; recent expulsion of
embryos is suggested. Five specimens collected in September and
exceeding 62 mm. in length agree with the four above-mentioned
individuals except that developing ovocytes are larger ( about 2 mm.
in diameter) and more yolk is present. The oviducts of one speci-
men (14447), collected in September, contained five (two in the
right, three in the left) small structures interpreted as unfertilized
eggs, which had a diameter of 2 mm. A specimen (23915), 60 mm.
in length (collection date unknown) had ovaries 2 mm. in length
with one ovocyte 1 mm. in diameter; the oviducts were 1 mm. wide
and slightly wrinkled, but contained two similar-sized structures
(as in 14447) in the right oviduct. Two of these same structures
were present in the convoluted oviducts of a specimen 56 mm. in
length (14457) collected in October. Five specimens smaller than
50 mm. had ovaries measuring 4 mm. in length with ovocytes 1 mm.
in diameter, but the oviducts were straight and narrow.

The data indicate that deposition of young, which varies from
four to six, occurs in June and July; of 14 adult females collected
from June 26 to July 28, ten contained well-developed embryos.
Five adult females collected in September contained no embryos
but larger developing ovocytes than those collected in June. The
smallest female definitely considered sexually mature is 62 mm. in
snout-vent length, although a length of 56 mm. is suggested.

Males. — Seventeen specimens ranging from 58 to 70 mm. in
length, and collected in June and July had testes 5 to 8 mm. in
length and 3 to 5 mm. in width. Ten specimens ranging from 59
to 70 mm. in length, and collected in August and September, had

Mexican Lizards of Genus Mabuya 1313

smaller testes with dimensions of 4 to 6 mm. and 3 to 4 mm. Thus
there is only a slight seasonal decrease in testes size. The smallest
male examined and regarded as sexually mature (22238) is 56 mm.
in length (no collection date), and has testes measuring 5 mm. in
length and 3 mm. in width. One specimen (14456), 58 mm. in
length and considered immature, was collected in October and had
a testis measuring 3 mm. in length and 1 mm. in width; another in-
dividual ( 14444 ) collected in October and 51 mm. in length also
had testes 3 mm. in length. A specimen (19106), considered sex-
ually mature and 58 mm. in length was collected in July and had a
testis 5 mm. in length and 3 mm. in width; this inconsistency in the
relation of testis size to snout-vent length is suggestive of the ap-
proximate size at sexual maturity. This interpretation is question-
able if a seasonal decrease in testis size is more marked in October.
The size at sexual maturity is thought to be approximately 56 mm.

LITERATURE CITED

Buhger, W. Leslie.

1952. Notes on the Latin American Skink, Mabuya mahouya. Copeia,
1952, No. 3: pp. 185-187.
Cope, Edward D.

1875. On the Batrachia and Reptilia of Costa Rica. Journ. Acad. Nat.
Sci. Philadelphia, ser. 2, vol. 8, 1876 (1875) pp. 93-154, 6 pis.
Dunn, Emmet R.

1936. Notes on American Mabuyas. Proc. Acad. Nat. Sci. Philadelphia,
vol. 87, pp. 533-557.
Mertens, Robert.

1952. Die Amphibien und Reptilien von El Salvador. Abh. Senckenb.
Naturf. Ges., No. 487, pp. 1-83, 1 map, 92 figs.
Smith, Hobakt M., and Edward H. Taylor.

1950. An Annotated Checklist and Key to the Reptiles of Mexico Exclu-
sive of the Snakes. Bull. U. S. Nat. Mus., No. 199, pp. i-vi, 1-253.
Taylor, Edward H.

1956. A Review of the Lizards of Costa Rica. Univ. Kansas Sci. Bull.,
vol. 38, pt. 1, pp. 3-322, figs. 1-76.
Werler, J. E., and Hobart M. Smith.

1952. Notes on a Collection of Reptiles and Amphibians from Mexico,
1951-1952. Texas Journ. Sci., vol. 4(4): pp. 551-573, 23 figs.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 195S [No. 18

The Integumental Anatomy of the Monarch Butterfly
Danaus plexippus L. (Lepidoptera: Danaiidae)*

BY

Paul R. Ehrlich **

Abstract: This paper is the first section of a work on the integumental
morphology, phylogeny and classification of the butterflies. The external and
internal integumental anatomy of the common monarch butterfly is described
and figured in detail and a descriptive terminology adopted which will be
employed in the comparative work to follow.

INTRODUCTION

This is the first section of a work on the integumental morphology,
phylogeny and classification of the butterflies ( Papilionoidea ) .
Despite the great popular interest in this group of insects they have
been the subject of relatively little modern systematic work above
the level of the generic revision. The interrelationships of the major
groups are poorly understood, and some recent workers ( e. g., Clark,
1948) have given family rank to such obviously nonequivalent en-
tities as the papilionids and the argynnids.

It seems apparent that a mere rearranging of the butterflies on
the basis of well-studied characters such as wing venation or color
pattern, or the introduction of a new phyletic arrangement and
classification on the basis of one or two previously unstudied char-
acters, would be of little significance. Therefore an attempt will
be made to reconstruct the phylogeny of the group and arrive at a
reasonable classification by utilizing as much published work as
possible in conjunction with a study of the comparative integu-
mental morphology. Unfortunately practical considerations will
not permit simultaneous study of the characters existing in the im-

* Contribution no. 949 from Department of Entomology, University of Kansas, Law-
rence.

** Research Associate, Chicago Academy of Sciences, 2001 N. Clark St., Chicago 14,
Illinois.

(1315)

1316 The University Science Bulletin

mature forms, the visceral anatomy, the fine details of the micro-
scopic anatomy, etc., nor of all the species or even all genera. How-
ever, it is hoped that sufficient characters and an adequate diversity
of species can be studied so that major errors will be avoided.

This first section forms the basis for the comparative morphologi-
cal study which is to follow. As far as could be determined this is
the first detailed integumental anatomy of a butterfly. Several
features of the monarch's anatomy, however, have been described
and figured in varying detail by other authors (Burgess, 1880;
Kellogg, 1893; Shepard, 1930).

The monarch was chosen for this study because it is common,
widespread, and large. Both dried specimens and individuals pre-
served in Kahle's fixative were used, and these specimens were
sometimes treated with chlorox or KOH. Brushes and fine forceps
were used for the tedious job of removing the scales. The pro-
cedure adopted in preparation of the figures was as follows: every
structure studied was examined in a minimum of two specimens,
in one specimen when the drawing was prepared, and in a second
when the description was written. The vast majority of the struc-
tures have been seen in four or more individuals. The simpler
illustrations were prepared from the first specimen and merely
checked in the second, while others, although outlined from one
specimen, have been altered and adjusted after examination of ad-
ditional specimens whose preparation showed certain features more
clearly. The drawings were all made with the aid of an eyepiece
grid to give proportions correctly.

The terminology used in this paper will be employed throughout
the comparative study to follow. Most of the terms used by
Michener (1952) in his study of the satumiid moth, Eacles im-
perialis (Drury), have been retained; however, the nomenclature
of the male and female genitalia is that of Klots (1956).

Thanks are due to Dr. C. D. Michener, Dr. K. C. Doering and
Dr. Ryuichi Matsuda, all of the University of Kansas, for their aid
on many facets of the work. Thanks go also to my wife, Anne H.
Ehrlich, for aid in the inking of drawings and the preparation of
the plates.

MORPHOLOGY

1

The Head

(Figures 1-7)

The most prominent features of the hypognathous head are the
compound eyes; they are approximately hemispherical and their
combined width is almost one half that of the entire head. The

Anatomy of Monarch Butterfly

1317

vertex

fiagellum

oedicel -
scape

antennifer -

labrum -

anterior tentorial pit

mandibular rudiment''

maxillary palp

transfrontal suture
aterofacial suture

paraocular area

clypeolabral suture
•eye

labial palp

\ ^Npilifer

galea

frontoclypeal sclerite

transfrontal suture
laterofacial suture -

temporal area
paratemporal suture

postocciput
frontoclypeal sclerite

maxillary palp

pilifer

hypostoma.

eye

,„-- paraocular area

- - - vertex

-•*- occiput

temporal suture

2

mandibular rudiment
.- stipes

occiput'

-- cardo
■socket of labial palp

labial sclerite
2mm

Figs. 1-3. Danaus plexippus. 1. Front view of head. 2. Dorsal view
of head. 3. Ventral view of head.

sclerites of the frontal portion of the head between the eyes are
termed here, collectively, the face. The homologies of these scle-
rites are uncertain, but it seems unlikely that the terminology used
by Michener (1952), which is based on that of DuPorte (1946),

1318 The University Science Bulletin

reflects the true situation. The nomenclature employed here for
the facial sclerites is based on DuPorte's more recent work ( 1956 ) .
The central area of the face is occupied by the large protuberant
roughly circular frontoclypeal sclerite. This structure is bounded
above by a sulcus connecting the inner margins of the antennal
sockets, the transfrontal suture. This suture is only weakly in evi-
dence externally, but is represented by a fairly strong ridge in-
ternally. Dorsolaterally the sclerite is bounded by the antennal
sockets, at the edges of which it is infolded to form strong ridges
along the lower parts of the sockets. These ridges connect with
the ridge of the transfrontal suture and each bears on its lateral end
a small dorsal projection, the antennifer, which is an articulation
point for the scape of the antenna. Laterally the frontoclypeal
sclerite is bounded by the curved laterofacial sutures, which con-
tain the prominent anterior tentorial pits and run from the dorso-
lateral edges of the labrum up to the ventral margins of the antennal
sockets. These sutures, although not extremely prominent ex-
ternally (they lie at the base of the forward thrust plateau of the
frontoclypeal sclerite), are represented internally by large ridges
which are major strengthening features of the forward wall of the
head. Approximately the lower fifth of the frontoclypeal sclerite
is separated from the main portion by a transverse heavily sclero-
tized band, designated here the transclijpeal band.

Below the frontoclypeal sclerite and separated from it by a weak
clypeolabral suture is a small transverse sclerite, the labrum. The
labrum has on its ventrolateral edges cylindroconical projections
called pilifers, each of which bears a row of bristles on its inner
edge.

The anatomical facts presented by DuPorte (1956, pp. 113-114,
fig. 4) have been confirmed by dissection by the present author.
The validity of DuPorte's interpretation of these facts rests in large
part on the answers to some of the more basic questions of insect
morphology. These questions, concerning the value of musculature
and innervation as morphological landmarks, the significance of the
position of the anterior tentorial pits and their relationship to the
so-called epistomal and frontogenal sutures, and the strictness with
which interordinal homologies of facial sclerites or areas can be
drawn, cannot be taken up in this paper.

The areas between the laterofacial sutures and the inner margins
of the eyes are known as the paraocular areas (Michener, 1944).
Near the ventral limit of each paraocular area, behind and slightly

Anatomy of Monarch Butterfly 1319

lateral to the pilifer, is a small protuberance, the mandibular rudi-
ment.

The antennae, situated at the dorsolateral margins of the fronto-
clypeal sclerite, are long (each being more than three times as
long as the head is wide) and clubbed. The basal segment, or
scape, of each antenna is relatively large, ringlike, and wider an-
teriorly than posteriorly. It has on its anterodorsal edge a small
articulatory process. The second segment, or pedicel, is a simple
ring approximately one-half the size of the scape. The remainder
of the antenna is the flagellum, which in the monarch is composed
of 43 segments. The segments increase very gradually in length,
and the diameter of the antenna gradually becomes slightly greater
from the proximal end to the vicinity of segment 33. In this region
the width of the segments (and, of course, of the whole antenna)
rapidly increases, while the length of the segments is somewhat re-
duced. The greatest width is reached in the vicinity of segments
39 and 40, while segments 41, 42 and 43 become progressively nar-
rower. The resultant club is somewhat more than one sixth the
length of the antenna.

The ventral surface of the head between the eyes is occupied by
the proboscidial fossa, which is very shallow in this species. The
maxillae occupy most of the anterior section of the fossa. Most
prominent are the galeae, which are greatly elongated, concave
mesally, and grooved together to form a tube through which liquid
food is drawn by the sucking pump. Supporting the galeae, and
extending laterocaudally from them, are the stipites, each bearing
a small tubercle directly behind and slightly lateral to the galea, the
maxillary palp. Each stipes is infolded and longitudinally divided
into two sections by an area of light sclerotization ( shown by heavy
stippling in figure 3 ) . The inner margins of the stipites are bilobed.
Behind each stipes is a small, triangular sclerite, the cardo. The
central and posterior parts of the fossa are occupied by the some-
what triangular labial sclerite, which bears caudally the large sockets
of the labial palps. An invagination along the mid-line of the
sclerite produces an internal ridge, the labial apodeme. The anterior
rim of each palpal socket has two short articular processes. The
anterior parts of the walls of the proboscidial fossa, lateral to the
maxillae, are the hypostomal areas. Projecting forward from their
sockets at the rear of the labial sclerite and up across the face are
the large, three-segmented labial palps. The middle segment of

10—8050

1320

The University Science Bulletin

each palp is the longest; the distal one is the shortest. All segments
are essentially cylindrical, but the distal one is terminally produced
into a point.

The posterior surface of the head is broken centrally by a large
opening, the foramen magnum, which is bisected by a transverse
bar, the tentorial bridge. At the lateral ventral corners of the bridge

postocciput temporal suture

temporal fossa .
postoccipital suture

\ vertex /

tentorial bridge

transoccipital band

posterior tentorial pit' /
articulation of cervical sclerite

anterior tentorial pit

tentorium-

sensory areo

paratemporal suture

-occiput
foramen magnum

para labial suture

labial apodeme
labial palp
labial sclerite

mouth

posterior tentorial pit

,floorof sucking pump
( hypopharynx)

tentorial bridge
2mm

ventral half of

ocular diaphragm

Figs. 4-5. Danaus plexippus. 4. Posterior view of head (the line
marked by short cross lines represents attachment of cervical membrane
to head). 5. Dorsal view of dissection of head to show tentorium (dia-
grammatic ) .

are two depressions, the areas of articulation of the cervical sclerites.
Along its dorsal and dorsolateral margins the foramen is bordered
by the postocciput. This sclerite is well defined dorsally by the
arched postoccipital suture, but the suture is indistinct laterally as
it runs down to the posterior tentorial pits. The latter portions of
the suture are shown as dotted lines in figure 4. The ventral border
of the foramen is the main portion of the labial sclerite behind the

Anatomy of Monarch Butterfly

1321

sockets of the palps; the ventrolateral borders consist of thin upward
growths of the posterior corners of the labial sclerite. The suture
separating this portion of the labial sclerite from the occiput is
called here the paralabial suture. Internally the dorsal portion of
the postoccipital suture is represented by a strong ridge, while the
lateral portions can be detected only as areas of heavy sclerotiza-
tion. The labial suture is, however, represented by a rather strong
ridge, at least in the ventral two thirds of its length.

flagellum

galea of left maxilla

r-

5mm

" 1

Figs. 6-7. Danaus plexippus. 6. Lateral view of head. 7. Antenna.

Lateral to the labial and postoccipital sutures, and covering the
greater portion of the caudal surface of the head capsule, is the
occiput. Approximately the inner one third of the occiput on each
side of the foramen is separated from the remainder by two arcuate,
heavily sclerotized streaks, the transoccipital bands.

In the center of the dorsal surface of the head is the vertex, an
area delimited by the transfrontal suture anteriorly, the antennal
sockets anterolateral^, and the temporal suture laterally and cau-
dally. The temporal suture is represented internally by a rather

1322 The University Science Bulletin

strong ridge, but both the suture and the ridge are indistinct near
the antennal sockets and in the center of the caudal portion. Lateral
to the temporal suture, and running parallel to it from the antennal
sockets to the postoccipital suture, is the paratemporal suture, rep-
resented internally by a ridge which is not as distinct as that of the
temporal suture. On the dorsal surface of the head the area be-
tween the two sutures is here called the temporal area; on the caudal
surface of the head there is a U-shaped depression between the
temporal suture and the paratemporal and postoccipital sutures,
called here the temporal fossa. Within the fossa are two reniform
areas (outlined with dotted lines in figure 4) in which there are a
great many setae. These areas, possibly sensory, may be the
chaetosemata of Jordan ( 1923 ) .

It should be noted here that the homologies of the various features
of the dorsal and caudal surfaces of the head capsule are much con-
fused by the presence of secondary sclerotizations. The above in-
terpretation has been necessarily arbitrary, in an attempt to arrive
at names which can be employed throughout the comparative work
which will follow this paper.

The tentorium consists of the posterior tentorial bar already de-
scribed and two simple anterior arms running between the anterior
and posterior tentorial pits. The anterior arms are somewhat thicker
anteriorly than posteriorly. Between the anterior arms in the front
of the head and attached to the cranial wall near the lower edge of
the labrum is the sclerotic ventral part of the sucking pump. The
structure is roughly semicircular and is made up principally of the
hypopharynx (see Schmitt, 1938). The dorsal portion of the pump
is not sclerotized and does not concern us here.

The compound eyes are separated from the head capsule by thin
ocular diaphragms. These are membranous disks perforated by
large oval openings (long axis dorsoventral ) through which pass
the optic nerves. The diaphragms each have a small sclerotized
area bordering the opening.

Thorax

(Figures 8-21)

Prothorax: The prothorax is much smaller than either of the
segments of the pterothorax. The pronotum is considered to be
divided into three parts, a curved roughly triangular dorsal plate,
and two flat, dorsomedially fused lateral plates. The dorsal plate is
pointed at its posterior end, the beaklike point being curved ventrally

Anatomy of Monarch Butterfly

1323

patogium

dorsal plate of pronoturn

lateral plate of

pronoturn

— inflection of

discrimen

>furcal lamella

spina '
spinasternum

pleural articulation of coxa I

8

2mm

Fig. 8. Danaus plexippus. Posterior ( internal ) view of prothorax.

and articulated with the prescutum of the mesothorax. The fused
portions of the lateral plates form a Y-shaped structure which artic-
ulates with the dorsal plate at the tips of the arms of the Y; the
crotch of the Y is membranous. Just above the lateral plates of the
pronoturn and forward of the cephalic margin of the dorsal plate
are the large, roughly hemispherical patagia. These well-sclerotized
paired structures are the most conspicuous features of the dorsum
of the prothorax. The ventral ends of the lateral plates of the
pronoturn are fused to the dorsal ends of the propleura, which in
turn are fused with one another mid-ventrally, the fusion being
indicated by a faint discrimen. Between the ring formed by the
lateral plates of the pronoturn and the propleura and the foramen
magnum of the head is the membranous cervix. Bridging this
cervical membrane lateroventrally on each side are the cervical
sclerites. They are T-shaped, and each has a circular sclerotic pad
bearing numerous setae on the stem of the T; the pads are called
here cervical organs. The stem of the T articulates internally with
the lateral extremity of the tentorial bridge and the upper arm of

1324

The University Science Bulletin

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Anatomy of Monarch Butterfly 1325

the T externally with the dorsal part of the cephalic margin of the
propleuron. Just forward of the line of fusion of the propleura, but
not cut by the discrimen, is a narrow mid-ventral sclerite projecting
into the cervix, called the presternum.

Internally the discrimen is represented anteriorly by a very weak
inflection and caudally by a small intercoxal lamella. Laterocau-
dally on the rim of the coxal socket is the pointed pleural articula-
tion of the coxa. Externally the discrimen may be traced between
the bases of the coxae as a mid-line marking the base of the inter-
coxal lamella; it ends at the caudal margin of the coxal sockets at
the oval furcasternum. In the center of the furcasternum can be

line of membrane attachment

A
l\

postnotum

10 5mm

Fig. 10. Danaus plexippus. Lateral view of mesothorax, showing phragmata.

seen a dark area representing the furcal pit. Internally the fur-
casternum is produced into a heavily sclerotized two-pronged furca,
which is joined to the pleuron by a plate which is to a large extent
transparent, the furcal lamella.

Bridging the pro-mesothoracic intersegmental membrane mid-
ventrally is a narrow sclerite, the prothoracic spinasternum. The
spinasternum tapers to a point anteriorly where it joins the furcaster-
num of the prothorax, and gradually widens posteriorly, so that
where it meets the mesothoracic katepisternum it is nearly as wide
as the furcasternum. The caudal end of the spinasternum is notched

1326

The University Science Bulletin

so that it joins the katepisternum at two points with a membranous
triangle between them. Near its middle the spinasternum is deeply
invaginated along with the adjacent membrane, forming an internal
projection, the spina.

Mid-laterally in the pro-mesothoracic intersegmental membrane is
the first spiracle. The upper half of the anterior border of the
spiracular opening is occupied by the narrow anterior spiracular

extensor of cervix

external articulation

with pleuron
s

anterior spiracular sclerite

II \

\ \ cervical organ

apodemes^-^ '•£■;

line of membrane attachment

12

internal articulation with head

Imm

posterior spiracular sclerite

13

"~~-apodeme-

2mm

Figs. 11-14. Danaus plexippus. 11. Lateral view of cervical sclerite.
12. Mesal view of first thoracic spiracle. 13. Mesal view of mesothoracic
basalare. 14. Mesal view of mesothoracic subalare.

sclerite, which bears near its ventral end a long apodeme. The en-
tire posterior border of the opening is made up of the bandlike
posterior spiracular sclerite, which has a small apodeme at its lower
end.

Mesothorax: The mesonotum occupies the greater part of the
dorsum of the pterothorax. It is divided into three sclerites, the
prescutum, scutum and scutelhim. The smallest of these, and the

Anatomy of Monarch Butterfly

1327

most anterior, is the prescutum. It is curved strongly ventrally in
front where it articulates with the pronotum, and bears on its
anteroventral margin the thin bilobed first phragma. Arising from
the lateral margins of the first phragma and hanging free in the
body cavity are a pair of phragmal arms. Each lower lateral angle
of the prescutum is produced into a long, slender process or prealare

patagium
tegular arm.

-pronotum

-prescutum 2
- tegula

, — sural are 2
— -scutum 2
— adnotole 2

posterior notal wing

/' process 2

•axillary cord 2

^scuto-scutellar

suture 2
scutellum 2

^, - anterior notal wing

process 3

!]____— postnotum 2

^-scutum 3

"posterior notal wing

process 3

\ n scuto-scutellar suture 3

\ scutellum 3

/

3V

postalor plate 2 I axillary cord 3

I
postnotum 3

process of postalor portion of epimeron 2

5 mm

/

/

Y

■K

Fig. 15. Danaus plexippus. Dorsal view of thorax. Broken line rep-
resents outline of left tegula, removed to show structures beneath it.

extending latero-caudoventrally to just in front of the tegular arm.
The suture between the prescutum and scutum is represented in-
ternally by a weak ridge.

The mesoscntum is the largest sclerite of the thorax. The lateral
edges of the anterior part of the scutum are produced into sloping
plates, the suralares. Internally each suralare is separated from the

1328

The University Science Bulletin

main part of the scutum by a strong ridge, the scutal ridge, which
runs from the posterior margin of the prescutum to the posterior
margin of the scutal incision. The scutal incision is a deep notch
in the lateral edge of the scutum just behind the suralare. From
the scutum just behind the incision a plate, the adnotale, projects
forward forming a lateral border for the posterior part of the inci-
sion. The first axillary sclerite articulates with both the suralare
and the adnotale, which together make up the anterior notal wing
process. Separating the scutum from the smaller mesoscutellum is
the inverted V-shaped scuto-scutellar suture. This suture is rep-
resented internally by a strong ridge. Projecting forward and

scutum 2

scuto-scutellar ridge 2

/
/I

scutum 3

scuto-scutellar

suture 3

scutellum 3

\adnotale /

1st phragma \ ,

phragmal arm ^ j

scutal ridge 2
1st axillary sclerite

h

postnotum
scutal ridge 3
2nd phragma
phragmal process

scuto-scutellar ridge

phragmal articulation

5mm

1

16

Fig. 16. Danaus plexippus. Mesal (internal) view of right half of meso-
and metathoracic terga. Metapostnotum and third phragma omitted.

laterally from the end of the scuto-scutellar suture is a horizontal
shelf, the postalar plate, the posterior portion of which is mem-
branous. The anterior mesal part of the plate is in the form of a
sclerotic arch which is continuous with the caudal part of the
adnotale (this arch is hidden by the scutum in figure 28). The
anterolateral corner of the postalar plate is produced as the posterior
notal wing process, to which is fused the fourth axillary sclerite. Be-
hind the membranous part of the plate is a thin sclerotic strip, and
mesal to this strip is the base of the membranous axillary cord.
Lateral to the plate and mesal to the axillary cord a rounded process

Anatomy of Monarch Butterfly

1329

of the postalar portion of the epimeron projects through the mem-
brane. This process is partially fused to the sclerotic part of the
postalar plate.

A narrow membranous area separates the postnotum ( the phrag-
ma-bearing plate) from the mesoscutellum. The second phragrrw.
is very large and somewhat triangular in lateral view, with dorsal

pleural articulation of coxa I

f urcasternum I -
preepisternum II

katepisternum 11—^

discrimen II

sternopleurite II —

marginopleural— '"'
suture II

katepisternum III—
marginopleural— ~~
suture III

epimeron III-
prespiracular bar-
sternum \*
sternum 2-

presternum

cervical sclerite

--discrimen I
--pleuron I

furcal pit

— -pit of spina

spinasternum I

-preepisternal suture H

— - precoxal suture
—epimeron II

ventral articulation
of coxa II

Hfcr~ -^-discrimen III

ventral articulation
of coxa III

« pleuron 2

4 spiracle 2

5mm

H

-I

17

Fig. 17. Danaus plexippus. Ventral view of thorax and base of abdomen.

and ventral angles posteriorly and a mid-lateral angle forward. The
anterior angles are paired, and each articulates with the mesoscutum
in the socket of a protuberance of the ventral edge of the scutum
immediately behind the caudal end of the adnotale, the phragmal
articulation. Dorsally, at the rear of the phragma, are two promi-
nent triangular projections, the phragmal processes, whose tips serve

1330 The University Science Bulletin

for muscle attachment. The surface of the phragma itself displays
a fairly complex pattern of ridges and varying sclerotization. Some
major features of this pattern are a lightly sclerotized, anteriorly re-
curved area at the lower angle of the phragma, a heavily sclerotized
bracing strut along the lower edge ( sloping from the articulation
almost to the lower angle), and two well-sclerotized ridges running
ventrally from the lower bases of the phragmal processes to the
lower angle. When viewed from behind, the lower angle of the
phragma is not pointed but is fairly broad and bilobed. Continuous
with the top of the phragmal process on each side, and arching
cephaloventrally from it, is a ridge which joins the marginal strut
and with it forms a small lateroventral protuberance. On this ridge
is attached the meso-metathoracic intersegmental membrane. This
line of membrane attachment separates the internal phragma from
the external postnotum.

The more prominent features of the sternopleural region of the
mesothorax are the episternum, epimeron and coxa. The mesepi-
sternum consists of a large katepisterniwi with a tiny anepisternum
nestled between the dorsocaudal corner of the katepisternum and
the ventral side of the basalare. Approximately the lower third of
the katepisternum is separated from the rest of the sclerite by the
precoxal suture, which runs from the pleural suture to the anterior
margin of the katepisternum at the point where it is joined by the
prothoracic spinasternum. This part of the katepisternum is called
here the sternopleurite. Immediately in front of the coxa the
sternopleurite is traversed by the marginopleural suture, which runs
from the pleural suture to the discrimen. The internal margino-
pleural ridge is strongest where it merges with the pleural ridge,
becoming progressively weaker until it meets the base of the lamella
of the discrimen. Internally the precoxal suture forms a strong
ridge, the precoxal ridge, which is continuous with the thickened
upper edge of the lamella of the discrimen. Above the precoxal
suture the katepisternum is separated by a vertical suture, the pre-
episternal suture, from a narrow anterior sclerite, the pre-episternum.

The pre-episternal suture is represented internally by the pre-
episternal ridge. This ridge is rather small at its origin near the
anterior part of the precoxal ridge, but becomes increasingly promi-
nent as it curves dorsally and merges with the pleural ridge.

Mid-ventrally the sternopleurites unite in a suture, the discrimen,
whose inflection forms the very high transparent lamella of the dis-
crimen. The base of this lamella, in the form of two narrow strips

Anatomy of Monarch Butterfly

1331

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1332 The University Science Bulletin

of the sternopleurites, extends backward between the bases of the
coxae to the ventral articulations of the coxae. Posteriorly the
lamella of the discrimen merges into the mesothoracic furca, which
arises from the discrimen above the coxal articulations. The furca,
when viewed in caudal aspect, is roughtly Y-shaped, the arms of
the Y (the secondary f ureal arms) fusing with the ventrocaudal
corners of the prealar portions of the epimera. Running from the
pleural ridges to the anteromesal parts of the secondary arms are
the tendonlike primary furcal arms. The furca is a complex struc-
ture exhibiting varying degrees of sclerotization in different areas.
Major features are three thin rounded lamellae projecting posteriorly
from the main stem of the furca and from near the lateral borders
of each of the secondary arms.

The mesepimeron is divided into two parts, an anterior prealar
portion and a posterior postalar portion. The dorsal edge of the
prealar portion curves strongly downward near the middle, leaving
a relatively large area between it and the ventral side of the subalare.
Near the dorsal part of the anterior border of the epimeron a small
plate is separated from it, the pre-epimeron. The postalar portion
is separated from the dorsocaudal corner of the prealar section by
a line. The anterior end of this portion is inflected and curved
downward into the body cavity as an apodeme and upward as a
process which penetrates the two membranes mesal to the axillary
cord and emerges lateral to the postalar plate. Externally the
postalar portion of the epimeron appears as a long strip which fuses
caudally with the dorsal part of the postnotum.

Between the epimeron and the episternum is the deeply inflected
pleural suture. The internal manifestation of this suture, the pleural
ridge, is the most prominent feature of the mesal wall of the meso-
thorax. Near its dorsal limit, at the point of attachment of the pri-
mary furcal arms, the ridge is produced mesocaudally into a small
plate. The inflection producing this plate is responsible for the
formation of the pre-epimeron, although the deep inflection of the
pleural suture in this area makes the exact method of its formation
difficult to determine. Narrow strips of the anepisternum and
epimeron, carrying with them the pleural suture, project dorsally
between the basalare and subalare as the pleural wing process.
From this process just mesal to the caudoventral corner of the
basalare a tubular internal process projects anteriorly behind the
basalare, the tegular arm. The arm terminates just in front of the
basalare in two lobes, one mesal to the other, whose surfaces are

Anatomy of Monarch Butterfly 1333

external. Articulating with the lateral lobe of the arm is the large
bilobed tegula. The smaller lower lobe of the tegula curves beneath
the leading edge of the wing.

In the region of the pleural coxal articulation the pleural ridge
becomes a quite complex structure as it is joined by the precoxal
ridge, the marginopleural ridge, and the ridges formed by the in-
flected dorsal and anterior margins of the meron. The coxa ( which
is discussed here because it is an integral part of the thoracic cap-
sule) consists of two sclerites, a relatively narrow anterior eucoxa
(Madden, 1944, not Michener) and a bulbous posterior meron. The
suture between the two ( the coxal suture ) seems to be a line suture,
with the internal ridge representing the inflection of the border of
the meron alone. The dorsomesal border of the eucoxa is inflected
and produced into the relatively large eucoxal apodeme. The
caudo-ventromesal part of the coxa is membranous, containing one
round sclerotic island, the coxal sclerule. A small caudomesal
lip of the upper part of the meron, the postcoxal sclerite, is sepa-
rated from the rest of that structure by the postcoxal suture, which
is represented internally by a weak ridge continuous with the in-
flected border of the meron. At the top of the coxal suture there is
a tiny sclerite, a lip of the meron beyond its inflected edge, the
basicoxite.

Above the episternum in front of the pleural wing process is a
roughly diamond-shaped plate, the basalare. Internally a triangular
cavity occupies the central part of the sclerite. A large, for the
most part lightly sclerotized apodeme is attached by an almost
transparent tendon to the anterior corner of the basalare. Above
the epimeron, and separated from it by a considerable expanse of
membrane, is the other epipleurite, the large elongate suhalare.
Internally the subalare has a concavity anteroventrally, and bears
a conspicuous winglike apodeme projecting from its mid-section
into the body cavity.

In the meso-metathoracic intersegmental membrane close to the
upper part of the prealar portion of the epimeron, is the second
spiracle. It is bordered in front and behind by fringed, lightly
sclerotized plates, the anterior and posterior spiracular sclerites.

Metathorax: In the metanotum the metascutum is divided into
two lateral portions by the metascutellum. The scutoscutellar
ridge is very strong and wall-like, dividing the bulging upper part
of the metanotum into three compartments. The anterior notal
wing processes are merely small projections of the anterolateral

The University Science Bulletin

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Anatomy of Monarch Butterfly 1335

walls of the scutum, while the posterior notal wing processes are
long slender projections from the posterior part of the scutum. The
scutal ridge is present in the same position as in the mesoscutum,
but is, of course, much shorter. There is no scutal incision. A
thin membranous area separates the complex metapostnotum dor-
sally from the metascutellum. The salient features of the postnotum
are a ventrally truncated heart-shaped area at the mid-line of the
dorsum, and the phragma which is divided into two arms, each
tipped with a flat oval plate, which project into the body cavity
from near the lateral extremities of the postnotum. The whole
postnotum is so well fused with the tergum and epimeron laterally
that its limits are difficult to ascertain.

As can be seen from the figures, the positions of the various
sclerites of the metathoracic pleural and sternal areas are very
similar to the positions of the homologous sclerites of the meso-
thorax, although their shapes are quite different. Therefore only
major differences will be discussed here. The anepisternum is a
small but well-defined sclerite just above the shoulder of the
katepisternum. It bears numerous bristles, and although it is
closely associated with the basalare and pleural wing process it is
separated from each by a distinct line suture. The precoxal suture
is absent, leaving the katepisternum as an undivided sclerite. There
is an insignificant pre-episternum, which does not continue ventrally
to the level of the coxa. The meron is sharply reduced in favor of
the epimeron.

Internally the metathorax presents quite a different aspect from
the mesothorax. The lamella of the discrimen is arched, arising at
the base of the pre-episternal ridge and terminating at the base of
the furca. The top of the lamella is thickened as in the mesothorax,
and there are two short pointed thickenings in the base of the la-
mella. There is, of course, no precoxal ridge. The eucoxal apodeme
has on its dorsum a small tubercle to which a muscle is attached. The
metafurca is entirely different from the mesofurca. Its most promi-
nent feature is a forward-thrust structure, shaped somewhat like an
arrowhead when seen in dorsal aspect, which overhangs the lamella
of the discrimen. The posteriorly projecting secondary arms are
fused together ventrally forming a V-shaped trough which becomes
progressively shallower caudally. The arms are thin and are broadly
fused to the epimera which are curved mesally and form the caudal
border of the thorax beneath the attachment of the abdomen. The

1336

The University Science Bulletin

secondary furcol arm
meron^

coxal sc'erule
trochanter

— epimeron

furca

postcoxal sclerite

postcoxal suture

20

scutum

posterior notal
wing process

— scutellem

coxa

5mm

h

— axillary cord
"^postnotum

— furca

trochanter

21

■*

Figs. 20-21. Danaus plexippus. 20. Posterior view of mesothorax,
tergum omitted (the line marked by short cross lines represents attach-
ment of meso-metathoracic intersegmental membrane). 21. Posterior
view of metathorax (line of metathorax-abdominal intersegmental mem-
brane indicated on left half of figure only).

Anatomy of Monarch Butterfly

1337

transparent, tendonlike primary furcal arms arise near the base of
the furca and attach to a plate which seems to be an outgrowth of
both the pleural ridge and the epimeron just posterior to it. There
is, however, no obvious pre-epimeron. The tegular arm and tegula
are absent.

The basalare is much smaller than that of the mesothorax, and
bears on its inner surface a relatively large blunt basalare apodeme.
The subalare is small and seems to be merely an external mani-
festation of a sclerotic cap to which the subalare muscle is attached.

Legs

(Figures 22-27)

The prothoracic legs of Danaus, as in all other so-called "four-
footed butterflies," are greatly reduced. The procoxa is long (ap-
proximately the same length as the profemur) and grooved on its
lateral face. The male tarsus is simple (not divided into tarsomeres)

position of scale brush
in metathorocic leg

tibia--

ungues*

5mm

r-

Figs. 22-25. Danaus plexippus. 22. Prothoracic leg of male. 23.
Protarsus of female. 24. Tibia and basal tarsomere of mesothoracic
leg of male. 25. Mesothoracic leg of female.

1338

The University Science Bulletin

while that of the female is club-shaped and divided into foui
tarsomeres, a long proximal one, and three compressed distal ones.
On the caudal side of the distal end of the first three tarsomeres
are paired spines. These are complemented by lobes on the caudal
side of the proximal end of the last three tarsomeres which bear
bundles of setae which cup around the bases of the spines.

The meso- and metathoracic legs are similar in both sexes. In
specimens which have the scales intact, however, there is a small
brush of narrow setalike scales somewhat more than halfway up
the mesal side of the tibia of the mesothoracic leg. This is absent

unguitractor tendon

I
Mi

unguitractor plate,.. / / \~V~ — distitarsus

empodium ^

unguis

26

"pulvillus

arolium

27

imm

Figs. 26-27. Danaus plexippu.s. 26. Detail of distitarsus of pro-
thoracic leg of female. 27. Ventral view of pretarsus of pterothoracic
leg.

in the metathoracic leg. The tibiae of the mesothoracic and meta-
thoracic legs in the females bear numerous spines, while there are
only a few scattered spines in the males. The spines of the proximal
tarsomere are also more prominent in the female.

In the pretarsus, the unguifer ( the dorsal plate to which the tarsal
claws or ungues are articulated) is only slightly sclerotized. The
most prominent feature of the ventral side of the pretarsus is the
large flat unguitractor plate. This plate is tapered internally into
a long thin apodeme, the unguitractor tendon. Just in front of the
unguitractor plate, between the basal parts of the claws, is a small,

Anatomy of Monarch Butterfly 1339

membranous lobe, the empodium. Just dorsal to the empodium
is a small lightly sclerotized protuberance, the arolium. Lateral to
the unguitractor plate are two membranous lobes, the pulvilli. The
ungues are not notched, and are abruptly hooked terminally and
thickened basally.

Wings and Wing Bases

(Figures 28-31, 33)

The areas of the wing articulations present an extremely complex
picture. Each area consists of two membranes and associated
sclerotic plates; an upper membrane connecting the dorsum of the
wing with the tergum, and a lower membrane connecting the
venter of the wing with the pleurites. These membranes are called
the upper and lower alary membranes respectively.

In these membranes are found a series of plates which are
among the most important structures of the wing articulation, the
axillary sclerites. In the mesothorax there are four axillary sclerites.
The first axillary is visible only from the upper side. It is roughly
Y-shaped, with the base of the Y articulating with the adnotale, the
anterior arm with the suralare mesally and the second axillary
laterally, and the posterior arm with the second axillary. The sec-
ond axillary is bilobed when viewed from above. The mesal lobe
articulates with the first axillary mesally and the third axillary
posterolaterally; the lateral lobe is narrowly fused to its anterolateral
corner. Distally the lateral lobe is in contact with the complex
basal sclerotizations of veins Sc, R, Cu and 2V. Beneath the upper
alary membrane both lobes of the second axillary send plates
ventrally which fuse into a blunt process. This process articulates
with the pleural wing process, a small portion being exposed
ventrally.

The third axillary is V-shaped, with the distal arm of the V lying
in the upper membrane and appearing from above as a slightly
arcuate oblong sclerite. The proximal arm projects ventromesally
between the membranes from the caudal end of the distal arm. A
small portion of it penetrates the lower membrane and can be seen
as a narrow strip above the subalare. The muscle of the third
axillary sclerite is attached to the crotch and proximal arm of the
V. The fourth axillary lies in the upper membrane and is fused to
the posterior notal wing process, from which it is differentiated by
a constricted area. Its distal end lies under the flap of membrane
enclosing the third axillary muscle and works against the proximal
arm of the third axillary.

1340

The University Science Bulletin

costal sclerite

basal process of Sc

axillary cord -""
oosterior notal wing process''

postalar portion of epimeron 2'"

--2nd axillary

suralare 2

^— --=Hs1 axillary

• ~ — -3rd axillary
4th axillary

-^postalar plate
28

katepisternum

pleural suture

i

vepimeron

^pleural wing process
2mm

llary cord
-4th axillary

~3rd axillary
fr~ subalare

scle to
3rd axillary

process
nd axillary

29

Figs. 28-29. Danaus plexippus. 28. Dorsal view of mesothoracic
wing base, wing held horizontal. 29. Lateral view of mesothoracic wing
base, wing held vertical.

Anatomy of Monarch Butterfly

1341

costol sclerite

♦R^ -J //' i.

Sc

Rs+Mr
Cu

2V 3V

axillary cord

basal process of Sc

^anterior notal wing process

— -2nd axillary
1st axillary

; 3rd axillary

-muscle to

3rd axillary

30
/ posterior notal wing process

Rs + M

Sc ♦ Rt-

radial sclerite — •

basalare —
pleural wing process--''

anepisternum' // \ I \ \ posterior notal

/ v / \! \ wing process

2nd axillary7

axillary cord

\ 3rd axillary

/ / ^

pleural suture / subalare

muscle to 3rd axillary
2mm

31

Figs. 30-31. Danaus plexippus. 30. Dorsal view of metathoracic
wing base, wing held horizontal. 31. Lateral view of metathoracic wing
base, wing held vertical.

1342 The University Science Bulletin

In front of the first and second axillary sclerites is the pointed
basal process of the subcosta (Sc), which articulates with the
suralare and the anterior arm of the first axillary. On the ventral
side the base of the fused Sc + R is expanded into a bilobed sclerite,
the radial plate.

In the metathorax there are only three axillaries, the fourth
axillary being absent. The pattern of the sclerites is similar to that
of the mesothorax, although their shapes are quite different. The
first axillary is long and thin, with the posterior arm of the Y re-
duced to a mere bulge. The second axillary is very irregular and
is fused anteriorly with the basal process of the Sc. Near the lateral
edge, the second axillary bears a ventral process which penetrates
the lower alary membrane, producing a rather large, ventrally ex-
posed sclerite which articulates with the pleural wing process.

The third axillary is shorter than that of the mesothorax, and the
muscle is smaller. It is more triangular than V-shaped. The por-
tion visible from above is oblong, and the anterior and posterior
ends of this portion are inflected ventrally and fused together to
form a process which penetrates the lower membrane. The muscle
is attached to the upper posterior angle of the rough triangle thus
formed. The posterior notal wing process articulates with the
caudomesal side of the lower apex of the triangle.

Just mesal to the base of Sc -f- Ri and projecting anteriorly is a
large costal sclerite. On the underside a bilobed radial sclerite is
present.

Figures of the wing venation are presented for completeness and
for orientation in connection with the figures of the wing bases.
The venation of the monarch has been figured numerous times be-
fore, and nothing new is added here. The system of naming the
veins is adopted from Klots ( 1951 ) with the substitution of vannul
veins for "anal veins."

Abdomen

(Figures 32, 34-41)

Pre genital segments: Because of the modification of the eighth
sternum of the male into pseudovalves, it can be said that there are
seven pregenital abdominal segments in both sexes of Danaus
plexippus. The first abdominal segment is highly modified, as in
most higher insects, for articulation with the metathorax. The
anterior part of tergum 1 is membranous. The sclerotic portion of
the tergum is dorsocaudally bulged, giving a ponchlike effect. Its

Anatomy of Monarch Butterfly

1343

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1344 The University Science Bulletin

caudal margin slightly overhangs the second abdominal tergum.
On each side of the first tergum, near its margins, is a deep inflec-
tion, the tergal groove. The heavy internal ridge of this inflection
is called the tergal brace. The anterolateral corners of the tergum
are produced laterally into small protuberances, the tergal lobes.
The first abdominal sternum (sternum J) is almost completely
membranous, only a small posterior sclerotic portion remaining
which is fused to sternum 2. This portion of the sternum is also
fused to a thin process of the posterolateral corner of the tergum,
the postspiracular bar. Just in front of this point of fusion another
thin process is emitted by the sternum which crosses the pleural
area below the first abdominal spiracle and terminates just below
the tergal lobe (not fusing with it). This process is called the
prespiracular bar. The abdominal spiracles are all similar to the
first thoracic spiracle but the abdominal anterior spiracular sclerites
resemble the thoracic posterior spiracular sclerites and vice versa.
There are brushes of bristles extending externally and caudally
from the anterior spiracular sclerites of the abdomen.

The first, second, and third terga and sterna are fused together
and lack intersegmental membranes. From the 3-4 intersegmental
area onward there is an increase in the amount of intersegmental
membrane and the resultant amount of possible overlap of segments.
The size and shape of pregenital segments 2-7 can be seen in figure
32. None of them bear significant internal processes.

Male genital segments: The eighth tergum is somewhat reduced.
The eighth sternum is fairly normal, though lightly sclerotized, for
the first one fourth of its length. Its caudal portion, however, is
heavily sclerotized and inflected, forming a U-shaped structure be-
low and around the genitalia proper. The extreme caudal portions
of the sternum extend beyond the pleural membrane as hollow
bilobed protuberances. These eighth sternal structures have the
appearance of paired valvae, and are called here pseudovalves,
following Viette (1948). It should be noted that Klots (1956) is
in error in regarding Viette's term as applying to enlargements of
the ninth sternum ( probably a lapsus calami ) .

Between the eighth and ninth sterna there is on each side an
invagination of the intersegmental membrane which contains a
hair pencil. The ninth sternum is a narrow U-shaped sclerotic band,
termed by taxonomists the vinculum. Mid-ventrally it bears a
blind tubular apodeme directed anteriorly, the saccus. Articulated
to the lateral arms of the vinculum are the paired valvae, morpho-

Anatomy of Monarch Butterfly

1345

anterior spiracular sclerite

aedeagus

uncus

— valvo

sternum 8
sternum 7

35

— diophrogmo
aneilus

— saccus

Figs. 34-38. Danaus plexippus. 34. Mesal view of fourth abdominal
spiracle. 35. Ventral view of apex of male abdomen. 36. Lateral view
of male genitalia proper ( setae of valvae omitted ) . 37. Mesal view of
right pseudovalve (setae omitted), hair pencil and sternum 9. 38. Pos-
terior view of male genitalia proper ( setae omitted ) .

1346 The University Science Bulletin

logically probably the gonocoxites of the ninth segment. Each
valva bears a sharp elongate caudomesal process. The ninth tergum,
which in most Lepidoptera makes up the major portion of the
heavily sclerotized tegumen, is membranous. The region of the
tenth tergum is occupied by two fairly lightly sclerotized lateral
lobes, termed collectively the uncus. There is no sign of a gnathos,
or any other sclerotization of the tenth sternum. A membrane
closes the rear end of the abdominal cavity, running from the bases
of the valvae and the vinculum to the uncus and anus. This terminal
membrane is refered to as the diaphragma. The anus is situated
in the lobe of the membrane which separates the parts of the uncus.
The diaphragma is pierced roughly in its center by the long slender
heavily sclerotized aedeagus. The eversible cone of membrane
around the aedeagus is termed the (melius. The only sclerotization
in the diaphragma proper is a roughly triangular fuxta lying below
the aedeagus and presumably helping to support it.

Female genital segments: Although it is not properly a genital
segment, it should be mentioned that the seventh sternum is some-
what modified to extend up and around the anterior borders of
the sinus vaginalis. The sinus vaginalis is a conspicuous cavity in
the ventral side of the abdomen, just caudal to the seventh sternum.
The sinus contains two sclerotic plates, a large anterior deeply in-
curved lamella antevaginalis, and a smaller posterior lamella post-
vaginalis. The two lamellae are joined laterally and enclose a small
membranous area in the center of which is the receptive opening,
the ostium bursae. Internally the heavily sclerotized ductus bursae
leads from the ostium to the large mebranous corpus bursae. In
the wall of the corpus bursae is a pair of spined signa, joined by a
yoke at the anterior end of the corpus. The corpus, ductus and
ostium bursae together comprise the bursa copidatrix. The lamella
antevaginalis is a structure of the 7-8 intersegmental membrane and
can be differentiated from the seventh sternum, while the lamella
postvaginalis, also presumably intersegmental in origin, cannot be
distinguished from the eighth sternum. The eighth tergum is
divided into two lateral plates, its dorsum being membranous.

The only well-sclerotized structures beyond the eighth segment
are two lateral plates fused to the papillae anales, which bear the
apophyses posteriores, strongly sclerotized paired apodemes which
project forward into the body cavity. These plates are considered
to represent the ninth tergum. The papillae anales are lightly

Anatomy of Monarch Butterfly

1347

lergum 5

sternum 5

papilla analis (retracted)
( tergum 9 + 10)

\ \ slernum 9+10?
\ \amella postvaginalis

lamella antevaginalis sinus v°9inalis

apophysis posterioris

39

signage

bursa copulatrix-

tergum 8
,tergum 9
papilla analis
(extruded)

corpus bursae / :,
ductus bursae 's
ostium bursae/v-^--.

sternum 7 '
lamella antevaginalis

-anus

oviporus

sternum 9+ 10?
lamella postvaginalis

40

tergum 8— — .

lamella postvaginalis

sternum 7

— papilla analis (retracted)

^-ostium bursae

-•lamella antevaginalis

_-i_vi :\ sternum 6

5mm

41

Figs. 39-41. Danaus plexippus. 39. Lateral view of apex of female
abdomen. 40. Lateral view of apex of female abdomen showing internal
structures. 41. Ventral view of apex of female abdomen.

1348 The University Science Bulletin

sclerotized setose lobes on either side of the membranous bulge of
the anal area. They are presumably derived from the ninth or tenth
terga or both. Just below and anterior to the papillae is a small
lightly sclerotized area which may be a remnant of the ninth or
tenth sterna or both. The anus and oviporus, the former above and
behind the latter, lie between the papillae. The ninth and tenth
segments can be retracted within the eighth so that only the tips
of the papillae can be seen, or they may be completely extruded so
that the bulging lightly ridged membrane around the anus protrudes
between the papillae.

VARIATION

In the descriptive section of this work no mention is made of
individual variation between the specimens examined. Disregard-
ing sexual dimorphism two types of variation were observed: varia-
tion in size and variation in shape. However, the structures studied
were invariably present.

Variation in size was not extreme. The length of a primary
(mesothoracic) wing from the base of the subcosta to the apex has
been studied in order to give an idea of the magnitude of this varia-
tion. Twenty-one male specimens from Lawrence, Douglas County,
Kansas showed a range for this measurement of 48-55 mm., with a
mean of 51.2 ± .48 mm. Fourteen females from the same locality
showed a range of 49-57 mm., with a mean of 52.1 ± .67 mm.

Variation in shape was also relatively slight. Some of it can be
attributed to the circumstances of the insect's death (whether it
was pinched, placed in a cyanide jar, or placed directly in fixative);
the outline of the thoracic terga or the amount of membrane ex-
posed between the meso- and metathoraces or the katepisternum
and eucoxa of the metathorax may be affected. Proportions of
various structures may vary, such as the ratio of the length of the
mesoscutellum to the mesoscutum. The courses of most of the
sutures show variability, some ( e. g., the mesothoracic scuto-scutel-
lar) being rather variable, while others (e.g., the mesothoracic
pleural) being relatively stable. Sclerites in which greater than
average variation in shape has been observed are, among others,
the epimera of the pterothorax, the mesothoracic subalare, and the
mesoscutellum. However, it should be emphasized that this varia-
tion is still relatively minor and that in no case is the characteristic
shape of the sclerite lost.

Anatomy of Monarch Butterfly 1349

LITERATURE CITED
Burgess, E.

1880. Contribution to the anatomy of the milkweed butterfly. Anniv.
Mem. Boston Soe. Nat. Hist., 16 pp.

Clark, A. H.

1948. Classification of the butterflies with the alloeation of the genera
occurring in North America north of Mexico. Proc. Biol. Soc.
Washington, vol. 61, pp. 77-84.

DuPorte, E. M.

1946. Observations on the morphology of the face of insects. Jour.

Morph., vol. 79, pp. 371-417.
1956. The median facial sclerite in larval and adult Lepidoptera. Proc.

R. Ent. Soc. London (A), vol. 31, pp. 109-116.

Jordan, K.

1923. On a sensory organ found on the head of many Lepidoptera. Nov.
Zoologicae, vol. 30, pp. 155-158.

Kellogg, V. L.

1893. The sclerites of the head of Danaus archippus. Kansas Univ.
Quart., vol. 2, pp. 51-59.

Klots, A. B.

1951. A field guide to the butterflies. Boston, Houghton Mifflin Co., 349
pp., 40 pis.

1956. Lepidoptera. In Tuxen, Taxonomist's glossary of genitalia in in-
sects. Copenhagen, Ejnar Munksgaard, 284 pp., figs. 1-215.

Madden, A. H.

1944. The external morphology of the adult tobacco hornworm (Lepi-
doptera, Sphingidae). Ann. Ent. Soc. America, vol. 37, pp. 145-
160.

Michener, C. D.

1944. Comparative external morphology, phylogeny, and a classification
of the bees ( Hymenoptera ) . Bull. Amer. Mus. Nat. Hist., vol.
82, pp. 157-326.

1952. The Saturniidae (Lepidoptera) of the Western Hemisphere. Mor-
phology, phylogeny and classification. Bull. Amer. Mus. Nat. Hist.,
vol. 98, pp. 341-501.

ScHMrTT, J. B.

1938. The feeding mechanism of adult Lepidoptera. Smithsonian Misc.
Coll., vol. 97, no. 4, pp. 1-28.

Shepard, H. H.

1930. The pleural and sternal sclerites of the Lepidopterous thorax.
Ann. Ent. Soc. America, vol. 23, pp. 237-260.

VlETTE, P.

1948. Morphologie des genitalia males des lepidopteres. Rev. francaise
Ent., vol. 15, pp. 141-161.

THE UNIVEKSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 19

A Revision of the Subgenus Melanonemotelus of America
North of Mexico (Diptera, Stratiomyidae)*

BY

Wilford J. Hanson

Abstract. This is a revision, based primarily upon male genitalic char-
acters, of the American species of Melanonemotelus, genus Nemotelus, north
of Mexico. Previous to this paper 15 species from this region had been de-
scribed. Nine new species are described, and all old species are redescribed.
Five species are placed in synonymy. In addition, the range of the European
species, Nemotelus nigrinus Fallen, is extended to this region, as four American
species are placed in synonymy with it.

New species described: Nemotelus politus, N. jamesi, N. variabilis, N.
centralis, N. halophilus, N. sabroskyi, N. communis, N. picinus, N. tenuistylus.

Species placed in synonymy: Nemotelus carneus Walker 1849 (syn. of N.
nigrinus Fallen 1817), N. crassus Loew 1861 (syn. of N. nigrinus Fallen
1817), N. unicolor Loew 1861 (syn. of N. nigrinus Fallen 1817), N. car-
bonarius Loew 1865 (syn. of N. nigrinus Fallen 1817), N. fulvicornis James
1936 (syn. of N. glaber Loew 1872).

INTRODUCTION

The subgenus Melanonemotelus of the genus Nemotelus has
long been in need of revision. These flies possess few distinct ex-
ternal characters which are useful in their determination. For this
reason and because of great individual variation, there has been
much taxonomic confusion. Examination of the male genitalia
showed the extent of this confusion. It became apparent that
several described species were poorly defined and that several were
as yet undescribed. This revision of the subgenus Melanonemotelus
is based primarily on male genitalic characters. These characters
generally exhibit noticeable diversity between species which other-
wise appear the same. They are, however, relatively stable within
species and are thus considered reliable. Because of the simplified

* Contribution number 925 from the Department of Entomology of the University of
Kansas.

(1351)
11—8050

1352 The University Science Bulletin

structure of the female genitalia, no usable characters were found
there. Thus as might be expected, the females of several species
are very similar, and determination is often impossible without
males.

The range of Melanonemotelus includes North, Central and
South America, the West Indies, northern Europe, and presumably
northern Asia. This revision includes only those species of North
America north of Mexico. The type specimens of the three de-
scribed species from Mexico have been destroyed, and because of
the lack of sufficient material and absence of genitalic description
or illustration of the types, further consideration of species from this
region is not feasible at this time.

The occurrence of these flies is sporadic. At certain times they
appear by the thousands on their favorite host flowers. The adults
are sluggish and, while feeding, may be taken by hand. Umbellifers
and composites such as goldenrod are among the preferred host
flowers.

Little is known of the immature stages. The larvae of Nemotelus
(s. str.) are known to be aquatic, some living in saline water. It
seems likely that larvae of Melanonemotelus occur in similar habi-
tats. N. (Melanone7notelus) melanderi and sabroskyi are limited
to the Atlantic coast. N. (Melanonemotelus) halophilus has been
collected only along the Texas gulf coast and from salt marshes in
New Mexico and Kansas. The author has collected N. (Mela-
nonemotelus) nigrinus along mountain streams in Utah.

The flies of this subgenus are very small in comparison with most
other Stratiomyidae and are shining black, usually with yellowish
markings. In two species from Florida and the West Indies the
males differ from all others of the subgenus in having abdomens
predominantly whitish. The face of Melanonemotelus, as in all
Nemotelus, is produced anteriorly into a cone to accommodate the
long proboscis. The male and female differ strikingly in the head.
The males are holoptic, the eyes very large and covering most of
the head. The frontal spots, when present, are contiguous. In the
female the eyes are smaller, the frontal spots separated, and the
facial prominence longer (Figs. 1-8). The wing venation is much
reduced, the posterior veins merely impressions in the wing mem-
brane.

An attempt was made to find some characters in the genitalic
structure of each species which might indicate relationships. How-
ever, where it was possible to group species according to similarities

Revision of the Subgenus Melanonemotelus 1353

in genitalia, it was found that the component species of each group
usually differed greatly in external appearance. On the other hand,
certain species very similar externally show striking differences in
the genitalia (e. g., N. melanderi and N. halophilus) . The obvious
similarities are indicated in the discussions of the individual species.
The paratypes of new species herein described have been de-
posited in the Snow Entomological Museum at the University of
Kansas and in the collection of the institution or person from whom
they were borrowed. Where several paratypes were designated,
pairs have been deposited in various other institutions including the
United States National Museum.

ACKNOWLEDGMENTS

The autiior wishes to thank those who have helped in the prog-
ress of this study. Especial thanks are due to the following: Dr.
C. D. Michener, University of Kansas, under whose direction this
work was completed; Dr. M. T. James, State College of Washington,
for suggesting this study, for his advice and information concerning
types, and for the loan of specimens from his private collection and
that of the State College of Washington; Dr. G. F. Knowlton for
his interest in collecting specimens for this work and for the loan
of specimens in the Utah State College collection; Dr. Willis W.
Wirth and Mr. Curtis W. Sabrosky, United States National Museum,
Dr. P. J. Darlington and Dr. W. L. Brown, Jr., Museum of Compara-
tive Zoology, and Dr. C. H. Curran, American Museum of Natural
History, for making possible the study of types in those museums
and for the generous loan of specimens.

Thanks are also due to the following other individuals and in-
stitutions for the loan of specimens: W. F. Barr, University of
Idaho; E. T. Cook, University of Minnesota; Henry Dietrich, Cor-
nell University; R. Dreisbach, Midland, Michigan; W. R. Enns,
University of Missouri; C. L. Fluke, University of Wisconsin;
Leonora K. Gloyd, Illinois State Natural History Survey; Paul D.
Hurd, Jr., University of California; E. L. Kessel, California Academy
of Sciences; Jean Laffoon, Iowa State College; A. T. McClay, Uni-
versity of California at Davis, California; L. W. Quate, University
of Nebraska; H. J. Reinhard, Texas Agricultural and Mechanical
College; G. E. Shewell, Canadian National Collection; W. P.
Stephen, Oregon State College; George Steyskal, Grosse He, Michi-
gan; H. V. Weems, Jr., State Plant Board of Florida; R. L. Wenzel,
Chicago Natural History Museum.

1354 The University Science Bulletin

MALE GENITALIA

(Figs. 10, 11, and 12)

Techniques. Having been removed from the specimen, the
genitalia were placed in a ten-percent solution of KOH and left
overnight. They were rinsed in distilled water and placed in
glycerin. For preservation they were kept in a microvial contain-
ing glycerin and attached to the pin of the corresponding specimen.
For study it was found necessary to separate the genital capsule
from the terga which form a flap over the capsule. It was not nec-
essary to remove the aedeagus from the gonocoxites. So doing
often resulted in damage to certain parts because of the small size
of the genitalia.

Description. The gonocoxites are fused both dorsally and
ventrally, the ventral fusion being much broader. From the base
of the dorsal side of the gonocoxites there extends basally a pair of
long fingerlike apodemes. On each side of these apodemes, and
sometimes between them, the gonocoxites are widely emarginate.
Each of these emarginations may be completely or partially filled
by a thin lamella (basal lamella of gonocoxites). The absence or
extent of these lamellae is a useful character in determining cer-
tain species. The gonostyli are freely articulated to the latero-
posterior portion of the gonocoxites. Mesal to the gonostyli and
forming dorsoventral bridges adjacent to them are a pair of stylus-
like processes extending apically. The posterior margin of the
ventral gonocoxal bridge may be deeply or shallowly emarginate.
The aedeagus consists of two main branches: a dorsal process
and the aedeagus proper. The dorsal process is bifid distally in all
species of this subgenus. It is attached to the dorsal gonocoxal
bridge by a narrow membrane. The ventral branch or aedeagus
proper consists of three lobes: a median penis and two lateral lobes,
here called penis valves. The latter are membranous like the
penis except in one species (N. jamcsi) in which they appear
sclerotized and pincerlike ( Fig. 17b ) . At the base of the aedeagus
is the aedeagal apodeme, which varies in length according to
species.

TAXONOMIC SECTION

The genus Nemotelus of North America was revised by Melander
(1903), who suggested the possible significance of vein R.4+5 when
he noted its unforked condition in certain species. Enderlein

Revision of the Subgenus Melanonemotelus 1355

(1936) divided the genus and placed N. nigrinus Fallen, the only
European species without the anterior branch of vein R4-5, in a
separate genus, Nemotelinus. Because this name was preoccupied
in the family Staphylinidae of Coleoptera, Lindner (1938) pub-
lished the name Melanonemotelus as a subgenus of Nemotelus to
include N. nigrinus.

The genus Nemotelus has been divided according to the follow-
ing combinations of characters: vein R4+5 forked (R4 and Rr, present
and separate distally), and males with mostly whitish abdomen
(Nemotelus s. str.): vein R4+5 not forked (only R5 present distally),
and males nearly entirely black (Melanonemotelus). While these
combinations of characters apply well to the European species, they
fail to hold up when all of the North American species are con-
sidered. N. (Nemotelus s. str.) bruesi Melander and pallipes Say
are both black but have vein R4+5 forked. N. (Melanonemotelus)
flavicornis Johnson and slossonae Johnson both have mostly whitish
abdomens in the males, but vein R4 is not forked.

A study of the male genitalia substantiates the significance of the
nature of vein R4 in defining the subgenera. In all forms in which
vein R4 is simple, the dorsal process of the aedeagus (Fig. 10) is
well developed and has two apical prongs. Examination of several
species having vein R4 forked, revealed that the dorsal process of
the aedeagus had no apical prongs. Besides North American
species, forms from Europe, Central America and South America
were examined.

Key to Subgenera of Nemotelus Geoffroy
Vein R4 present (vein R4 r. forked distally); dorsal process of aedeagus

without apical prongs Nemotelus (s. str.)

Win R4 absent (vein R4-5 not forked distally) (Fig. 9); dorsal process of

aedeagus with apical prongs (Fig. 12) . . . Melanonemotelus Lindner

Subgenus Nemotelus Geoffroy

Nemotelus Geoffroy, 1762, Histoire abregee des Insectes qui se trouvent aux

environs de Paris, dans laquelle ces animaux sont ranges suivant un ordre

methodique. Paris, Durand, vol. 6, p. 542.
Nemotela Latreille, 1796, Precis des caracteres des Insectes disposes dans in

Ordre natural, p. 164.
Nematotelus Osten Sacken, 1863, Cat. of Desc. Dipt, of N. Amer., Arner.,

Smithson. Inst. Misc. Coll., vol. 16, p. 48. Type species: Musca pantherina

Linnaeus, 1758.

A key to the North American species of this subgenus has been
presented by James ( 1936 ) .

1356 The University Science Bulletin

Subgenus Melanonemotelus Lindner

Nematotelus Seguy, 1926, Faune de France. Paris, No. 13, p. 30. (nee

Nematotelus Osten Sacken, 1863. )
Nematolinus Enderlein, 1936, Zweifliigler Diptera. Die Tierwelt Mitteleuropas.

Leipzig, vol. 6, teil 3, p. 79 (type species: Nemotelus nigrinus Fallen, 1817;

monobasic. )
Melanonemotelus Lindner, 1938, Die Fliegen der Palaearktischen Region.

Stuttgart, E. Schweizerbart'sche verlagsbuchhandlung, vol. 4, p. 107 (type

species: Nemotelus nigrinus Fallen, 1817; monobasic).

The body surface in this subgenus is shining black, often with a
bluish or greenish iridescence. The mesonotum is rugulose, es-
pecially toward the lateral margins, but in some species it may be
completely smooth and highly polished. The pile is usually short
and inconspicuous, but may be long and dense in some species,
especially on the dorsum of the thorax. The facial prominence
varies in length, but less so than in Nemotelus (s. str.). Two
creamy-white spots are present on the frons in several species.
The antennae are situated noticeably higher up on the facial
prominence than in Nemotelus (s. str.) and vary little in form. In
the smaller species they are usually shorter and more compact. In
one species (glaber) the basal segments are typically yellowish.
The absence of vein R4 in the wing is the most distinctive character.

In the following key it will be noted that certain couplets apply
only to males (couplets 9-15, 20, and 25). Good external char-
acters being very few in this subgenus, it was found necessary to
use such difficult characters as length and abundance of pile and
relative length of the facial prominence. The latter is measured
from its apex diagonally toward the nearest margin of the eye. The
pile in the female is usually considerably shorter than in the male;
thus, couplet 16 may be difficult to use.

Key to the species of the subgenus Melanonemotelus of America

north of Mexico.

1. Humeral callus with at least anterior half black 6

Humeral callus with less than anterior half black, may be entirely

yellowish white 2

2. Abdomen wholly black tenuistylus

Abdomen whitish or with a pale yellow lateral line on each side 3

3. Thorax bare or nearly so, hairs of dorsal anterior part separated by

length of a hair or more; male with abdomen mostly whitish

(Florida) 4

Thorax with conspicuous pile, hairs of dorsal anterior part separated
by much less than length of a hair; male with abdomen mostly
black 5

Revision of the Subgenus Melanonemotelus 1357

4. Thorax shining black; middle tibiae brown; humeral callus with an-

terior corners black slossonae

Thorax with distinct blue iridescence; middle tibiae yellow; humeral
callus wholly creamy white flavicornis

5. Apices of abdominal terga five and six black, not creamy white; length

of facial prominence less than one third width of eye; female with

large postocular creamy white spot lambda

Apices of abdominal terga five and six creamy white; length of facial
prominence at least one third width of eye; female without creamy
white postocular spot albimarginatus

6. Frons with two creamy white or buff spots, contiguous in male 7

Frons immaculate or with dark brown spots 16

7. Hind tibiae mostly yellowish, with only narrow black band around

mesal portion; abdomen with thin, pale yellow lateral line on each

side; large species canadensis

Hind tibiae black, at most with the bases and apices yellowish; abdo-
men without pale yellow lateral line on each side 8

8. All tibiae entirely black except for brownish extremities; body color

shining black tristis

Fore and middle tibiae yellowish; body color black, usually with bluish
or greenish iridescence 9

9. Pile on dorsum of thorax long over entire surface 10

Pile on dorsum of thorax mesally very short and inconspicuous, much

longer near margins of mesonotum 12

10. Male: Pile on dorsum of thorax very fine, long but inconspicuous,

erect or nearly so, whitish; posterior half of humeral callus creamy
white as seen from dorsal aspect; gonocoxal ventral bridge not
deeply emarginate, but with an apparent double notch in posterior
margin, gonostyli curved, with blunt apices ( Fig. 29 ) . . communis
Male: Pile on dorsum of thorax shining and conspicuous, sometimes
giving a "shaggy" appearance, often with a buff tinge; humeral
callus usually mostly black; gonocoxal ventral bridge deeply emar-
ginate, without an apparent double notch; gonostyli either curved
and with sharp apices, or straight and with blunt apices (Figs. 15,
16, and 18) 11

11. Gonostyli strongly curved, apices pointed (Fig. 16c) (Distribution

mainly Great Basin and Rocky Mountain regions ) politus

Gonostyli straight or only slightly curved, apices rounded, blunt ( Fig.
18c) (Distribution mainly west of Sierra Nevada and south along
Pacific coast ) arator

12. Knobs of halteres chalky white 13

Knobs of halteres pale yellowish buff or partly brown 15

13. Male: Gonocoxites with basal lamellae absent or only slightly evident

( Fig. 14 ) politus

Male: Gonocoxites with basal lamellae complete (Figs. 19 and 20) . . 14

14. Penis valves black, pincerlike, sclerotized (Fig. 19) jamesi

Penis valves transparent, membranous, narrow and inconspicuous

( Fig. 20 ) variabilis

1358 The University Science Bulletin

15. Knobs of halteres wholly pale yellowish buff, sometimes paler beneath

(Atlantic Coast) sabroskyi

Knobs of halteres with clearly defined brown or buff blotch restricted
to dorsoposterior area picinus

16. Dorsum of thorax with pubescence over entire surface, but often

extremely short and inconspicuous mesally especially in female; at

least a few hairs immediately anterior to scutellum 17

Dorsum of thorax bare mesally, sometimes with a few very short hairs
anteriorly, but never immediately anterior to scutellum 21

17. Facial prominence of male less than one third width of eye (Fig. 8a);

small species, about 3 mm. in length 18

Facial prominence of male at least one third width of eye as seen
from lateral aspect (Figs. 1 and 4); length 3.5 mm. or longer. ... 19

18. Abdomen with buff or light yellow lateral line on each side, these

lines very thin in female; basal lamellae of gonocoxites absent ( Fig.

23 ) Jialophilus

Abdomen without buff lateral lines; basal lamellae of gonocoxites com-
plete ( Fig. 22 ) melanderi

19. Facial prominence very long, its length in male half of width of eye

viewed laterally; its length in female about equal to width of eye;

f rons never with spots ( Fig. 1 ) beameri

Facial prominence not noticeably long and tapering, its length in
male less than half the width of eye viewed laterally; its length in
female less than width of eye; frons with or without brown spots
(Fig. 4) 20

20. Dorsal process of aedeagus with apical prongs convergent; base of

aedeagus strongly curved (Fig. 20) (southern species, California

to Florida and into Mexico ) variabilis

Dorsal process of aedeagus with apical prongs divergent; base of
aedeagus not strongly curved (Fig. 21) (Great Lakes region west
to South Dakota ) centralis

21. All tibiae completely black or dark brown glaber

Fore tibiae at least partly yellow 22

22. At least basal segments of antennae yellow or yellowish brown 23

Antennae wholly black or dark brown 24

23. Gonostyli with strongly curved outer edges, apices pointed; penis

valves pointed; gonocoxal apodemes short, convergent (Fig. 14).
Knobs of halteres with brown or buff blotch only on dorsal half

glaber
Gonostyli slightly curved, not so sharply pointed; penis valves trun-
cated, widest at apices; gonocoxal apodemes long, parallel (Fig.
30); knobs of halteres with dorsal brown or buff blotch nearly
covering distal end picinus

24. Knobs of halteres partly brownish or entirely white; aedeagus brown,

sclerotized, the penis and penis valves forming three well-separated
prongs (Fig. 13); facial prominence of female as long as width of
eye or nearly so, as seen from lateral aspect ( some beameri females
may key out here. Compare Figs, lb and 2b) nigrinus

Revision of the Subgenus Melanonemotelus 1359

Knobs of halteres always partly brownish; aedeagus membranous,
penis and penis valves contiguous; facial prominence of female
short, about half as long as width of eye as seen from lateral

aspect 25

25. Basal lamellae of gonocoxites large, extending nearly to apices of
apodemes; penis merely a short lobe between penis valves (Fig.
21 ) centralis

Basal lamellae of gonocoxites absent, apodemes long, parallel; penis
long, longer than prongs of dorsal process ( Fig. 30 ) picinus

Nemotelus nigrinus Fallen

(Figs. 2, 13)

Nemotelus nigrinus Fallen, 1817, Diptera Sveciae, vol. 6, p. 3; Meigen, 1822,
Syst. Beschreib. der bekannten Eur. Zwiefliig. Insecten, vol. 3, p. 1175;
Macquart, 1934, Hist. Nat. des Ins. Dipteres, vol. 1, p. 266; Zetterstedt,
1842, Dipt Scand., vol. 1, p. 151; Walker, 1851, Insecta Brittanica, vol. 1,
p. 26; Schiner, 1862, Fauna Austriaca, Die Fliegen, vol. 1, p. 5; Siebke,
1874, Insectorum Norvegicorum, fasc. 1, p. 7; Van der Wulp, 1877, Diptera
neerlandica, p. 447; Aldrich, 1905, Cat. N. Amer. Diptera, Smithson, Inst.
Misc. Coll., vol. 46, p. 190; Lundbeck, 1907, Diptera Danica, part 1, p. 30;
Verrall, 1909, Brit, flies, vol. 4, p. 125; Seguy, 1926, Faune de France, no.
13, p. 30; James, 1951, Proc. Ent. Soc. Wash., vol. 53, p. 343.

Nemotelus carneus Walker, 1849, List of the Specimens of Dipterous Insects
in the Collection of the British Museum, part 3, p. 521; Melander, 1903,
Psyche, vol. 10, p. 178; Aldrich, 1905, Cat. N. Amer. Diptera, Smithson.
Inst. Misc. Coll., vol. 46, p. 190 (new synonymy).

Nemotelus crassns Loew, 1861, Diptera Americae Septentrionalis Indigena,
Cent. Ill, p. 109; Williston, 1885, Can. Ent., vol. 17, p. 128; Melander,
1903, Psyche, vol. 10, p. 175; Aldrich, 1905, Cat. N. Amer. Diptera, Smith-
son. Inst. Misc. Coll., vol. 46, p. 190; Curran, 1927, Roy. Soc. Canada,
Trans., sect. 5, vol. 21, p. 225 (new synonymy).

Nemotelus unicolor Loew, 1861, Diptera Americae Septentrionalis Indigena,
Cent. Ill, p. 109; Williston, 1885, Can. Ent., vol. 17, p. 128; Melander,
1903, Psyche, vol. 10, p. 176; Aldrich, 1905, Cat. N. Amer. Diptera, Smith-
son. Inst. Misc. Coll., vol. 46, p. 190; Curran, 1927, Roy. Soc. Canada,
Trans., sect. 5, vol. 21, p. 224; Leonard, 1928, Cornell Agric. Expt. Sta.,
Mem. 101, p. 753; Knowlton, 1931, Can. Ent., vol. 63, p. 153 (new
synonymy ) .

Nemotelus carbonarius Loew, 1865, Diptera Americae Septrentrionalis Indigena,
Cent. VIII, p. 119; Melander, 1903, Psyche, vol. 10, p. 177; Aldrich, 1905,
Cat. N. Amer. Diptera, Smithson. Inst. Misc. Coll., vol. 46, p. 190; Leonard,
1928, Cornell Agric. Expt. Sta., Mem. 101, p. 753 (new synonymy).

This is a widespread northern European and northern North
American species. Although the type was not examined by this
author, several specimens from England and northern Sweden were
available and their genitalia were examined. The form known as
unicolor, which is widespread in Canada, Alaska, and northern
United States, shows no significant differences, either externally or
in the genitalia, from European nigrinus, except that in some speci-
mens the knobs of the halteres are somewhat darkened. The type
of carbonarius, although smaller, has genitalia identical with ni-

1360 The University Science Bulletin

grinus. Its halteres are darkened, unlike typical nigrinus, and the
thoracic lateral lines are almost obliterated. However, intermediate
conditions between the two are frequently encountered, and no
distinct line of separation was found possible. The type of crassus,
a female, appears to be merely a slight variant of nigrinus with the
second antennal segment yellowish apically. In all other respects
it is like nigrinus, which cannot be confused with other species found
in Rhode Island, the type locality of crassus. The type of carneus,
also a female, was not available to the author, but the description
agrees with that of nigrinus, and the type locality (St. Martin's
Falls, Albany River, Ontario ) is within the range of only that species.

In general, the body proportions, especially in the head, are be-
tween beamed and picinus, but some individuals approach or equal
these species in this respect. Examination of the male genitalia is
usually necessary for determination. However, the bare mesal
portion of the dorsum of the thorax distinguishes it from beameri
males, and the wholly black or dark brown antennae nearly always
distinguishes it from glaber, in which the basal segments are yellow
or yellowish brown.

Male: Body color shining black, usually with bluish iridescence
on facial prominence and frons. Facial prominence moderately
long (Fig. 2a); frons without pale spots; antennae brownish black
to black. Thorax with rather sparse, white pile, becoming bare to-
ward mesal portion of mesonotum; scutellum nearly bare; meso-
notum with yellowish lateral margins forming a thin line on each
side; humeral calli with small yellowish spot above each spiracle.
Knobs of halteres white (in some specimens ranging from tan to
dark brown on apical portions ) . Femora black with yellow apices;
fore and middle tibiae yellow, hind tibiae mostly black with yellow
bases and apices; tarsi pale yellow, apical segments brownish.
Abdomen concolorous with thorax, but less pilose. Gonostyli with
outer edges curved subapically; emargination of ventral bridge of
gonocoxites V-shaped; apodemes short; basal lamellae moderately
developed, covering nearly half of emarginations; aedeagus fork-
like, the penis valves pointed, more or less straight but directed
slightly mesad; penis shorter than, and distinctly separated from
penis valves from point of junction. Length, 3.5-4.5 mm.

Female: Like male except for sexual differences in head: eyes
smaller, separated; facial prominence longer (Fig. 2b).

Distribution: Arizona: Oak Creek Canyon. California: Lone
Pine, Inyo Co.; Tejon Canyon, Kern Co.; White Mountains, Mono

Revision of the Subgenus Melanonemotelus 1361

Co. Colorado: Florissant; Gold Hill; Grant. Idaho: Blooming-
ton; Cub River Canyon; Moscow; Paris. Illinois: Algonquin
Calumet; Carbondale; Gary; Chicago; Elburn; St. Joseph; Urbana
White Heath. Indiana: Chesterton; Hammond; Lafayette. Iowa
Alleman; Ames; Polk City; Sioux City. Kansas: Douglas Co.
Muscotah; Sun City. Massachusetts: Beverly. Michigan: Bay
City; Detroit; East Lansing; Eaton Rapids; Hemlock, Saginaw Co.;
Livingston Co.; Mason; Salem; Traverse City; Vandalia. Min-
nesota: Beltrami; Clear River; Eagle Bend; Grand Rapids; Houston
Co.; Itasca Park; John Latch State Park, Winona Co.; Mantorville;
Minneapolis; Owatonna; Plummer; Ramsay Co.; St. Anthony; St.
Peter. Mississippi: "Agricultural College." Nebraska: Kayapaha
Co.; Valentine. New Hampshire: (no specific locality). New
Jersey: Mullica Hill. New Mexico: Moriarty. New York:
Slaterville; Caroline. South Dakota: Big Stone City; Canton.
Utah: Hayden; Holt; Kelton; Smithfield. Wisconsin: Dane Co.
Wyoming: Burntfork; S. Pass City. Alberta: Banff; Laggan.
British Columbia: 290 miles. Manitoba: Fort Churchill, Hudson
Bay Ry. mile 505; Sundown. Northwest Territories: Saw Mill
Bay. Ontario: Orillia; Ottawa; Sandford; Simcoe; Toronto; Tren-
ton. Quebec: Abbotsford; Aylmer; Montreal; Outrem't; Rupert
House. Yukon Territory: Whitehorse. Alaska: Fairbanks;
Matanuska Valley.

The location of the type of nigrimis is unknown to the author,
that of cameus from St. Martin's Falls, Albany River, Ontario, is in
the British Museum, that of crassus from Rhode Island, that of
unicolor from Illinois, and that of carbonarius from Lenox, Massa-
chusetts, are in the Museum of Comparative Zoology at Harvard
University.

Flight records: March 18 (Mississippi) to July 29 (Wyoming).

Nemotelus glaber Loew
(Figs. 4, 14)

Nemotelus glaber Loew, 1872, Diptera Americae Septentrionalis Indigena.

Cent. X, p. 232; Melander, 1903, Psyche, vol. 10, p. 177; Aldrich, 1905,

Cat. N. Amer. Diptera, Smithson. Inst. Misc. Coll., vol. 46, p. 190; Curran.

1927, Roy. Soc. Canada, Trans., Sect. V, vol. 21, p. 224.
Nemotelus ftavicornis James, 1932 (not Johnson, 1894), Amer. Mus. Nov., no.

517, p. 7 (new synonymy).
Nemotelus fulvicornis James, 1936, Bull. Brooklyn Ent. Soc, vol. 31, p. 86

( new name for ftavicornis James ) ( new synonymy ) .

This species has two forms; one with nearly entirely black legs,
the other with extensive yellow areas on tibiae and fore femora.
No intergrades have been found. Both occur throughout the range

1362 The University Science Bulletin

which extends from Michigan and New York, southwestward into
Mexico, and both have been taken together by this author. The
type of glaber belongs to the black-legged form, but the yellow-
legged form appears to be more common. Fulvicornis, the holotype
of which is a female, has the typical orange-tan base of the anten-
nae and olive-tinged thorax of glaber and is here regarded as a
synonym. Should the yellow-legged form prove to be a distinct
species, the name fulvicornis is available for it. The dorsum of the
thorax is nearly bare. The body proportions are like those of
picinus and centralis, and it is smaller and has a shorter facial
prominence than most nigrinus individuals.

Female: Body color shining black with olive tinge, especially
on dorsum of thorax. Facial prominence moderately short, stout
(Fig. 4a); frontal spots absent; antennae short, compact, yellowish
brown except for brownish black terminal pseudosegment. Thorax
nearly bare, pubescence at margins of dorsum short, sparse, absent
mesally. Mesonotum with thin pale yellow line on each lateral
margin extending onto humeral callus. Femora and tibiae black,
the apices with very small yellowish areas; apical two segments of
fore tarsi and apical segments of middle and hind tarsi infuscated,
otherwise pale yellow. Halteres with knobs white, each with brown
blotch dorsally, covering about half of distal end. Abdomen con-
colorous with thorax. Length, 3.3 to 3.5 mm.

Male: Like female except for the usual sexual differences in head
(Fig. 4b). Genitalia small; gonostyli strongly and evenly curved,
with pointed apices; emargination of ventral bridge of gonocoxites
deep, nearly evenly rounded; apodemes convergent; basal lamellae
small; aedeagus similar to that of nigrinus but smaller, penis valves
more blunt and slightly convergent (Fig. 14). Length, 3.3 mm.

Distribution: Illinois: Dubois; Homer; Monticello; Odin; St.
Joseph; Urbana; White Heath; Winchester. Indiana: Lafayette.
Iowa: Lacey-Keosauqua State Park, Van Buren Co.; Lewis and
Clark State Park, Monona Co.; Onawa. Kansas: Lawrence; Mus-
cotah Marsh. Louisiana: Mound; Opelousas; Shreveport. Michi-
gan: Detroit; Saugatuck. Mississippi: "Agricultural College." Mis-
souri: Columbia; Jefferson City. New York: Babylon; Long
Island. Ohio: Summit Co.; Amherst. Texas: Brownsville; Col-
lege Station; Dallas; Devil's River; Harlingen; New Brunswick;
Victoria. Virginia: Potomac Creek. Guerrero: Acapulco. More-
los: Cuernavaca. San Luis Potosi : Pujal; Xilitla.

Revision of the Subgenus Melanonemotelus 1363

Holotype, female: Texas, in the Museum of Comparative Zoology
at Harvard University; that of fulvicornis, female, from Lawrence,
Kansas, is in the American Museum of Natural History.

Flight records: March 17 (Brownsville, Texas) to September 27.

Nemotclu.s canadensis Loew

(Figs. 3, 15)

Nemotelus canadensis Loew, 1861, Diptera Americae Septentrionalis Indigena,
Cent. Ill, pp. 109, 110; Melander, 1903, Psyche, vol. 10, p. 175; Aldrich,
1905, Cat. N. Amer. Diptera, Smithson. Inst. Misc. Coll., vol. 46, p. 189;
Curran, 1927, Roy. Soc. Canada, Trans., sect. V, vol. 21, p. 225; Leonard,
1928, Cornell Agric. Expt. Sta., Mem. 101, p. 753; Knowlton, 1931, Can.
Ent, vol. 63, p. 153.

This widespread species is related to orator and politus. It is
distinguished by the thin lateral yellowish abdominal lines and the
narrow black band around the mesal portion of the hind tibiae.

Male: Body color shining black with olivaceous tinge. Facial
prominence moderately long, stout (Fig. 3a). Frontal spots pale
yellow, contiguous. Antennae brownish black. Thorax densely
pilose, the hairs very long, erect, shining white with slight yellow
tinge, slightly longer near edges of mesonotum; mesonotum with
lateral margin narrowly pale yellow; humeral calli with yellow spot
covering less than one half dorsal surface. Halteres with knobs
white. Femora black with yellow apices; fore and middle tibiae
yellow, hind tibiae with narrow black transverse median band
covering less than one third length of tibiae; tarsi pale yellow, the
apical segments slightly infuscated. Abdomen concolorous with
thorax, less pilose; third, fourth, and fifth abdominal terga narrowly
margined laterally with pale yellow; gonostyli with curved lateral
edges, apices sharply pointed; gonocoxites with ventral bridge
deeply emarginate; aedeagus membranous, the penis valves pointed,
longer than penis ( Fig. 15 ) . Length 4-5 mm.

Female: Like male except for shorter pile and the usual sexual
differences in head. Length 4-5 mm.

Distribution: This species is widely distributed in the following
localities: Northwest Territories: Fort Resolution; Hyarling
River. Alberta: Lethbridge; Medicine Hat; Taber, Chin; Coal-
dale. British Columbia: One Hundred Mi^e House; Oliver.
Manitoba: Baldur; Russel. Saskatchewan: Little Quill; Pleasant
Creek. Alaska: Anchorage; Matanuska. California: Spaulding,
Lassen Co.; Lompoc. Colorado: Roggen. Idaho: Geneva; Henry,
Caribou Co.; Montpelier. Minnesota: Douglas Co.; Willmer.

1364 The University Science Bulletin

Montana: ( no specific locality ) . Nevada: Wells. North Dakota:
Turtle Mountains. Utah: Altonoto; Blue Creek; Brigham;
Ephraim; Fielding; Garden City; Garland; Goshen; Hyde Park;
Logan; Magna; Manti; Mona; Myton; Sage Junction; Salt Lake City;
Slaterville; Smithfield; Spanish Fork; Trenton; Vernal; Wellsville;
Woodside. Washington: Harrah; Roche Harbor; Sunnyside; Top-
penish; Yakima. Wyoming: Yellowstone National Park.

The types, from Fort Resolution, N. W. T., are in the Museum of
Comparative Zoology at Harvard University.

Flight records: May 1 to September 7.

Nemotelus polit.us sp. nov.

(Fig. 16)

Nemotelus simplex Snow, 1904, Univ. Kansas Sci. Bull., vol. 2, no. 12, p. 341
( nomen nudum ) .

This species is related to orator and canadensis which it closely re-
sembles, especially in the form of the genitalia (compare Figs. 15,
16 and 18). It is often difficult to distinguish from communis,
especially the females. However, the pilosity is different, that of
communis being thinner, less conspicuous, and the hairs more
evenly spaced.

Male: Body color shining black with slight bluish iridescence.
Facial prominence moderately long, as in communis ( Fig. 6a ) ; frons
with two pale yellow contiguous spots; antennae brownish black.
Thorax rather densely pilose, the hairs silvery white, suberect,
longer near margins of mesonotum, long on scutellum. Meso-
notum with lateral margins pale yellow; humeral calli with pale
yellow spot above each mesothoracic spiracle, covering about one
third of each callosity (in a few specimens observed about half
of callosity is pale yellow). Knobs of halteres white. Femora
black with yellow apices; fore and middle tibiae yellow, hind tibiae
mostly black with yellow bases and apices; tarsi pale yellow, apical
segments slightly infuscated. Abdomen concolorous with thorax,
less pilose, the hairs shorter and sparser than those on thorax. Gono-
styli with curved outer edges, the apices sharply pointed; ventral
bridge of gonocoxites deeply emarginate at apical margin; penis
valves somewhat pointed, longer than penis (Fig. 16). Length,
4-4.5 mm.

Female: Like male except for sexual differences in head; pilosity
on dorsum of thorax shorter. Length, 4-4.5 mm.

Distribution: California: Deep Spring, Inyo Co.; Kern Co.;
Owen's Valley, Inyo Co.; Paradise Spring, Monterey Co.; Sardine

Revision of the Subgenus Melanonemotelus 1365

Creek, Mono Co.; Victorville, San Bernadino Co. Idaho: Mesa;
Middleton; Nampa; Star; Wilder. Montana: (no specific locality).
Nevada: Austen; Fallon; Reno. Oregon: Elgin. Utah: Allen
Canyon; American Fork; Blue Creek; Brigham; Duchesne; Ephraim;
Farmington; Garland; Heber; Hooper; Huntsville; Lay ton; Logan;
Mapleton; Midvale; Morgan; Pintura; Promontory; Smithfield;
Vernal. Washington: Walla Walla. Wyoming: South Pass City.

Holotype, male: Midvale, Utah, Aug. 7, 1937 (Knowlton); allo-
type, female: Huntsville, Utah, June 8, 1938 (Knowlton and Hardy),
both in the Snow Entomological Museum at the University of
Kansas. Paratypes ( six males, two females ) : Logan, Utah, June 5
and 7, 1939 (G. F. Knowlton and G. S. Stains); American Fork,
Utah, Aug. 23, 1936 (G. F. Knowlton); Mapleton, Utah, June 16,
1937 (G. F. Knowlton); Pintura, Utah, August 11, 1929 (R. H.
Beamer); Wilder, Idaho, June 27, 1945; Clarkston, Utah, June 10,
1944 (S. L. Wood); Huntsville, Utah, June 27, 1937 (G. F. Knowl-
ton); Midddletown, Idaho, July 19, 1944 (W. E. Shull); Fallon,
Nevada, Aug. 12, 1940 (L. J. Lipovsky); Blue Creek, Utah, June 3,
1954 (G. F. Knowlton); Vernal, Utah, June 29, 1954 (G. F. Knowl-
ton).

Flight records: April 4 (Monterey Co., California) to August 12
( Nevada ) .

Ne?notelus arator Melander

(Fig. 18)

Nemotelus arator Melander, 1903, Psyche, vol. 10, p. 179; Aldrich, 1905, Cat.
N. Amer. Diptera, Smithson. Inst. Misc. Coll., vol. 46, p. 189; Curran, 1927,
Roy. Soc. Canada, Trans., sect. V, vol. 21, p. 225.

The commonest species of California west of the Sierra Nevada
Mountains, this species is similar to politus, to which it is closely re-
lated. Most individuals have a greenish tinge, and the gonostyli
are straight and blunt, while politus has gonostyli with curved
lateral edges and pointed apices. Another similar species, canaden-
sis, has lateral abdominal stripes which this species lacks. The
types were not seen by the author; however, several topotypical
paratypes were available and agree with Melander's description
of the species.

Male: Body color shining olivaceous to bluish black. Facial
prominence moderately long, as in communis (Fig. 6a); frons with
two pale yellow contiguous spots; antennae brownish black. Thorax
covered with rather long, dense, pale yellow to white pile, longer
at lateral margins of mesonotum and on scutellum. Mesonotum

1366 The University Science Bulletin

with lateral margins pale yellow, forming a narrow line on each
side; humeral calli with small, pale yellow spot above each meso-
thoracic spiracle. Knobs of halteres white. Femora black with
yellow apices; fore and middle tibiae yellow, hind tibiae mostly
black with yellow bases and apices; tarsi pale yellow; apical seg-
ments slightly infuscated. Abdomen concolorous with thorax, less
pilose. Gonostyli straight, rounded at apices; aedeagus as in politus
( Fig. 16 ) . Length, 4-5 mm.

Female: Like male except for sexual differences in head; pilosity
of dorsum of thorax shorter. Length, 4-5 mm.

Distribution: California: Albany, Alameda Co.; Antioch;
Bolinas; Claremont; Davis; Fairfax; La Jolla; Los Angeles; Modoc
Co.; Monterey Co.; Mt. Diablo; Palo Alto; Pomona; Pt. Reyes,
Marin Co.; Redwood City; Rio Vista; San Diego Co.; Santa Clara
Co.; Sardine Creek, Mono Co.; Tolay Creek, Sonoma Co.; Tracy,
San Joaquin Co.; Ventura Co.; Yolo Co. Washington: Kenne-
wick; Prosser; Walla Walla.

Holotype and allotype: San Diego County, California, in the
Melander collection, Riverside, California.

Flight records: March 5 (San Diego Co., California) to August
31 (Sonoma Co., California).

Nemotelus tristis Bigot

(Fig. 17)

Nemotelus tristis Bigot, 1887, Ann. Soc. Ent. France, ser. 6, vol. 7, p. 30.
Melander, 1903, Psyche, vol. 10, p. 177.

I have not seen the type, but although the original description is
extremely brief, the combination of pale frontal spots and nearly
completely black legs leaves little doubt as to the identity of this
species.

The black, nonmetallic coloration of this species and the wholly
black femora and tibiae distinguish it from the closely related
canadensis, arator and politus, which have very similar male geni-
talia but possess extensive yellow areas on the legs.

Male: Body color shining black. Facial prominence moderately
long (Fig. 6a); frontal spots creamy white, contiguous; antennae
black. Thorax with rather dense white pile covering entire dorsum,
longer near lateral margins, much shorter on scutellum; mesonotum
with lateral margins narrowly yellow, forming a very thin line on
each side. Each humeral callus with small yellow spot above
mesothoracic spiracle. Femora black, tipped with yellowish brown;
tibiae entirely black except for small yellowish brown areas at

Revision of the Subgenus Melanonemotelus 1367

bases and apices; tarsi with basal segments yellowish brown, the
remaining segments progressively blacker toward apices. Halteres
with knobs pale creamy white. Squamae slightly infuscated. Ab-
domen concolorous with thorax, pile less dense. Gonostyli with
evenly curved outer edges, apices pointed; aedeagus with penis
valves pointed, longer than penis (Fig. 17). Length, 4-5 mm.

Female: Like male except for shorter, more appressed pile on
thorax and the usual sexual differences in head. Length, 4.8 mm.

Distribution: California: Davis; Mt. Hamilton, Santa Clara Co.;
Oakhurst Madera Co.; Panoche Creek, Fresno Co.; Putah Canyon,
Yolo Co.; Woody, Kern Co.

Types: Bigot apparently described this species from a series of
two males and two females collected in California and now in the
care of Mr. J. E. Collin in England. Both of the males are damaged,
and according to Mr. Collin, one appears to represent a distinct
species from the other three specimens in having pale fore and
middle tibiae.

Flight records: March 26 to June 8.

Nemotelus jamesi sp. nov.

(Fig. 19)

This species closely resembles a few other species within its
range, especially arator and politus. However, the male genitalia
are strikingly different in that the penis valves are sclerotized and
sharply pointed in this species. The form of the genital capsule
and strong curvature of the aedeagus indicates a close relationship
with variabilis, a more southern species. The females are without,
or with only minute, frontal spots.

Male: Body color shining black with slight bluish iridescence.
Facial prominence moderately long (Fig. 6a); frontal spots pale
yellow, small, contiguous; antennae brownish black. Thorax with
moderately dense pilosity near margins of mesonotum, becoming
sparse, short, and sub-appressed mesally; lateral margins narrowly
pale yellow; each humeral callus with small pale yellow spot above
mesothoracic spiracle covering less than one third of dorsal surface.
Femora black with yellow apices; fore and middle tibiae yellow,
slightly infuscated; hind tibiae mostly black with yellow apices and
more extensively yellow bases; tarsi pale yellow, apical segments
infuscated. Halteres with knobs white. Abdomen concolorous with
thorax, but less pilose. Gonostyli with apical portions bent inward;
penis valves black, sclerotized, pincerlike. Aedeagus strongly
curved dorsally (Fig. 19).

1368 The University Science Bulletin

Female: Like male except for absence of frontal spots, facial
prominence longer, eyes smaller, pubescence shorter.

Distribution: California: Escondido; Hallelujah Junction, Las-
sen Co. Idaho: Fish Haven. Nebraska: Norway; Valentine.
Utah: Logan Dry Canyon; Thatcher; Yost. Washington: Prosser.

Holotype, male, and allotype, female: Prosser, Washington, May
2, 1951 in the collection of the State College of Washington. Para-
types: ten males, nine females, same data as holotype.

Flight records: May 2 to Aug. 25.

Nemotelus variabilis sp. nov.

(Fig. 20)

This widespread southern species ranges from California to
Florida. It varies greatly in size and in coloration; the frontal spots
are usually absent but are present in some individuals. The geni-
talia are similar to those of jamesi in that the aedeagus is strongly
curved, but differ in being considerably narrower.

Male: Body color shining black with bluish iridescence. Facial
prominence relatively short (Fig. 4a). Frontal spots absent. (In
some, small yellowish or brownish spots are present.) Antennae
brown, basal two segments yellowish brown. Thorax with pile
short over entire dorsum, very short mesally, longer near lateral
margins, short on scutellum. Mesonotum with lateral margins
narrowly yellowish tan. Each humeral callus with small yellow
spot above mesothoracic spiracle. Halteres with knobs pale buff
(white in some). Femora black with yellow apices; fore and mid-
dle tibiae yellow, slightly infuscated; hind tibiae black medially
with yellow bases and apices; (in some individuals each hind tibia
has a narrow medial black band); tarsi pale yellow. Abdomen
concolorous with thorax. Gonostyli curved; ventral bridge of gono-
coxites strongly convex and deeply emarginate at basal margin;
basal lamellae complete, extending nearly to apices of adopemes.
Aedeagus strongly curved; prongs of dorsal process convergent;
penis and penis valves equal in size, very slender and inconspicuous
(Fig. 20). Length, 3.5-4.2 mm.

Female: Like male except for sexual differences in head and
shorter pile. Length, 3.2-4.5 mm.

Distribution: Arizona: Bill Williams Fork. California: Ar-
royo R.; Cazadero; Lone Pine, Inyo Co.; Little Lake, Inyo Co.;
Los Angeles; Olanche, Inyo Co.; Panamint Mountains, Inyo Co.
Florida: Biscayne Bay; Charlotte Harbor; Hibernia; Homestead;
Sanford; St. Augustine; Yankeetown. Nevada: Carson City. New

Revision of the Subgenus Melanonemotelus 1369

Mexico: Laguna, Valencia Co.; Las Cruces; Mesilla Park; Roswell.
Texas: Anahuac; Brownsville; Hidalgo; Southmost; Victoria.
Utah: St. George.

Holotype, male, and allotype, female: Hibernia, Florida, August
7, 1939 (R. H. Beamer), in the Snow Entomological Museum at
the University of Kansas.

Paratypes: seven males, two females, Sanford, Florida, April,
1908 and 1926 (Van Duzee); Biscayne Bay, Florida (C. W. John-
son); Yankeetown, Florida, July 7, 1948 (R. H. Beamer); Hibernia,
Florida, Aug. 7, 1939 (D. E. Hardy); St. Augustine, Florida; Char-
lotte Harbor, Florida.

Flight records: March 27 (southern Texas) to August 8 (Cali-
fornia ) .

Nemotelus centralis sp. nov.

(Fig. 21)

Almost indistinguishable from picinus except for the genitalia,
this species also has much the same range of distribution. The
complete basal lamellae of the gonocoxites, the short, convergent
apodemes, and the small lobelike penis distinguish it from that
species, which has no basal lamellae, long parallel apodemes, and
much longer penis.

Male: Body color shining black. Facial prominence as in glaber
(Fig. 4a), shorter than in nigrinus, rather pointed. Frontal spots
dark brown, not conspicuous. Antennae brownish black, short,
compact. Thorax with short pubescence on dorsum, nearly absent
mesally, longer toward margins. Mesonotum with lateral margins
narrowly pale yellow. Humeral calli with small yellowish spot
above each mesothoracic spiracle. Halteres with knobs brownish
apically and dorsally. Femora black with about one fifth of apical
portions yellow; fore and middle tibiae yellowish brown, hind tibiae
with middle half black, yellow basally and apically; tarsi pale yel-
low. Abdomen concolorous with thorax. Genital capsule rounded
as seen from dorsal aspect; apical process of gonocoxites short,
thick; basal lamellae complete, extending nearly to apices of
apodemes, which are short, convergent; penis valves short, together
forming rounded outline, not extending beyond prongs of dorsal
process; penis appearing as a small medial lobe between the longer
penis valves (Fig. 19). Length, 3.4 mm.

Female. Unknown.

Distribution: Illinois: Algonquin. Michigan: Detroit; Doug-
las Lake; Midland Co. Minnesota: Clear River; Eagle Bend;

1370 The University Science Bulletin

Wadena; Washington Co. Ohio: Summit Co. South Dakota:
Brookings. Ontario: Trenton; Simcoe.

Holotype, male; Clear River, Minnesota, June 22, 1922 (Gilmer)
in the collection of the University of Minnesota. Paratypes: three
males, Brookings, South Dakota, and Washington Co., Minnesota,
July 22, 1914; Simcoe, Ontario, June 20. 1939 (G. E. Shewell).

Flight records: June and July.

Nemotelus melanderi Banks

(Figs. 8, 22)

Nemoielus melanderi Banks, 1920, Canadian Ent, vol. 52, p. 65; Curran, 1927,
Roy. Soc. Canada, Trans., sect. V, vol. 21, p. 225; Leonard, 1928, Cornell
Agric. Expt. Sta., Mem. 101, p. 753.

This coastal species can easily be distinguished from other eastern
species by the small size and short facial projection. In these re-
spects it resembles very closely the southwestern species halophilus.
However, there are great differences in the genitalia (cf. Figs. 22,
23).

Male: Body color shining brownish black. Facial prominence
short (Fig. 8a) rugose, haired, with slight depression below anten-
nae. Frontal spots absent. Antennae brownish black. Thorax with
long thin pile on dorsum. Mesonotum with lateral margins yellow-
ish brown, forming very thin lines which extend onto humeral calli
above mesothoracic spiracles. Halteres with knobs ivory-white.
Femora dark brown, the apices pale; tibiae brown with pale bases
and apices; tarsi pale yellow. Abdomen concolorous with thorax.
Genitalia small, the gonostyli curved; gonocoxal apodemes short,
convergent; basal lamellae large, extending nearly to apices of
apodemes; penis valves appear as small lobes on each side of penis
which extends slightly beyond them (Fig. 22). Length, 3mm.

Female: Like male except for sexual differences in head (Fig. 8b).

Distribution: Florida: Paradise Key; Royal Palm Park. Mary-
land: Chesapeake Beach. Massachusetts: Beverly; Gloucester
Woods Hole. New Jersey: Lake Branch; Morgan. New York
Calverton, Long Island; Cold Spring Harbor; Montouk, Long Island
Orient, Long Island; Southold. Virginia: Mathias Point. Nova
Scotia: Baddeck; Cape Breton.

The types, twelve males, Cheasapeake Beach, Maryland, two
females, Bayville, New York, are in the Museum of Comparative
Zoology at Harvard University.

Flight records: April 4 (Florida) to September 4 (Virginia)

Revision of the Subgenus Melanonemotelus 1371

Nemotelus lambda James

(Figs. 7,24)

Nemotelus lambda James, 1933, Jour. Kansas Ent. Soc, vol. 6, pp. 70-71; 1936,
Bull. Brooklyn Ent. Soc, vol. 31, pp. 86-91.

The globose head, short facial projection, and large pale markings
on the head, thorax and abdomen set this species apart from all
others from this area.

Male: Body color shining black with ivory-white markings.
Facial prominence very short, depressed between apex and origin
of antennae (Fig. 7a). Frontal spots large, ivory-white, contiguous.
Antennae black. Thorax densely pilose, the hairs erect, very long
over entire dorsum. Humeral calli and lateral margins of meso-
notum ivory-white, the latter broad. Knobs of halteres white.
Femora black with whitish apices; fore and middle tibiae yellow,
hind tibiae with black middle portion and yellow bases and apices;
tarsi yellowish white. Abdomen concolorous with thorax, less pilose;
second, third, and fourth tergites with conspicuous lateral margins,
ivory-white, obsolete on first and fifth segments; second and third
sternites with lateral and posterior margins whitish. Gonostyli very
pointed at apex, inner edge broadly lobate in middle region (Fig.
24); apical processes of gonocoxites shorter than gonostyli.
Gonocoxites deeply emarginated on each side of basal apodemes;
basal lamellae absent; basal apodemes very long, parallel. Aedeagus
attached to gonocoxites by broad membrane, apical prongs of dor-
sal process slightly convergent ( Fig. 24b ) . Length, 4-4.5 mm.

Female: Like male except for the following: Facial prominence
short, without depression above. Eyes widely separated. Frontal
spots much larger, not contiguous; posterior ocular orbits promi-
nently ivory-colored on lower half (Fig. 7b). Thorax with pile
much shorter. Abdomen with narrow but definite whitish lateral
margins.

Distribution: New Mexico: Albuquerque; Jemez Springs; Roose-
velt County.

Holotype, male: Roosevelt County, New Mexico, July 7, 1927,
in the Snow Entomological Museum at the University of Kansas.
Allotype, female: Jemez Springs Mountains, New Mexico, May, in
the collections of Purdue University.

Flight records: May and July.

1372 The University Science Bulletin

Nemotelus halophilas sp. nov.

(Fig. 23)

Although resembling melanderi very closely, this species appears
to be more closely related to the following two species as indicated
by the genitalia ( Figs. 22-27 ) . Externally halophilus may be dis-
tinguished by the buff or yellow lateral abdominal line. This
species has been found only near salt marshes in New Mexico and
Kansas and along the gulf coast of Texas.

Male: Body color shining brownish black. Facial prominence
short (Fig. 8a), haired, profile with slight depression below anten-
nae. Frontal spots absent. Antennae brownish black. Thorax
with long thin pile on dorsum. Mesonotum with lateral margins
narrowly yellowish brown, forming thin lines which extend onto
humeral calli above mesothoracic spiracle. Halteres with knobs
white. Femora dark brown with yellowish brown apices; tibia©
dark brown with yellowish brown bases and apices; tarsi pale
yellow. Abdomen concolorous with thorax except that lateral
margins of terga and sterna three to six are yellowish buff. Pilosity
sparser than on thorax. Genitalia small; emargination of ventral
bridge of gonocoxites evenly rounded, moderately deep; gonocoxal
apodemes long, divergent; basal lamellae absent; aedeagus short;
prongs of dorsal process pointed, divergent; penis slightly shorter
than penis valves. Length, 3 mm.

Female: Like male except for sexual differences in head, (Fig.
8b), pilosity shorter on dorsum of thorax, abdominal lateral lines
narrower. Length, 3 mm.

Distribution: Kansas: Clark Co.; Liberal; Stafford Co. Salt
Marsh. New Mexico: 25 mi. west of Tularosa. Texas: Corpus
Christi (holotype, allotype, and 15 paratypes); Galveston; Padre
Island.

Holotype, male, and allotype, female: Corpus Christi, Texas,
January 1, 1946 (R. H. Beamer), in the Snow Entomological Mu-
seum at the University of Kansas.

Flight records: December 12 and January 1 (southern Texas),
June 22 (Kansas).

Nemotelus flavicornis Johnson

(Fig. 27)

Nemotelus jlavicornis Johnson, 1894, Proc. Acad. Nat. Sci. Philadelphia, p.
272; Melander, 1903, Psyche, vol. 10, p. 191; Aldrich, 1905, Cat. N. Amer.
Diptera, Sm. Inst., p. 190.

This species is closely related to slossonae. These two species
are unique among the species of this subgenus in having largely

Revision of the Subgenus Melanonemotelus 1373

whitish abdomens in the males. The thorax of flavicornis has a
greenish blue iridescence and the humeral calli are completely
creamy white.

Male: Facial prominence black, shining, one third of width of
eye. Antennae yellowish tan, terminal pseudosegment darker,
flagellum black. Thorax shining black with bluish luster. Pubes-
cence short, sparse. Lateral margins of mesonotum pale yellow
forming broad lateral lines. Humeri entirely pale yellow. Halteres
with knobs white. Fore and middle femora yellowish with brown-
ish tinge, hind femora mostly yellow with medial black band; fore
and middle tibiae yellowish, hind tibiae with basal half and apical
tip yellow, apical half excepting tip, black; tarsi pale yellow. Wing
vein R4 absent. Abdomen pale yellow, fourth tergite with median
black triangle, fifth tergite black basally. Genitalia small, widest to-
ward apex, ventral bridge shallowly emarginate at apical margin;
apical processes of gonocoxites short, more widely separated than
their lengths ( Fig. 27 ) . Length, 3 mm.

Female: Unknown.

Distribution: Florida: Key Largo, July 19, 1939 (R. H. Beamer).
The only other locality record at present is the type locality in
Jamaica.

Holotype, male: Kingston, Jamaica, in the Museum of Compara-
tive Zoology at Harvard University.

Nemotelus slossonae Johnson

(Fig. 25)

Nemotelus slossonae Johnson, 1895, Proc. Acad. Nat. Sci. Philadelphia, p. 304;
Melander, 1903, Psyche, vol. 10, p. 181; Johnson, 1913, Bull. Amer. Mus.
Nat. Hist., vol. 32, p. 49.

This small species has a black thorax and mostly white abdomen
in the male. The female has white lateral abdominal lines. It is
closely related to flavicornis, another Floridian species, the genitalia
of both being very similar.

Male: Facial prominence shining, less than one third of width of
eye. Frontal spots creamy white. Antennae brown. Thorax shin-
ing black; pubescence on dorsum very sparse. Mesonotum with
lateral edges pale yellow, forming broad lines from humeral calli
to wing bases. Humeral calli creamy white except anterior corners
which are brownish. Halteres with knobs white. Femora black
except for pale yellow apices; fore tibiae yellow, slightly infuscated,
middle and hind tibiae brownish black with pale yellow bases and
apices; tarsi pale yellow. Wing vein R4 absent. Abdomen mostly
creamy white; first tergite with black spot immediately beneath

1374 The University Science Bulletin

scutellum; fourth tergite with three indistinct brownish spots; fifth
tergite with anterior half brownish black. Sternites mostly white
with indistinct brownish blotches. In one specimen the fourth
tergite has only one medial triangular spot. Genitalia very small,
similar to that of flavicornis; apical prongs of dorsal process of
aedeagus divergent, longer than the separation between them ( Fig.
25). Length, 3 mm.

Female: Like male except for sexual difference in head, and in
abdomen which is black with thin lateral pale yellow lines on mar-
gins of terga two to five. Length, 3.4 mm.

Distribution: Florida: Big Pine Key; Fruitville; Homestead;
Key West; Merritt Islands; Mims; Saddlebunch Keys; Tampa.

Holotype, male: Charlotte Harbor, Florida, in the Museum of
Comparative Zoology at Harvard University. Allotype, female:
Punta Gorda, Florida, Nov. 11, 1911, in the American Museum of
Natural History, New York City.

Flight records: January 29, February 12, March 18, April 4, June
22, July 20, 25, August 11, December 29.

Nemotelus albimarginattis James

(Figs. 5, 26)

Nemotelus albimarginatus James, 1936, Bull. Brooklyn Ent. Soc, vol. 31, pp.
86, 87.

This species is easily distinguished from the similar canadensis
by the creamy-white apices of the fifth and sixth abdominal terga
and larger pale areas on the sides of the thorax and abdomen.

Male: Shining black with slight bluish iridescence. Facial promi-
nence moderately long, about one third to one half width of eye
viewed laterally; frontal spots contiguous, large, creamy white;
antennae brownish black. Thorax densely pilose, hairs very long,
white; suberect; mesonotal lateral margins broadly creamy white;
humeral calli nearly entirely creamy white; supra-alar calli marked
with creamy white; halteres with knobs white. Femora black with
pale yellow apices; fore and middle tibiae yellow, hind tibiae yellow,
each with transverse black mesal annulus; tarsi pale yellow, terminal
segment slightly infuscated. Abdomen concolorous with thorax
except the lateral margins of both terga and sterna ( except the first
segment) and apices of fifth and sixth terga and third, fourth, and
fifth sterna which are creamy white. Gonostyli broad with distinct
lobe on inner edge; emargination of ventral bridge of gonocoxites
evenly rounded, moderately deep; penis valves narrow, pointed
apically as seen from dorsal aspect (Fig. 26). Length, 5 mm.

Revision of the Subgenus Melanonemotelus 1375

Female: Like male except for shorter pilosity, especially on dor-
sum of thorax, and the usual sexual differences in head.

Distribution: California: Benton, Mono Co.; Big Pine and
Little Lake, Inyo Co.; Panocho Creek, Fresno Co.; Wood Lake.
Tulare Co. Idaho: Caldwell; Wilder. Nevada: Valley Hot
Spring, Douglas Co. Oregon: Burns. Utah: Beaver; Delta
Duchesne; Farmington; Hayden; Hooper; Kingston; La Point
Myton; Ogden; Plain City; Riverdale. Washington: Walla Walla
Yakima. British Columbia : Oliver.

Holotype, male: Kingston, Utah, June 19, 1935 (C. J. Sorenson),
in the United States National Museum.

Flight records: March 24 (Fresno Co., California) to August 6
(British Columbia).

Nemotelus sabwskyi sp. nov.

(Fig. 28)

This coastal species has rather small frontal spots and pale
yellowish buff halteres which distinguish it from the slightly larger
but similar communis. The pile on the dorsum of the thorax is also
much shorter.

Male: Body color shining black with slight bluish iridescence,
more pronounced on face. Facial prominence moderately long
(Fig. la); frontal spots pale yellow, small, contiguous; antennae
brownish black. Thorax rather sparsely pilose, hairs white, moder-
ately long near edges of dorsum, becoming very short and nearly
absent mesally, extremely short on scutellum. Mesonotum with
lateral margins narrowly yellowish tan. Each humeral callus with
small yellowish spot above mesothoracic spiracle. Fore femora
black with distal one third yellowish brown, second and third
femora black with distal one fourth yellowish brown; fore and
middle tibiae yellowish brown; hind tibiae with broad black band
on middle portion, covering one third to one half of tibial length,
basal and distal areas yellowish brown; tarsi yellowish tan. Halteres
with knobs pale yellowish buff, paler beneath. Abdomen con-
colorous with thorax, pile very short, sparse. Gonostyli sharply
pointed, expanded on inner edges before apices; aedeagus mem-
branous, darker apically, penis with well-marked indentation at apex
( Fig. 28 ) . Length, 4 mm.

Female: Like male except for shorter pile and the usual sexual
differences in head. Frontal spots usually very small, yellowish tan
or sometimes yellow. Length. 4 mm.

1376 The University Science Bulletin

Distribution: Maryland: Chesapeake Beach. New Jersey:
Atlantic City. North Carolina: Southport.

Holotype, male U. S. N. M. no. 63263, and allotype, female:
Southport, North Carolina, October 10, 1948 (C. W. Sabrosky) in
the United States National Museum. Paratypes: 10 males, 14
females, same data.

Flight records: September 2 (New Jersey) to October 10 (North
Carolina).

Nemotelus communis sp. nov.
(Figs. 6, 29)

Specimens of this species, usually determined as canadensis,
differ from that species in lacking yellowish lateral margins on the
third, fourth, and fifth abdominal terga, and in having a more taper-
ing facial prominence. The pile on the dorsum of the thorax is finer
and the black areas on the hind tibiae are much more extensive.
This species resembles more closely politus, which occurs in much
of its range* of distribution. The humeral spot in communis is
usually larger, covering half of each humeral callus, and the male
genitalia differ considerably. In politus the ventral bridge is deeply
emarginated apically and the aedeagus is short, the penis shorter
than the penis valves. In communis the ventral bridge is not
emarginate but has a double notch effect. The penis is equal in
length to the penis valves ( Figs. 16 and 29 ) .

Male: Body color shining black with slight bluish iridescence
more pronounced on dorsum of thorax. Facial prominence rather
long and tapered (Fig. 6a) with two contiguous creamy white spots
above antennae. Antennae brownish black. Thorax pilose on
dorsum, the hairs rather evenly spaced, white, erect, their lengths
about equal over entire dorsum. Humeral calli with large creamy
white spot covering about half of dorsal surface of each. Lateral
margins of mesonotum creamy white, moderately broad. Halteres
with knobs white. Femora black with yellow apices; fore and
middle tibiae yellow, hind tibiae mostly black with yellow bases
and apices; tarsi pale yellow, the apical segments tinged with light
brown. Abdomen entirely black with long white pile, sparser than
on thorax. Genitalia large; ventral bridge of gonocoxites with an
apparent double notch on posterior margin; gonostyli curved, with
rounded apices; aedeagus membranous, the penis valves bent out-
ward at apices; penis equal in length to penis valves; aedeagal
apodeme short, directed dorsad (Fig. 29). Length, 4.5mm.

Revision of the Subgenus Melanonemotelus 1377

Female: Like male except for shorter pile and the usual sexual
differences in head ( Fig. 6b ) . Length, 4.8 mm.

Distribution: Arizona: Apache Co.; Chiricahua Mountains;
Faraway Ranch. Colorado: Macedonia; Manzanola; Roggen;
Sugar City; Vineland; Wray. Idaho: Pine Lodge; Stone. Illinois:
Chicago. Indiana: Lafayette. Iowa: Ocheyedan. Kansas:
Leavenworth Co.; Liberal; Meade Co.; Medicine Lodge; Menlo;
Stafford Co. Minnesota: Big Stone Co. Montana: Hamilton.
Michigan: Midland Co. Nebraska: Morrill Co.; Willow Island,
Dawson Co. New Mexico: Albuquerque; Belen; Socorro. Oregon:
Rest Lake. South Dakota: Aberdeen; Jefferson; Rapid City.
Utah: American Fork; Antelope; Bert; Blue Creek; Brigham; Claw-
son; Delta; Duchesne; Goshen; Huntsville; Lampo; La Point; Low;
Lyndyl; Murray; Myton; Plain City; Promontory; Richfield; River-
ton; Roosevelt; Salt Lake City; Santaquin; Springville; St. George;
Syracuse; Tremonton; Tridell; Vernal. Wyoming: Teton Pass.
Manitoba: Baldur; Beulah; Red Deer River.

Holotype, male, and allotype, female: Duchesne, Utah, June 29,
1954, on goldenrod (Knowlton), in the Snow Entomological Mu-
seum at the University of Kansas. Paratypes: thirty males and
thirty females, same data as holotype.

Flight records: June 6 (Utah) to September 15 (New Mexico).

Nemotelus beameri James

(Figs. 1,32)
Nemotelus beameri James, 1933, Jour. Kansas Ent. Soc, vol. 6. p. 70.

The relatively long, tapering facial projection is distinctive in
most individuals of this species, but in a few of the specimens avail-
able the projection is not noticeably different from that of some
specimens of nigrinus. The large genitalia with membranous
aedeagus and lanceolate penis are distinctive ( Fig. 32.) .

Female: Body color shining black. Facial prominence long, its
upper margin, viewed laterally, forming a continuous gentle slop-
ing line with the vertex (Fig. 3b); frontal spots absent; antennae
black. Thorax sparsely pilose, the hairs white and short, longer
and more dense at lateral edges of mesonotum, very short on scutel-
lum. Mesonotum with lateral margins pale yellow, resulting in a
thin line on each side; humeral calli with small, pale yellow spot
above each mesothoracic spiracle. Knobs of halteres white, slightly
buff in some. Femora black with yellow apices; fore and middle
tibiae yellow; hind tibiae mostly black with yellow bases and apices;

1378 The University Science Bulletin

tarsi pale yellow. Abdomen concolorous with thorax, pile more
sparse. Length, 4.5-5 mm.

Male: Like female except for sexual differences in head (Fig.
la), dorsum of thorax with pilosity much longer, less sparse, but
short on scutellum. Genitalia large; gonostyli long, rather straight;
apical process of gonocoxites longer than gonostyli; ventral bridge
of gonocoxites very shallowly emarginate; apodemes long, parallel;
basal lamellae large, extending to near apices of apodemes. Aedea-
gus very long; penis valves slightly longer than penis, which is
lanceolate and indented at apex. Length, 4.5-5 mm.

Distribution: Arizona: Tucson. Colorado: Boulder; Floris-
sant; Glen Haven; Greeley. Illinois: Chicago. Iowa: Dan
Green's Stough, Clay Co.; Ocheyedan; Ruthven. Kansas: Meade
Co. Minnesota: Lesueur Co. Montana: Big Timber; Drum-
mond. Nerraska: Lakeside, Sheridan Co. Nevada: Ely. New
Mexico: Sapelle. South Dakota: (no specific locality). Utah:
Duchesne; Greenville; Helper; Kanab; Manila; Springville. Wyo-
ming: Lander; Lusk; Moran; Tie Siding.

Holotype, female: Northgate, Colorado, August 20, 1931, (R. H.
Beamer) in the Snow Entomological Museum at the University of
Kansas. Allotype, male (here designated): Drummond, Montana,
August 11, 1931 (J. O. Nottingham), also in the Snow Entomologi-
cal Museum.

Flight records: March 8 (Arizona) to September 12 (Minnesota).

Nemotelus picinus sp. nov.

(Fig. 30)

This species is distributed in the Great Lakes region west to
Minnesota and Iowa, and is almost indistinguishable externally
from centralis. The differences in male genitalia, as given in the
discussion of that species, are so distinct that determination of the
males is not difficult. Unfortunately, no adeqaute differences in
the females have been found.

Male: Body color shining black (some individuals with slight
bluish luster). Facial prominence as in glaber, pointed (Fig. 4a).
Frontal spots absent (a few individuals with small yellow spots).
Antennae brown (basal segments slightly orange brown in some
individuals), small. Thorax with short pubescence on dorsum,
longer near lateral margins, becoming absent mesally. Mesonotum
with lateral margins narrowly yellow. Humeral calli with small
yellow spot above each mesothoracic spiracle. Halteres with knobs

Revision of the Subgenus Melanonemotelus 1379

browa dorsally and apieally. Femora black with yellow apices;
fore tibiae yellowish brown, middle and hind tibiae brownish black;
tarsi pale yellow. Abdomen concolorous with thorax. Genital
capsule with rectangular shape as seen from dorsal aspect; gonostyli
slightly curved, with subapical lobe on inner margin. Gonocoxal
apodemes long, parallel; basal lamellae absent; penis valves longer
than penis, truncated and wider at apices (Fig. 30). Length, 4 mm.

Female: Like male except for sexual differences in head. Length,
4 mm.

Distribution: Illinois: Algonquin; Chicago; Monticello; Prince-
ton. Indiana: Lafayette. Iowa: Ames. Michigan: Cheboygan
Co.; Lenawee Co.; Vandalia. Minnesota: Minneapolis; St. Anthony
Park; Wadena. Wisconsin: Dane Co.; Madison; Wingra Lake.
Ontario: Toronto. Quebec: Hemmingford.

Holotype, male: Chicago, Illinois, July 6, 1895 (Wheeler); allo-
type, female: Algonquin, Illinois, July 20, 1907 (Nasen), both in
the collections of the Illinois Natural History Survey, Urbana,
Illinois. Paratypes ( ten males and five females ) : Ames, Iowa, July
10, 1951 (M. Cochran), July 12, 1947 and June 10, 1951 (J. Laffoon),
Minneapolis, Minnesota, July 5, 1922 (A. Hertig), Wadena, Min-
nesota, July 4, 1922 (W. Hoffman), Dane County, Wisconsin, July
6, 1951, Princeton, Illinois, July 2, 1936 (Burks).

Flight records: June and July.

Ne?notelus tenuistylus sp. nov.
(Fig. 31)

The broad lateral thoracic lines and nearly entirely yellow
humeral calli are conspicuous characters which separate this species
from all others with completely black abdomens. It is a wide-
spread western species, its relationship to other species indefinite.

Male: Body color shining black with slight bluish iridescence.
Facial prominence moderately long as in communis (Fig. 6a);
frontal spots creamy white; antennae brownish black. Thorax with
short, subappressed pile on dorsum, longer near lateral margins;
mesonotum with lateral edges broadly yellow, the lateral line widest
posteriorly; humeral calli nearly entirely pale yellow, except for
anterior corner which is black. Halteres with knobs white. Fore
femora with basal halves black, apical halves yellow, middle and
hind femora with less yellow at apices; fore and middle tibiae
yellow, hind tibiae with narrow black mesal bands; tarsi pale yellow.
Abdomen concolorous with thorax, sparsely pilose. Gonostyli

1380 The University Science Bulletin

nearly straight, long, at least distal half tapered; emargination of
ventral bridge of gonocoxites shallow, V-shaped, acute; gonocoxal
apodemes long; aedeagus long, narrow, penis valves sharply di-
vergent at apices; prongs of dorsal process several times longer than
their basal separation ( Fig. 29 ) . Length 3.4 mm.

Female: Like male except for sexual differences in head and
shorter pilosity on dorsum of thorax. Length, 3.2 mm.

Distribution: California: Needles. Idaho: Nampa. Kansas:
Liberal; McPherson Co.; Meade Co. Nevada: Carson City; Fallen.
New Mexico: Las Cruces. Utah: Corinne; Garland; Grantsville;
Randlett; St. George.

Holotype, male U. S. N. M. no. 63262: Las Cruces, New Mexico,
June 16, 1917 (Aldrich); allotype, female: same locality, Sept. 25,
1895 ( Cockerell ) , both in the United States National Museum.

Flight records: June 15 (Utah) to September 25 (New Mexico).

LITERATURE CITED
Enderlein, G.

1936. ZweifliAgler Diptera. Tierwelt Mitteleur. Leipzig, Vol. 6, Teil 3,
p. 79.

James, M. T.

1936. Notes on Nemotelus (Dipt., Stratiomyidae). Bull. Brooklyn Ent.
Soc, vol. 31, pp. 86-91.

Lindner, Erwin.

1938. Die Fliegen der Palaearktisehen Region. Stuttgart. Band 4, pp.
106-148.

Melander, A. L.

1903. A Review of the North American Species of Nemotelus. Psyche,
vol. 10, pp. 171-183.

1382 The University Science Bulletin

Figures 1-9

Figs. 1-8. Lateral aspects of heads of certain species of Melanonemotelus;
a. male, b. female.

Fig. 9. Melanonemotelus wing.

Revision of the Subgenus Melanonemotelus 1383

Figs. 1-9

1. beameri

3. canadensis

5. albimarginatus

mgrinus

4. glaber

6. communis

7. lambda

8. melanderi

9

12—8050

1384 The University Science Bulletin

Figures 10-14

Fig. 10. Laterodorsal aspect of genital capsule of male.

Fig. 11. Lateroventral aspect of genital capsule of male.

Fig. 12. Laterodorsal aspect of aedeagus.

Figs. 13 and 14. a. Dorsal aspect of genital capsule, b. Dorsal aspect of
aedeagus. c. Lateral aspect of gonostylus. d. Lateral aspect of apex of
genital capsule, showing apical process of gonocoxites.

Revision of the Subgenus Melanonemotelus
Figs. 10-14

1385

10

apical process of gonoxites .
gonostylus

ventrol bridge of gonocoxites

gonocoxite

basal lamella of gonocoxites
dorsal bridge of gonocoxites

.gonocoxal apodeme

penis valve 1 1
penis

prong of dorsal process
dorsol process of aedeagus

aedeagal apodeme

13. nigrinus

1386 The University Science Bulletin

Figures 15-20

?ct of genital ca;
o^ct of gonostylus.
genital capsule, showing apical process of gonocoxites

Figs. 15-20. a. Dorsal aspect of genital capsule of male. b. Dorsal aspect
of aedeagus. c. Lateral aspect of gonostylus. d. Lateral aspect of apex of

-lital r*ar»eiil<» cVinwincr tinirril nrnpptc nf ffnnnrnvitp«

Revision of the Subgenus Melanonemotelus

Figs. 15-20

138'

15. canadensis

1 7. tristis

r§

19. jamesi

16. politus

18. arator

20. variabilis

1388 The University Science Bulletin

Figures 21-27

Figs. 21-27. a. Dorsal aspect of genital capsule of male. b. Dorsal aspect
of aedeagus. c. Lateral aspect of gonostylus. d. Lateral aspect of apex of
genital capsule, showing apical process of gonocoxites.

Revision of the Subgenus Melanonemotelus

1389

Figs. 21-27

22. melanderi

24. lambda

23. lialophilus

e 25. slossonae

26. albimarginatus

27. flavicornis

1390 The University Science Bulletin

Figures 28-32

Figs. 28-32. a. Dorsal aspect of genital capsule of male. b. Dorsal aspect
of aedeagus. c. Lateral aspect of gonostylus. d. Lateral aspect of apex of
genital capsule, showing apical process of gonocoxites.

Revision of the Subgenus Melanonemotelus

1391

Figs. 28-32

28. sabroskyi

<0 b

30. picinus

31. tenuistylus

32. beameri

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 20

Two New Fossil Plants from the Pliocene of
Northwestern Texas *

BY

A. Byron Leonard **

Abstract. Two new fossil plants, Biorbia papillose and Panicum eliasi,
are described from the Ogallala Formation of Lubbock County, northwestern
Texas.

Investigations of Cenozoic stratigraphy and paleontology in the
Texas High Plains, now in progress by John C. Frye and myself
under the auspices of the Bureau of Economic Geology of the Uni-
versity of Texas, have resulted in the discovery of several out-
crops of Pliocene (Ogallala formation) deposits containing assem-
blages of fossil seeds. One of the most prolific of these exposures
is situated on the prominent Pliocene escarpment a few miles west
of the town of Post, Garza County, Texas. Included in the assem-
blage are an undescribed species of Biorbia ( Boraginaceae ) repre-
sented by a large series of nutlets of this plant and an undescribed
species of Panicum (tribe Paniceae, millet grasses) represented
by the fertile glumes that cover the caryopsis or grain. The starchy
caryopsis or grain, and the chafflike infertile glumes are invariably
wanting in these seeds, but the delicate pericarp may sometimes
be preserved. Fortunately, both the nutlets of borages and the
fertile glumes ( lemma and palea ) of grasses bear some of the most
diagnostic characters of these plants, so that accurate determina-
tions are possible from these parts alone.

The seeds described here occur in partly cemented sands and
gravels; the cementing substance is calcium carbonate for the

* Published with the permission of the Director, The University of Texas Bureau of
Economic Geology.

* Department of Zoology, University of Kansas, Lawrence, Kansas.

( 1393 )

1394 The University Science Bulletin

most part, but the seeds are opalized, which permits their being
cleaned by dissolving the calcite with dilute acid, thus freeing
the seeds of adhering sand and gravel. Seeds are collected by
handpicking from the exposed face of the outcrop; no more pro-
ductive method of collecting seeds from these sediments has been
devised, because of the cementation present.

These and other fossil seeds from the Pliocene of the Great Plains
are catalogued with the collections of Cenozoic mollusks main-
tained at the Museum of Natural History of the University of
Kansas.

Subclass DICOTYLEDONAE

Series GAMETOPETALEAE

Family Boraginaceae

Genus Biorbia Elias, 1932

Biorbia fossilia * is the name applied to the nutlets of the boragi-
naceous plant abundantly represented in late Tertiary sediments of
the Great Plains by these well-preserved fruits. Since there exists
in the literature some confusion concerning the appropriate name
of the plant that produced these nutlets, a review of the nomencla-
ture seems in order.

The genus Biorbia was erected by Elias (1932, p. 350) to receive
Biorbia rugosa (Berry) Elias, originally described by Berry (1928,
art. 13, p. 1) as Lithospermum fossilium var. rugosum. At the
time, Berry had before him a collection of Tertiary seeds collected
by John B. Hatcher; these Berry described as three varieties of a
single new species, viz., Lithospermum fossilium var. rugosum, L. f.
var. glabrum, and L. /. aristatum. Later in the same year, Brooks
(1928) described the same nutlets as Celtis microendocarpiea, but
since Berry's name has date priority, Brooks' name has no real
bearing on the history of the name of the plant in question. In
1931 Elias, realizing that Berry's material consisted of fossil seeds
not properly assigned to the genus Lithospermum, and recognizing
that Berry's L. f. rugosum was actually a borage, while the other
two varieties of L. fossilium were grasses, invented the new name
combination Biorbia rugosa for the boraginaceous plant; L. f.
glabrum and L. f. aristatum were assigned to a new genus Berrich-
loa. These were nomina nuda since they were not accompanied by
descriptions or figures. The next year, however, Elias ( 1932, p.
350) validated the name Biorbia rugosa (Berry) by publishing an

* Prof. G. RuKnieri, University of Palermo, has called to my attention that the correct
name combination is Biorbia fossilis.

Two New Fossil Plants from Texas 1395

adequate description, accompanied by figures. Cockerell v1933,
p. 15), noting Elias' use of the name combination Biorbia rugosa,
called attention to the fact that there was no justification in the
suppression of the species name L. fossilium, and expressed the
view that the correct name combination was Biorbia fossilia (Berry),
on the grounds that the "first and most abundant variety . . .
should be considered the typical form of the species" (Art. 63, 70,
Int. Code Bot. Nomen.). The confusion was compounded by Elias
(1942, p. 106), who used the name combination Biorbia fossilia
(Berry) Elias, when it is obvious that Cockerell is the authority
for the new combination. There seems no doubt that all these
authors were writing of the nutlets of the same plant, so that the
correct name of the species is Biorbia fossilia (Berry) Cockerell.

The history of the name, Biorbia fossilia (Berry) Cockerell may
be summarized as follows:

Lithospermum fossilium var. rugosum Berry, 1932, Proc. U. S.
Nat. Mus., vol. 73, art. 13, p. 1, PI. 1, Figs. 1-10, May 1.

Celtis microendocarpica Brooks, 1928, Annals Carnegie Mus.,
vol. 18, p. 297, PI. 18, Fig. 1, June 1.

Biorbia rugosa (Berry) Elias, 1931, State Geol. Surv. Kansas,
Bull. 18, vol. 32, no. 7, p. 150, April 1. ( nomen nudum )

Biorbia rugosa (Berry) Elias, 1932, Kansas Univ. Sci. Bull., vol.
20, p. 350, PI. 29, Figs. 2a, 2b, 2c, 3, May 15. New name combina-
tion.

Biorbia fossilia ( Berry ) Cockerell, 1933, Torreya, vol. 33, no. 1,
p. 15. Jan. -Feb. New name combination.

Biorbia fossilia (Berry) Elias, 1942, Geol. Soc. America, sp.
papers, no. 41, p. 106, PI. 15, Fig. 12. November 25. Incorrect
authority for name combination.

The genus Biorbia was described as follows (Elias, 1932, p. 350):
"Only nutlets known. Nutlets keeled, asymmetrically inflated, being
more convex on the dorsal side. Surface rugose covered with ridges,
arranged in honeycomblike net. Dorsal keel turns to right or to
left, spoiling the apparent bilateral symmetry of the nutlets. Prob-
ably four nutlets, two right and two left ones, constituted a com-
plete fruit. Scar of attachment round, comparatively small and
elevated, consists of double ring and an elaiosome in the center.
Distinct canal in the ventral part of the scar."

"Genotype: Lithospermum fossilhnn var. rugosum Berry. U. S.
National Museum."

1396 The University Science Bulletin

Biorbia papillosa, sp. nov.
(Plate I, Figures 1, 2, 3)

Types. — Holotype, catalogue number 9917; 160 paratypic ex-
amples from the type locality, catalogue number 9918. Collected
by A. B. Leonard and John C. Frye, June 30, 1955; original number,
A. B. Leonard, 742. Five paratypic examples, catalogue number
9983, from Ogallala formation in Tierra Bianca Canyon, near dam
of Buffalo Lake, Lubbock County, Texas. Collected by John C.
Frye and A. B. Leonard, July 6, 1955; original number, A. B.
Leonard, 765.

Type locality. — Partly cemented sand and gravel at the rim of the
escarpment produced by resistant Pliocene deposits, 3.8 mi. west of
intersection of U. S. Highway 380 and U. S. Highway 84 near the
center of the town of Post, Garza County, Texas.

Diagnosis. — A small nutlet of a boraginaceous plant; nutlet gen-
erally ovoid in shape; base inflated, tapering toward pointed apex,
but also compressed laterally; scar of attachment circular, rim ele-
vated; surface sculpture densely papillose; similar in general form
to the nutlets of Biorbia fossilia (Berry) Cockerell, but differing in:
smaller size (total length less than 2.0 mm), densely papillose
rather than heavy reticulate surface sculpture, and in details of
scar of attachment.

Description of holotype. — Nutlet small, total length 1.7 mm.,
transverse diameter, 1.3 mm., vertical diameter, 1.3 mm.; ovoid in
shape, swollen near base, tapering toward cuspidate apex, but also
compressed; ventral, bladelike ridge, arising from base of style
at anterior rim of scar of attachment, extending forward in midline
over apex, thence backward along mid-dorsal axis of nutlet for a
short distance before extending diagonally to the left, over side of
the nutlet to reach scar of attachment; surface sculpture of closely
spaced, conical, papillose excrescences, varying in height from 0.2
to 0.3 mm.; in a few places along side of the nutlet, the papillae
fused to form short ridges; scar of attachment circular in outline,
depressed strongly in center, within outer rim a parallel groove,
followed medially by a parallel row of six low, rounded papillae or
bosses; in center of scar of attachment a small cylindrical elevation
(elaiosome ?) rising slightly above outer rim of scar; the style, sit-
uated on anterior rim of scar at point of origin of keel, elevated

Two New Fossil Plants from Texas 1397

well above rim of scar, spatulate, possessing a large median vein
and two smaller, parallel veins laterally.

Variations and comparisons. — Size varies little, the largest speci-
mens being slightly under 2.0 mm. in length. The ventral keeled
ridge, after rising over apex of nutlet, may turn either to the left,
as in the holotype, or to the right. This seeming random division
of the seeds into two groups on the basis of the course of the char-
acteristic keeled ridge is similar to the condition in Biorbia fossilia.
On certain paratypic individuals, the coalescence of papillae into
ridges has resulted in an anastomosing, reticulate pattern, like that
typically found on B. fossilia, except that the intervening spaces are
invariably papillose. No smooth seeds, like those commonly found
in large series of B. fossilia, have been observed among B. papillosa.
The scar of attachment of B. papillosa differs from that on the nut-
let of B. fossilia in the number of bosses on the inner circlet within
the depressed central area of the scar; B. fossilia commonly bears
twelve or more of these knoblike elevations. In general, B. papil-
losa is distinguished most readily from B. fossilia by its smaller size,
its papillose rather than reticulate surface sculpture, and its more
cuspidate apex. The ventral ridge on the nutlets of B. papillosa
invariably arises from the base of the style on the scar of attach-
ment; this is not always the case among nutlets of B. fossilia.

Remarks. — In the same assemblage with Biorbia papillosa were
seeds of the hackberry, Celtis willistoni Cockerell, and four grasses,
including the species of Panicum described below, a single example
of a seemingly undescribed Panicum related to the living Panicum
fasciculatum, Berrichloa tuberculoid Elias, and fragments of hulls
of an undescribed species of Stipidium.

It is difficult to make specific stratigraphic assessment of the sedi-
ments bearing this assemblage of seeds, beyond the reasonable
judgment that the outcrop should be assigned to the equivalent of
the Ash Hollow member of the Ogallala formation. Berrichloa
tuberculata is, so far as known, restricted to the Ash Hollow; all
previous records of Panicum are from this stratigraphic unit, and
all previous records of Biorbia are found within the span of Ash
Hollow deposition. Celtis willistoni has a vertical range that pre-
cludes its having any value for precise stratigraphic placement of
these outcrops.

1398 The University Science Bulletin

Subclass MONOCOTYLEDONAE

Order GLUMIFLORAE

Family Gramineae

Subfamily Panicoideae — Tribe Paniceae, The Millet Grasses

Genus Panicum Linnaeus

As far as the fruits are concerned, the genus Panicum Linnaeus
may be characterized as follows: glumes unequal, the first often
minute, the second subequal to the sterile lemma which often en-
closes a hyaline palea and rarely a staminate; fertile lemma and
palea chartaceous-indurated, veins obsolete, the margins of the
lemma inrolled; grain (starchy caryopsis) free within the rigid,
firmly closed lemma and palea.

Panicum eliasi, sp. nov.
(Plate I, Figures 7, 8, 9, 16)

Types. — Holotype, catalogue number 9975; 15 paratypic ex-
amples, catalogue number 9976, from type locality. Collected 30
June 1955 by A. B. Leonard and John C. Frye; original number,
A. B. Leonard, 752. Twenty-nine paratypic examples, catalogue
number 9978, from Ogallala deposits exposed along Acuff Road,
in north valley wall of North Fork of Double Mountain Fork of
Brazos River, 5 mi. S Acuff, Lubbock County, Texas. Collected
by A. B. Leonard and John C. Frye, 6 July 1955; original number,
A. B. Leonard, 765.

Type locality. — Partly cemented sand and gravel at the rim of
the Pliocene escarpment, 3.8 mi. W intersection U. S. Highway 380
and U. S. Highway 84 near center of the town of Post, Garza
County, Texas.

Diagnosis. — A small seed of a panic or millet grass, about 2.5 mm.,
depressed ovoid in shape, swollen posteriorly, narrowing anteriorly
to acuminate, produced apex; consisting of the fertile glumes;
lemma enclosing palea, but the latter broadly exposed; surface
sculpture of wrinkled transverse minute ridges, those on palea
superimposed on minute longitudinal ridges; generally similar to
Panicum clegans Elias, but differing in: size more variable, hull
more depressed, apex more sharply acuminate, and surface trans-
versely wrinkled, rather than smooth and glossy.

Description of holotype. — A small seed of a panic grass, repre-
sented by the fertile glumes; total length 2.4 mm., transverse diam-
eter, 1.7 mm., vertical diameter, 1.4 mm.; depressed ovoid in

Two New Fossil Plants from Texas 1399

shape, greatest diameter (both vertical and transverse) slightly
posterior to center of longitudinal axis, narrowing slowly to acumi-
nate, produced apex; apex acute, bent slightly downward, con-
sisting of central cylindrical spikelike projection and two smaller,
lateral flanking ridges; inflated lemma enclosing palea, but latter
broadly exposed; palea less inflated than lemma, greatest exposed
diameter, 1.4 mm.; palea excavated anteriorly, especially just be-
low reflexed apex, forming pitlike depression; surface sculpture of
lemma wrinkled, transverse ridges, about 20 in a distance of 1 mm.;
sculpture of palea similar but finer in texture, and superimposed
upon minute longitudinal ridges; scar of attachment (not well pre-
served) compressed ovoid, 1.4 mm. in transverse diameter, the
lesser diameter about one-third as great; surface impressed, pitted,
style ( ? ) on dorsal rim.

Variations and comparisons. — Size is variable, the largest speci-
mens being slightly over three millimeters long, but otherwise ob-
served characters are remarkably constant. Panicam eliasi differs
from P. fasciculatum var. reticulatum (Torrey) Beal, a panic
grass living in western Texas and widely dispersed elsewhere, in
being less elongate and less conspicuously sculptured; P. eliasi is
similar in size and shape to P. schribernerianiim Nash, another panic
grass living in the Great Plans, but differs in: acute rather than
blunt apex, and in transverse rather than longitudinally arranged
ridges.

Panicum eliasi is named in honor of Dr. Maxim K. Elias, noted
among his many accomplishments for his pioneering ( 1932 ) and
monographic studies (1942) of the floras of the Tertiary deposits
of the Great Plains.

Remarks. — This is the second species of Panicum to be recognized
in Tertiary deposits of the Great Plains, and represents the Gemi-
nata group, as evidenced by the sculptured surface of the fertile
glumes. P. elegans is found in the upper part of Ash Hollow de-
posits as the Ogallala formation is understood elsewhere, but Elias
(1942) has described a variety of P. elegans which occurs low in
the Ash Hollow member in the lowest part of the Biorbia fossilia
zone. Since the genus is known to occur throughout the Ash
Hollow, the occurrence of P. eliasi throws no light on the precise
stratigraphic placement of the beds, but it may be assumed that
they occur within the equivalent of the Ash Hollow.

At the type locality, Panicum eliasi occurs with the assemblage
of fossil seeds listed above; at the Acuff Road exposure, no other
seeds were found with it.

1400 The University Science Bulletin

It has generally been assumed (and often informally stated)
that fossil seeds are rare or absent in Piocene sediments south of
west central Kansas. Indications from the collections made dur-
ing a rapid reconnaissance of Cenozoic deposits in the Texas High
Plains serve to dispel this erroneous impression, although it is
possible that the rich prairie floras so characteristic of more northern
portions of Pliocene deposits in the Great Plains (Frye and Leon-
ard, 1956) may be less well represented. It is worthy of note,
however, that the typology of the floral assemblage at the Post
locality is similar to many assemblages found in northwestern Kan-
sas ( Frye and Leonard, 1956 ) , and indicates that the southern High
Plains in middle Pliocene time supported a mixed prairie vegetation,
although individual species differ in certain cases.

REFERENCES
Berry, E. W.

1928. Fossil nutlets of the genus Lithosperinum. Proc. U. S. Nat. Mu-
seum, vol. 73, art. 13, pp. 1-3, 1 pi., May 1.
Brooks, Betty W.

1928. A new species of fossil hackberry ( Celtis ) from the "Loup Fork"
Miocene of Phillips County, Kansas. Annals Carnegie Mus., vol. 18,
p. 297, pi. 18, fig. 1, June 1.

COCKERELL, T. D.

1933. The name of a fossil boraginaceous plant. Torreya, vol. 33, no. 1,
p. 15. January-February.
Elias, M. K.

1931. The geology of Wallace County, Kansas. State Geol. Surv. Kansas,
Bull. 18, vol. 32, no. 7, 254 pp., 42 pis., 7 figs., 2 tabls., April 1.

1932. Grasses and other plants from the Tertiary rocks of Kansas and
Colorado. Univ. Kansas Sci. Bull., vol. 32, pp. 115-159, pis. 12-16,
May 15.

1942. Tertiary prairie grasses and other herbs from High Plains. Geol.

Soc. America, sp. paper no. 41, 176 pp., 17 pis., 1 fig., 6 tabls.,

November 25.
Frye, J. C, and Leonard, A. B.

1956. Stratigraphy of the Ogallala formation (Neogene) of northern

Kansas. State Geol. Surv. Bull. 118, 92 pp., 9 pis., 5 figs., April 11.

1402 The University Science Bulletin

EXPLANATION OF FIGURES, PLATE I

Figures 1, 2, 3. Holotype of Biorbia papillosa, catalogue number 9917,
from partly cemented sand and gravel in the Pliocene escarpment, 3.8 mi. W
junction U. S. Highway 84 and U. S. Highway 380 in Post, Garza County,
Texas. Lateral, ventral and dorsal views, respectively. Enlarged 10 times.

Figures 4, 5, 6. Biorbia fossilia ( Berry ) Cockerell, catalogue number 9648,
from Ash Hollow deposits in NW )i NW % Sec. 11, T. 3 S, R. 33 W, Rawlins
County, Kansas. Lateral, ventral and dorsal views of three different nutlets.
Enlarged 10 times.

Figures 7, 8, 9. Holotype of Panicum eliasi, catalogue number 9975, from
partly cemented sand and gravel in the Pliocene escarpment, 3.8 mi. W junc-
tion U. S. Highway 84 and U. S. Highway 380 in Post, Garza County, Texas.
Lateral, ventral and dorsal views, respectively. Enlarged 10 times.

Figures 10, 11, 12. Panicum elegans Elias, catalogue number 9639, from
Ash Hollow deposits, center W line, SW % Sec. 4, T. 4 S, R. 34 W, Rawlins
County, Kansas. Lateral, ventral and dorsal views, respectively. Enlarged
10 times.

Figures 13, 14, 15. Panicum fasciculatum var. reticulatum (Torrey) Beal,
catalogue number 9977, collected living 20 mi. S United States Conservation
Service Nursery, San Antonio, Texas. Dorsal, lateral and ventral views, re-
spectively. Enlarged 10 times. Courtesy of the Botany Department, The
University of Kansas.

Figure 16. Detail of surface sculpture of holotype of Panicum eliasi, en-
larged 120 times.

Figure 17. Detail of sculpture of Panicum faciculatum var. reticulatum
enlarged 120 times.

Figure 18. Paratypes of Biorbia papillosa with B. fossilia. Enlarged ap-
proximately 8 times.

Two New Fossil Plants from Texas

1403

PLATE I.

THE UNIVERSITY OF KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 21

Anatomical Findings at Autopsy Immediately
Following Hanging

BY

Paul G. Roofe, Howard A. Matzke, Irwin L. Baird and
Edwin C. Galbreath *

Abstract. Thirty-five minutes after hanging the head and neck were flushed
with citrated Ringer's via both common carotids. This was followed with
neutral 10 percent formalin. Examination of the remainder of the body was
performed immediately without fixation. The anterior atlanto-occipital mem-
brane and associated ligaments were ruptured as was the posterior atlanto-
epistrophic membrane and ligaments. Associated vertebral musculature showed
hemorrhagic areas. There was no evidence of any fractures in the cervical
vertebrae. The odontoid process and associated ligaments were intact. Gross-
examination of all major blood vessels, meninges, brain, and spinal cord showed
no significant deviation from normal. There was evidence of petechial hemor-
rhages bilateral in the intermediate gray matter of the second cervical segment;
no other histologic changes were noted on preliminary examination.

It was interesting to note that the pituitary gland was twice the normal size.
Death apparently occurred from suffocation within thirteen minutes.

Anatomical findings immediately following hanging do not indi-
cate sufficient structural damage to account for the death of the in-
dividual. We had a unique opportunity to do a complete dissection
immediately after death from hanging; the results are reported here.

Through prearranged agreements with the condemned man, per-
mission was granted to dissect the body as soon as possible after
death. Within 35 minutes after the man was pronounced dead the
head and neck were perfused through the common carotids with
citrated Ringer's solution, followed by 10 percent neutral formalin.
The brain was removed immediately following this procedure. It
was well hardened in situ. The spinal cord was fixed by injecting
10 percent neutral formalin into the spinal canal with a 50 cc hand

* Department of Anatomy, University of Kansas, Lawrence, Kansas.

( 1405 )

1406 The University Science Bulletin

syringe. The cord was removed 10 hours later by a complete
laminectomy.

Careful inspection showed no indication of hemorrhage or large
clot. If there had been damage to the blood vessels to the extent
that hemorrhages had occurred, this would not have been obscured
by the flushing of the vessels.

All coverings of the brain and upper cord were intact. No major
or minor cerebral blood vessel showed damage. If there was hemor-
rhage within the cranial cavity or brain we have been unable to
detect it. Certainly no significant gross structural deviation could
be detected.

Reports on the cause of death from hanging frequently state
"broken neck" or "fractured cervical vertebrae." "Broken neck" is
hard to define; clinically, it involves the crushing or fracture of
vertebrae. In this case we cannot use the term "broken neck."
However, we found that the anterior portion of the atlanto-occipital
membrane and its associated ligaments were ruptured and torn
slightly and, posteriorly, the atlanto-epistrophic membrane and liga-
ments were also torn. Hemorrhagic areas occurred in the intimately
associated vertebral musculature. The odontoid process with its
ligaments was intact.

The central nervous system showed little damage. A few pete-
chial hemorrhages were present in the intermediate gray at the level
of the second cervical segment. No other areas of the central
nervous system showed signs of damage which are referable to the
hanging. The pituitary was about twice its normal size. Blood
channels through it appeared unusually large.

In this individual there was scattered chromatolysis throughout
the cord; this was most marked in the lumbar and sacral areas.
Marked ependymal hyperplasia was pronounced from the mid-
thoracic area to the lower sacral. The ventral white commissure
was displaced. However, no neurological signs were mentioned
in his medical record. Scattered perivascular demyelination was
found in the cerebral white matter in the few areas studied. How
extensive this was is yet to be determined. None of the above
changes could be attributed to the hanging.

The cortices of both cerebrum and cerebellum appeared normal.
We have not yet investigated the medulla, pons, midbrain and
thalamus.

The cause of death could not be determined. It is suggested
that suffocation probably was the reason.

Findings at Autopsy Following Hanging

1407

Legally, hanging is classified as traumatic asphyxia due to strangu-
lation. The lumen of the upper portion of the larynx is effectually
occluded by the direct pressure of the ligature which forces the
base of the tongue upward and backward and depresses the epi-
glottis over the laryngeal opening. The compression applied to the
throat from front to back and from the sides affects the superior
laryngeal nerves, the vagus, the carotid arteries and the jugular
veins, causing shock and direct disturbances to the cerebral circula-
tion.

Figure 1.

Heart beat
ana
Pulsa

M inutcs

The chart is a composite compiled from 12 case histories scattered
in the literature, chiefly from E. Kalle, and indicates the sequence
of events related to heart rate and pulse rate. The attending physi-
cian's report follows very closely the sequence of events indicated
in the accompanying chart. Zero minutes corresponds to springing
of the trap. We can only speculate as to the physiology involved.

1408 The University Science Bulletin

Probably the traumatic irritation of the vagus and carotid body and
sinus and the sudden shock and immediate closure of the laryngeal
opening all contribute to this early low heart rate, if not its complete
stoppage. In some instances the heart rate goes to 150 or higher
in the 4 to 7 minute interval. The average time of death in these
cases was roughly 13 minutes and 30 seconds. The increase of
heart rate may be accounted for by the thoraco-lumbar activation
with excessive adrenalin. Spasms may or may not occur. Gasping
does not occur.

LITERATURE CITED
Kalle, E.

1933. Beobachtungen uber den Tod bei Hinrichtungen mit dem Strang.

Deutsch Zeitschrift fur die Gesamte Gerichtliche Medizin, vol.

22, pp. 192-203.

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 22

A Statistical Method for Evaluating Systematic

Relationships 1

BY

Robert R. Sokal and Charles D. Michener 2

Department of Entomology
University of Kansas, Lawrence

Abstract. Starting with correlation coefficients (based on numerous char-
acters) among species of a systematic unit, the authors developed a method
for grouping species, and regrouping the resultant assemblages, to form a classi-
ficatory hierarchy most easily expressed as a treelike diagram of relationships.
The details of the method are described, using as an example a group of bees.
The resulting classification was similar to that previously established by classi-
cal systematic methods, although some taxonomic changes were made in view
of the new light thrown on relationships. The method is time consuming, al-
though practical in isolated cases, with punched-card machines such as were
used; it becomes generally practical with increasingly widely available digital
computers.

INTRODUCTION

The purpose of the study reported here was to develop a quanti-
tative index of relationship between any two species of a higher
systematic unit, as well as to exploit such indices of association in
the establishment of a satisfactory hierarchy. The authors became
interested in the development of such a method when they at-
tempted to find a technique for classifying organisms that was free
from the subjectivity inherent in customary taxonomic procedure.

1. Contribution number 945 from the Department of Entomology, University of Kansas.

2. We wish to acknowledge the constructive criticism received in connection with this
and related work from the following individuals who kindly gave their time to read and
comment upon the manuscript: Paul R. Ehrlich, University of Kansas; Raymond B. Cat-
tell, University of Illinois; Alfred E. Emerson, University of Chicago; Warwick E. Kerr,
Universidade de Sao Paulo; Ernst Mayr, Harvard University; Louis L. McQuitty, Michigan
State University; G. G. Simpson, American Museum of Natural History; Peter C. Silvester-
Bradley, University of Kansas and University of Sheffield; and Paulo E. Vanzolini, De-
partmento de Zoologia, Secretaria de Agricultura, Sao Paulo. These persons, however, are
not responsible for the opinions which we have expressed.

Acknowledgment is also due to the University of Kansas General Research Fund for
assistance.

(1409)

1410 The University Science Bulletin

The systematic group chosen as a test of the feasibility of this under-
taking was one consisting of 97 species of solitary bees in the family
Megachilidae. This choice was made because one of us (C. D. M.)
has made recent systematic studies of these insects, so that conclu-
sions as to the relationships obtained by the usual systematic pro-
cedure could be compared with the results of the new method.

The findings of our study as well as the philosophical bases of
our attempts at quantifying systematic relationships have been re-
ported elsewhere (Michener and Sokal, 1957). In this paper we
propose to describe in some detail the actual method employed,
as well as our reasons for adopting it and for rejecting several alter-
nate procedures. It is our intention to illustrate the procedures in
sufficient detail so that persons with a limited knowledge of statisti-
cal methods will be able to follow our method. We expect our
system to be applicable to most organisms, provided they exhibit a
variety of characters, and the account to follow is consequently
phrased in general terms. However, our practical illustrations are
based on the bee group cited above in order to provide the reader
with concrete examples.

A quantitative method of finding the relationship between two
species must be based on a number of taxonomic characters in a
manner similar to the traditional systematic approach. However,
whereas the latter technique generally uses few characters and
weights these quite unequally and subjectively, the former method
employs numerous but unweighted characters. Our reasons for not
weighting characters have been detailed in the companion paper
( Michener and Sokal, 1957 ) . In the absence of an objective criterion
of character weight it seems best to rely on a large number of
equally weighted characters. In our bee study we employed 122
characters per species; however, we feel significant results may be
obtained from as few as 60 characters.

Our use of the word "character" will require some elaboration.
In its commonest taxonomic usage, a character is any feature of one
kind of organism that differentiates it from another kind. Thus the
red abdomen of one bse is a character distinguishing it from another
bee with the abdomen black. In this paper we use the word in a
second connotation only; that is, as a feature which varies from one
kind of organism to another. Now, to use the above example, ab-
dominal co'or is the character, which occurs in two "states" or alter-
natives, red and black.

Evaluating Systematic Relationships 1411

For each character the states were coded: 1, 2, 3, etc. In the
bee study the number of states per character ranged from two to
eight. Much variation in the number of states is undesirable from
the point of view of the methods discussed below. In the study we
undertook most characters had either three or four states. How-
ever, when variation exceeds desirable bounds it might be prefer-
able to divide the character state codes by a common denominator
or to normalize them.

The kinds of characters used in the bee study and the manner in
which they were coded are discussed at length by Michener and
Sokal ( 1957 ) . The possible effect of parallelism is also treated in the
same article. For purposes of the present paper the available data
might be summarized as follows: we have records of a given num-
ber ( n ) of species. For each species we have k records, k being
the number of characters considered in the study. The coded values
for any character may range from 1 to 9 depending on the number
of states in which this character occurs in the group under con-
sideration. As was mentioned previously it is desirable to have the
number of states not differ too widely for the various characters.
While it is not necessary to limit the number of possible character
states to nine, our particular computational setup was greatly fa-
cilitated by the use of a single digit code.

PROCEDURES

Character correlations and species correlations

Two obvious ways suggested themselves to the authors regard-
ing a procedure for deducing relationships from the character states
of a group of species. We could either correlate characters with
each other or species with each other. Since both of these methods
would lead to interpretable, although differing, results a brief dis-
cussion of the implications of the two approaches follows.

Sturtevant (1942) undertook a study of the genus Drosophila
with objectives and procedures somewhat similar to ours. He re-
corded 33 morphological, cytological and life history characters for
each of 56 species of Drosophila and two species of the genus
Scaptomyza. In his aim to develop a classification "as free from
personal bias as I could make it," Sturtevant set up two tables. The
first was a table of the total number of differences with respect to the
33 chosen characters between any two of the 58 species. These give
the degree of difference between the species concerned and are

1412 The University Science Bulletin

analogous to the complemental values of the "matching coefficient"
discussed in the section on Choice of a Correlation Coefficient be-
low.

A second table showed correlations between characters, expressed
as two-way frequency distributions. By examining the three high-
est character correlations Sturtevant found that six species con-
sistently fell into the exceptional classes of the two-way frequency
distributions. They were the two Scoptomyza species and four
species of Drosophila which he thereupon placed in separate sub-
genera. On the basis of the number of character differences be-
tween and within subgenera Sturtevant was able to confirm this
classification and arrive at some ideas on the relationships and ori-
gins of the various groups. He also performed a similar analysis on
29 characters of 40 genera of flies (Scatopliaga, Conops and 38 as-
sorted Acalypterae) to establish the relations of the family Droso-
philidae. Unfortunately the paper cited lists only summaries of
the above tables and it is therefore difficult to compare Sturtevant's
findings with ours.

Correlation between characters ( R-technique in the idiom of the
factor analysts) is the customary technique in biological and psy-
chological studies involving correlational analysis. In character
correlation matrices involving studies within one species each cor-
relation represents the sum total of the common forces acting on
any pair of characters. When analyzed by some method of factor
analysis, the matrix customarily yields a so-called general size factor,
a series of group factors affecting various groups of characters, and
residual specific factors affecting single characters only. The fore-
going is an example of a factor constellation involving morphologi-
cal characters and is not necessarily the only possible constellation.
As a matter of fact much psychometric work and the biometric
papers by Howells (1951) and Stroud (1953) use the method of
"simple structure" which a priori rejects solutions involving general
factors.

Regardless of the constellation preferred, the factors common to
two characters and causing them to be correlated could be visualized
as developmental forces, genetic or environmental in the final anal-
ysis. The range of these genetic or environmental forces is de-
pendent on the causes of variation within the sample of individuals
studied. Thus a sample of individuals from an inbred, isogenic, line
of animals would yield character correlations reflecting common
nongenetic, physiological (i.e., caused by microecological dif-

Evaluating Systematic Relationships 1413

ferences) factors only. Another sample comprising individuals from
various races or subspecies would provide correlations based on
common factors representing (1) genetic differences between in-
dividuals; (2) genetic differences between races; (3) nongenetic
physiological differences between individuals and (4) nongenetic
ecological differences between races. One of the authors ( R. R. S. )
has been able to accumulate a series of character correlation mat-
rices from various organisms representing these levels of variation.
Matrices on correlation of aphid characters within galls (clones)
and between galls have been published ( Sokal, 1952 ) while similar
matrices on aphid correlations between localities and morphological
correlations within and between strains of houseflies and Drosophila
await suitable analysis and publication.

When the sample transcends the bounds of the species the fac-
tors behind a character correlation matrix take on new meaning:
They now represent genetic divergence or the results of evolution-
ary processes. In the one case they were ontogenetic forces, in the
other they are phylogenetic forces. This type of analysis was
pioneered by Stroud (1953) who analyzed correlations of 14 char-
acters for soldiers of 48 species and imagines of 43 species of the
termite genus Kalotermes. He was able to interpret some factors
extracted from his correlation matrices as recognizable evolutionary
trends.

Another method of correlational analysis is called the transposed
matrix method or the Q-technique (as compared with the R-tech-
nique of character correlations, discussed above).3 It consists of
correlations between individuals based on measurements of char-
acters which they have in common. In psychology this involves
correlations between persons based on scores for common tests
which these persons have taken. In the Q-technique we are in effect
dealing with the same kind of raw data as in the R-technique, but
we compute the correlation coefficients by summing squares and
products at right angles to the direction previously taken (or we
transpose the matrix before computation which amounts to the
same thing ) .

A Q-technique correlation coefficient in a study correlating in-
dividuals of one species represents common forces or factors acting
on the two individuals concerned. In this case we cannot speak of
the "sum total of common forces" as we could in the case of the

3. In a recent paper Cattell (1954) has suggested restricting the Q and R symbolism
to studies involving factor analvsis and proposed Q' and R' for studies, such as the present

one, employing more superficial methods

1414 The University Science Bulletin

R-technique. Insofar as the characters used are indicative of the
entire spectrum of potential variation of the individuals we can
say that the resulting correlation coefficient is representative of the
real affinity between two individuals. When scanned for clusters
of high correlation coefficients the Q-type matrix reveals types of
individuals which are similar. It is thus especially suited to classi-
ficatory problems. When subjected to factor analysis the resulting
factors are now of a different nature. The general size factor has
been lost and in its place we find a general taxonomic group factor
which accounts for the overall correlations of all the individuals in
the study.

When, as in the present study, the correlation is between species
of a taxonomic unit the general factor is a general systematic factor
denoting overall relationship within the systematic group. The
species having the highest factor loading would be most representa-
tive of the group. Other factors would describe subgroups within
the systematic unit and describe the relationships of these subgroups
with each other and of the species to the subgroups. It should be
clear from the above that for purposes of biological classification
the relationships represented by a Q-technique matrix are more
meaningful by far than are those of a R-technique matrix. Except
for the above-mentioned work of Sturtevant ( 1942 ) which involved
not correlations but character differences, the only Q-type study
in systematics of which the authors are aware is in a publication
by one of them ( Sokal, 1958 ) containing factor analyses of selected
portions of the present data. A number of the phytosociological
coefficients of association and similarity can be considered as of the
Q-type.

Psychologists have used Q-technique repeatedly (e. g., Burt 1937,
Stephenson 1936), although R-technique is still preferred in most
studies. Cattell (1952) has listed 5 points of criticism of the Q-
technique. It is appropriate that we discuss briefly their relation to
the problems under study here. The first objection is that Q-tech-
nique loses the general size factor, yielding in its place a common
species factor. This latter is claimed to be trivial by Cattell, and
correctly so, for psychological work. However, in a matrix of cor-
relations between species such a general systematic factor delineates
the relation of individual species to the taxonomic group and in-
dicates the proportion of the variance of each species explained by
the general systematic factor.

Cattell's second objection to Q-technique is that it is unreasonable
to assume simple structure in the factorization of a Q-matrix. The

Evaluating Systematic Relationships 1415

authors agree with this argument, but for the purposes of the pres-
ent paper it is not important since they are not here undertaking a
factor analysis. Furthermore, they feel that simple structure ( i. e.,
each factor having essentially zero effects on certain species) is
not necessarily a very suitable constellation for many biological
factorizations.

The third objection refers to a customary shortcoming of Q-
matrices. They are based on few individuals and generalizations
about the entire population are drawn from them. In this study,
the matrix is of course of more than adequate size. Furthermore
our conclusions are not intended to extend to species not included
in our study.

It is true that the species recorded are an eclectic sample from
those extant in the world today. On the other hand we are of course
dealing with a sample obtained by natural selection from the multi-
tude of species or specieslike entities that have existed since the
origin of the four genera of this study. Hypotheses regarding these
extinct species will be valid only insofar as recent species reflect the
course of evolutionary history.

Another point in connection with the third objection is the num-
ber of characters employed. True relationships will become
apparent only insofar as the characters adequately represent the
sources of variation within the species.

A fourth objection relates to the lack of equivalence in recording
and interpreting the factors from the Q- and R-matrices. It com-
pares the relative permanence of psychological tests with the rela-
tive impermanence of persons. In this study we are confronted
with characters and species varying in their relative permanence,
but both equally permanent when based on the time scale of the
scientist investigating them.

The fifth criticism, labelling the Q-technique as descriptive rather
than predictive, again is invalid when applied to the present data.
Since the purpose of the study is historically descriptive and one of
our aims is to divide the population of species into categories, the
technique's fault for psychological research becomes a virtue in our
field of investigation.

There are two evolutionary situations under which it is important
to examine the two types of matrices. The first might be referred
to as breakage of correlation. It occurs when in a certain evolution-
ary line two characters that were correlated in ancestral lines and
are still correlated in related lines become independent of each

13—8050

1416 The University Science Bulletin

other. Under such conditions the R-matrix is a poor representation
of the true relation between the two characters. There is no good
way of representing such a correlation, close in one line, absent in
the other. On the other hand a Q-matrix is not affected by such
data.

Convergence of species for a number of characters is a second
disturbing phenomenon. Here the R-matrix is not affected while
the Q-matrix is affected if the convergent characters outweigh the
nonconvergent ones in numbers.

We do not believe this is likely if an adequate number of char-
acters is studied. In case of a preponderance of convergent char-
acters and in the absence of paleontological data it is doubtful
whether the systematists would be able to distinguish convergence
from relationship by descent.

From a consideration of the above arguments it follows that given
the objectives and material of the present study the Q-technique is
to be preferred to the R-technique and the objections made by
Cattell to the former method do not apply to our case. However,
besides the theoretical reason for adopting the Q-technique as re-
flecting relationships between species there were several practical
reasons for so doing. The problem of finding a suitable type of
correlation coefficient between characters would have been formid-
able in view of the coding system adopted. Since some of the char-
acters were present in two states only while others were present in
as many as eight states, there would probably not have been any
one type of correlation coefficient for all possible character com-
binations. A matrix based on correlation coefficients of different
types would be far from desirable. Furthermore, uniformity of
computational procedure was essential to efficient handling of the
data by International Business Machines (IBM) equipment.

Not to be underestimated is the saving in computation resulting
from adoption of a 97 x 97 species correlation matrix vs. a 122 x 122
character correlation matrix. The former requires the computation
of only 4656 correlation coefficients while the latter would neces-
sitate 7381 such coefficients.

The choice of a correlation coefficient
As a next step a suitable correlation coefficient had to be chosen
to represent the correlations between species. There were serious
considerations against the use of the product-moment correlation
coefficient since the variables (species) are anything but normally
distributed. Table 1 presents frequency distributions of state codes

Evaluating Systematic Relationships 1417

Table 1

Frequency distributions of state codes for the characters of species 19, 56,
83 and 84.

State Sp. 19 Sp. 56 Sp. 83 Sp. 84

code f f f f

1

54

56

48

46

2

31

40

42

41

3

31

14

23

26

4

3

11

7

6

5

2

1

2

2

6

1

7

1

vf 122 122 122 122

for four representative speeies. The distributions are highly asym-
metrical. Those for species 19 and 56 approach Poisson distribu-
tions for their means when the class codes are reduced by one.
Any interpretation of this agreement is dubious, however, in view
of the variable number of states possible per character.

Other correlation coefficients were considered and rejected. The
correlation ratio, 77, is unsuitable since x?7y does not necessarily equal
7rjx. Tetrachoric r would have lost some of the information avail-
able because it would necessitate reducing all characters to two
states. Furthermore the theoretical assumptions of underlying
normality essential to correct application of the tetrachoric correla-
tion coefficient cannot be defended for all characters.

Another method of demonstrating an association between species
would be the very simple one of counting the numbers of matches
in states for the 122 characters of any pair of species of bees and
then dividing this number by 122, the highest possible number of
such matches. The results for species 19, 56, 83, and 84 are shown
on table 2 where these "matching coefficients" are compared with
product-moment correlation coefficients. The "matching coeffici-
ents" are somewhat higher than the correlation coefficients but
resemble them in relative magnitude. In spite of this fact, "match-

Table 2

"Matching coefficients" ( below diagonal ) and product-moment correlation
coefficients (above diagonal) between species 19, 56, 83 and 84.

19 56 83 84

19

X

.40

.37

.37

56

.52

X

.47

.38

83

.53

.61

X

.93

84

.50

.54

.87

X

1418 The University Science Bulletin

ing coefficients" were not used since they have an unknown sampling
distribution, they distort resemblances by counting a 3 to. 4 mis-
match the equal of a 1 to 7 mismatch, and finally they would
have been harder to handle by the IBM equipment available to us.

Lacking a more suitable means of correlation we adopted the
product-moment r, in spite of nonnormal distribution of variates
and possible heteroscedasticity. Various ways of improving the
distributions by means of transformations were tried. Table 3 shows
the same correlation coefficients as the upper half of the matrix of
table 2, but based on -yj X and \/ X -|-.5 transformations. The
slight differences obtained do not justify the extra computational
labor involved.

We have already briefly touched on the desirability of coding the
data in such a way as to put all character states on the same scale.
In a character with two states the code 2 indicates a situation dif-
fering greatly from one given by code 2 in a character with 7 states.
This problem is also encountered in Q-matiices in psychology where
the scores for different tests are often not in comparable units. This
situation is usually met by normalizing the rows (tests, or in our
case characters ) of the raw score matrix. The authors did not per-

Table 3

Product-moment correlation coefficients between species 19, 56, 83 and 84
based on variates coded as VX (below diagonal) and as yX+.5 (above
diagonal). Compare with uncoded product-moment correlation coefficients in
table 2.

19 56 83 84

19

X

.42

.36

.37

56

.42

X

.50

.41

83

.36

.51

X

.93

84

.37

.41

.93

X

form this transformation since ( 1 ) it would have removed the com-
mon systematic factor from the matrix of correlations and would
thus have lowered the correlation coefficients considerably; (2) ap-
plication of the character state codes does standardize the data to
a certain extent because 76 percent of the characters have either
three or four states and only 3 percent have six or more states; (3)
although the additional labor of normalizing the variates would not
have been excessive the amount of IBM work involved in comput-
ing correlation coefficients would have been prohibitive, since a
one-digit code would not have sufficed for normalized data.

The authors are well aware that their methodology of coding and
correlation could profit by refinement. It is, however, our point of

Evaluating Systematic Relationships 1419

view that in a pilot study of this nature such refinements are pre-
mature. Should the general method prove of value, significant re-
sults will surely emerge in spite of minor imperfections in technique.

Computation

The computation of a large matrix of correlation coefficients such
as the 97 x 97 bee matrix presents serious technical difficulties. Only
high speed electronic computing machines are able to perform this
operation with real dispatch. At the time our bee data were being
processed we had only punched-card tabulating machines at our
disposal. It might be noted here that a computational operation of
this magnitude cannot reasonably be undertaken without some auto-
matic computing facilities. The equipment used by the authors is
that available in the University of Kansas IBM laboratory: a card
punch ( type 26 ) , a verifier ( type 56 ) , an accounting machine ( type
402) and a reproducing machine (type 514).

The computational problem was simplified somewhat by the fact
that the variates consisted of single digits only. This increased the
number of variables that the machine could process simultaneously.
Each IBM card represented a character with the state code of each
species for the particular character listed in separate columns. Since
there are only 80 columns per card, it was impossible to record all
species on any one card. A different approach was therefore
adopted and the card divided as follows:

Column 1 — Project code

Columns 2-4 — Character code number

Column 5 — Deck code (explained below)

Columns 6-8 — Left blank for possible subsequent use

Columns 9-44 — Multiplier columns for 36 species

Columns 45-80 — Multiplicand columns for 86 species.
The 97 species were divided into group I for species 1-36, group II
for species 37-72 and group III for species 73-97. Since group III
used only 25 columns another 5 columns were taken up by a repeti-
tion of data on species 1 through 5, which we used as a check on
computational procedure. Six decks of 122 cards each, one card
per character, were then prepared. The decks were constituted as
follows :

Deck

Multiplier

Multiplicand

1
2
3
4
5
6

Group I
Group II
Group III
Group I
Group II
Group I

Group I
Group II
Group III
Group II
Group III
Group III

1420 The University Science Bulletin

Different card colors besides a punched code were used to dis-
tinguish the decks.

By running these decks in succession through the tabulator we
were able to reduce rewiring of the board to one half of what it
would have been with the minimum number of decks (3).

The method of arriving at the 2x2 and 2xy was the customary
one of progressive digiting with interspersed "X-cards." Running
time on the 402 tabulator was some 24 hours. Punching and verify-
ing of the cards had taken a similar amount of time. Thus the
preparation of the 2x2 and 2xy for the entire matrix took about a
week. These values were computed for a half-matrix only. How-
ever, a test deck and five test variables detected wiring errors and
machine malfunction with a reasonable limit of safety.

The next step was the computation of the correlation coefficients.
This was done by computers using desk calculators.4 The matrix of
squares and products was subdivided into manageable sections, 30
variables (species) square. All computations were checked by a
different computer and, where possible, by different steps. The
computational procedure employed was the customary L method.5
It does not seem necessary to elaborate on the details of this method.
Any good textbook of statistics will contain a section on the com-
putation of a product-moment correlation coefficient. Furthermore,
each computation center has its own setup for correlation coefficients
depending on the capabilities of the machines and thus no general
account need be presented here.

The correlation coefficients were calculated to four significant
decimal places and entered on a matrix. Three decimal places
would have been quite sufficient for this study; however four were
computed in case later statistical work required greater refinement.
Total computation time for this phase of the work was 160 man-
hours. It should be emphasized that the time estimates given above
refer to the relatively simple equipment available to us. Digital
computers are now available which would handle the entire com-
putation, from raw data to completed correlation matrix without
human intervention in less than an hour. This would be only one-
two hundredths of the time it took us to compute the same informa-

4. The writers at this point wish to express their appreciation to Misses Betty Becker,
Marion Clyma, Jacqueline Johnson, Nonnandie Morrison, and Messrs. D. A. Crossley, Jr.,
Ralph Jones and Roger Price for their conscientious assistance with IBM work and desk
computation.

5. rxy = Lxy / VU VL7> where Lxy = N2XY — 2X2Y and

Lx = N2X2 — (2X)2, Ly = N2Y2— (2Y)2.

Evaluating Systematic Relationships 1421

tion! With every passing year electronic computers are becoming
more efficient and more widely distributed. Thus the computational
aspects of our method will become a progressively less important
impediment.

Since the matrix of correlation coefficients was unwieldy (it also
had to be subdivided into sections ) and since further work with the
correlation coefficients was contemplated, the latter were punched
on 4656 IBM cards, one to a card. These cards were duplicated by
means of the reproducing punch in order to obtain cards for a com-
plete matrix of 9312 correlation coefficients. Information on these
cards included matrix row and column numbers for the particular
correlation, the coefficient with sign, and a class code for the co-
efficient. These class code numbers (1-22) represented 22 classes
of a frequency distribution of the correlation coefficients arrayed
in ascending order of magnitude with class intervals of .05. In ad-
dition, the cards contained codes for the relationship between the
two species involved as evaluated by conventional systematic meth-
ods (by CD. M.).

The correlation coefficients on punched cards have so far been
put to the following uses: We have compiled a printed tape record
of the full matrix, column by column, which has been very useful
for reference and further computation. Another tape has been com-
piled giving a listing and frequency distribution of the correlation
coefficients grouped in the 22 size classes. This tape has been of
great value in various approaches to a classification of the relation-
ships demonstrated by the matrix. A third tape lists the sums of
the correlation coefficients, column by column. This has been nec-
essary for the B-coefficient method briefly described below. A
fourth tape presents a two-way frequency distribution showing the
relation between correlation coefficients and the relationship code
developed by conventional systematic methods. These tapes were
prepared in a few hours running time from the correlation coefficient
cards, which we still expect to use in a variety of ways.

The matrix of correlation coefficients
In the bee study the 4656 correlation coefficients computed in
the above manner ranged in magnitude from — .0626 for the cor-
relation between species 26 and 92, to .9747 for the correlation be-
tween species 43 and 44.6 As was mentioned previously, a fre-

6. For lack of space the matrix cannot be reproduced here. Microfilm or IBM-tape
or card copies can be obtained through the Secretary, Department of Entomology, Uni-
versity of Kansas. Lawrence.

1422 The University Science Bulletin

quency distribution of these coefficients, grouped into 22 classes
with class intervals of .05 was set up. The modal class showed a
class mark of .38; this represents the most frequent class of cor-
relation coefficients found between species in this study. How-
ever, a second mode was located at .78. This bimodality would
indicate that we are dealing with two populations of correlation
coefficients: those indicating close, possibly intrageneric relations
and others representing more distant relations. Codes representing
Michener's previous views on the relationships among the species
were correlated with the above coefficients. The single correla-
tion coefficient between the correlation matrix and Michener's codes-
was .80. It was encouraging to find that magnitude of the corre-
lation coefficients in our matrix was apparently an estimate of
systematic relationship as indicated by the previous classification.

Another way of examining these correlation coefficients is to study
frequency distributions of the coefficients for any single species
against all other species. By this means we were able to distinguish
members of closely related groups of species from isolated species
within a genus and these in turn from very isolated species represent-
ing monotypic genera or subgenera. For a detailed discussion and
illustrations of this procedure, the reader is referred to Michener
andSokal (1957).

The absence of significant negative correlations from our matrix
requires some discussion. Q-technique matrices of correlations be-
tween people (based on psychological tests) are quite likely to
yield such correlations. If there are distinct, antithetical types of
persons represented in the matrix, such as extroverts and introverts,
it is likely that a high score for one type will be a low score for the
other and vice versa. In our case evolutionary progress may be
represented by either an increase or a decrease in state codes. In
the majority of characters the supposedly primitive situation is an
intermediate state code with two diverging evolutionary trends rep-
resented by the lower and higher code numbers. Furthermore,
characters representing correlated trends were not necessarily coded
along the same scale or in the same direction. It is clear that under
such circumstances distantly related forms are likely to be uncor-
rected rather than negatively correlated.

The search for group structure

The matrix of correlation coefficients between species can be put
to a variety of uses and the analysis reported below represents
merely an initial effort at an exploitation of the data. The correla-

Evaluating Systematic Relationships 1423

tion coefficients serve as an absolute measure of relationship be-
tween any two species in our study, limited only insofar as the
characters chosen do not represent the total correlated variation of
the two species.

The search for structure among the correlation coefficients of the
matrix is of course no different in aim from the search by the sys-
tematist for a natural system in an array of species. Such a system
consists of a hierarchy of groups. Various methods can be used for
discovering a hierarchy in data such as ours. A customary, rather
simple device of the psychometrician is so-called "cluster analysis,"
developed to a fine art by Tryon ( 1939 ) .

A concise description of the procedure (the ramifying linkage
method ) is given in Cattell ( 1944 ) and Thomson ( 1951 ) . Because
of the simplicity of the procedure, cluster methods are used exten-
sively, although Cattell (1944, 1952) and others have pointed out
that cluster analysis cannot be considered a substitute for the more
involved factor analytic methods. Attempts to employ cluster anal-
ysis for finding structure in our matrix were only partially success-
ful, since the resulting clusters were partly overlapping, i. e., a given
species might be a simultaneous member of two clusters. This
makes good sense for intermediate forms in an abstract scheme of
relationships. In a systematic hierarchic classification, however,
groups at the same level have to be mutually exclusive for practical
as well as for theoretical reasons, except for low level groups exhibit-
ing reticulate evolutionary pattern (rare above the species level in
animals). A further reason for the unsuitability of cluster analysis
is the complexity of the clusters as more species are added to them.
Although clusters are therefore not convenient in an initial search
for structure, the diagram of relationships established by methods
to be described below could be easily recognized in the clusters
outlined by cluster analysis. A method essentially similar to cluster
analysis is the p- group and pF-group method of Olson and Miller
(1951) applied to three paleontological R-technique matrices. It
suffers from the same drawbacks as cluster analysis.7

7. After the present research and manuscript had been completed one of us ( R. R. S.)
became acquainted with the psychometric work of Professor Louis L. McQuitty of the
Michigan State University, who in recent years has developed a whole battery of refined
cluster methods (McQuitty, 1955, and a series of papers in press in The British Journal
of Statistical Psychology, Educational and Psychological Measurement, and Psychological
Monographs). Several of these papers deal with psychological problems which are closely
related to those of biological classification. One of the methods invented by McQuitty
bears a close resemblance to our variable group method developed below. It is interesting
(as well as reassuring to us) that workers in different fields had unknown to each other
developed some of the same formulations. We hope to try some of McQuitty's other methods
on our material. They have the advantage of simplicity and can be programmed for elec-
tronic computation without much difficulty. Indeed the time may not be far off when
computation for a study such as our bee work will be a minor matter routinely handled by
a computing center in a very few hours and the remaining problem will be the collection
of data for the machine and the interpretation of the voluminous answers that are produced.

1424 The University Science Bulletin

As a technique for grouping the species we experimented exten-
sively with the coefficient of belonging (B-coefficient) of Holzinger
and Harman (1941). It is the sum of the correlations among the
members of a group divided by the sum of the correlations of these
group members with the other variables ( species ) of the study.

Results of our B-coefBcient analyses for the bees were reasonably
good, as judged by the previous classification and by our subsequent
investigations. There was one main drawback, however. Large
species groups showed a lack of structure and relatively low B-
coefficients which would make the species in these groups appear a
good deal less related to one another than members of groups of
two or three species. The cause of this phenomenon is not hard to
find. In large species groups the denominator of the B-coefBcient
would include high correlations due to correlations of group mem-
bers with numerous other prospective members not yet included in
the group. This would tend to depress the B-coefficient values. By
the time all such members have been admitted to the group, it has
become so large that even the admission of a relatively unrelated
variable will effect the B-coefficient only slightly.

In view of the disadvantages of the B-coefficient we developed
our procedure which is presented below in a general manner to-
gether with some of the reasons for its adoption. This presentation
is followed by a detailed step-by-step account of the computational
procedure for readers who wish to become more familiar with it.

A nucleus of a group was established, using the two species hav-
ing the highest coefficient of correlation. Then species would be
added to this nucleus, one at a time, always adding first the species
having the highest average correlation with members of the group.
The limit of the groups could be found by decreases ( L„ + t - Ln )
in the level of the average correlation Ln, where the subscript refers
to the number of members in the group. As in the B-coefficient a
significant drop is empirically determined since sampling distribu-
tions of average correlations, such as L„, are unknown. By develop-
ing first lower groups (species groups), then by the same method
grouping these into larger groups (sometimes subgenera), and
these into still larger or higher groups, etc., it has been possible to
develop a hierarchy of groups for which the diagram of relationships
( figure 1 ) can serve as a representative. Each number in this figure
represents a different species; for a list of the species concerned see
Michener and Sokal (1957).

Since Ln is not amenable to rigorous statistical treatment it was
decided to recompute correlation coefficients ( using Spearman's sum

Evaluating Systematic Relationships

1425

«

E

u

£ J

b

o

a
E

— x

'2 a, -

° r M

f <U O

X « :

«- 2 -°

O .- 3

iooo n

900-

800-

700-

600-

500-

♦ 00-'

HI II 1

63 63 65 6T 70 75 7T 83 78 7J 14 85 SB 90 62 94 94 97
T6 66 Eg 71 f2 82 81 74 80 73 8T 89 91 83 35 8t

T

X

LN

&d

u

u

ANTHOCOPA

PROTERIADE5,

29,30 HOPLITIS, 40 &41.

\ ^

Fig. 1. Diagram of relationships for the genus Ashmeadiella obtained by
the weighted variable group method. Ordinate: magniture of correlation coef-
ficient multiplied by 1000. Exact correlations between any two joining stems
can be found by reading the value on the ordinate corresponding to the hori-
zontal line connecting the stems. This value becomes approximate and maxi-
mal in cases of multifid furcations. Broken lines used where more than three
stems join are for convenience only; the horizontal connecting line has the same
significance as elsewhere. "Roofs" over species numbers at the summits of the
lines delimit subgenera containing more than one species, as based on C. D. M.'s
previous findings and not on this study. Generic names are in small capitals.
The horizontal broken lines are not relevant to the present account; they are
explained by Michener and Sokal (1957).

of variables method ) after the group limits at each hierarchic level
had been reached. Thus we returned at the end of each grouping
procedure to a new matrix of correlation coefficients about which
confidence statements might be made. Two further considerations
in the final choice of a method for grouping remain to be mentioned:
We might have admitted only one new member for each group
at a given hierarchic level, thus obtaining a diagram of relationships
consisting of bifurcations only. We have called this method the

1426

The University Science Bulletin

pair-group method as contrasted with the variable-group method,
where any number of new members can be admitted to the group at
any one hierarchic level, the limit of the group being determined by
a significant drop in Ln. The pair-group method has some theoreti-
cal justification in that much evolutionary ramification is believed
based on speciational processes involving the splitting of one species
into two. However, there must also occur some speciation as a re-
sult of the splitting of a species into more than two isolates and, on
the assumption of equal evolutionary rates for these new lines, the
pair-groups method would fail to represent the true situation. More-
over, many of the groups must be markedly different, not merely
because of divergence, but because of extinctions of intermediates.
A group might be broken into any number of different subgroups
by different extinctions. Furthermore an empirical study of this
method ( see fig. 2 for an analysis of relationships in the subgenera
Chilosima and Ashmeadiella by the pair-group method and com-
pare with the left side of fig. 1 for the variable-group method) dem-
onstrates that in spite of the pair-group device we are forced into
multified furcations by drops in Ln too small to plot or by temporary

1000-n

900-

800-

700

600 -1

63 69 65 67 70 76 77 82 78 79 84 73
64 75 66 68 71 72 83 81 74 80 85

u

t

I _ _ _ _

Fig. 2. Diagram of relationships for the subgenera Chilosima (63-64) and
Ashmeadiella s. str. ( 65-85 ) , obtained by the method of pair-groups, i . e. dia-
grams would ideally consist of bifurcations only. Stems have been weighted.
Explanatory comments as for figure 1.

Evaluating Systematic Relationships

1427

A B C D

ABCDEFGH

a,

U

W X

Fig. 3. Hypothetical diagrams of relationships to illustrate effects of different
methods of weighting stems. For explanation see text.

reversals of Ln values, discussed below. Thus the variable-group
method was adopted as the more reasonable and flexible of the two.
A second consideration is how to weight the variables during the
recalculation of the correlation matrix after each grouping pro-
cedure. A simple diagram (fig. 3a) will make this issue clear. A
and B represent the two species with the highest correlation coef-
ficient. The Ln for C against A and B is significantly below rab, so
that A and B are represented as being closer to each other than they
are to C. When studying the relation of a fourth species D with
group ABC we face the following problem: Should we calculate the
correlation of ABC against D with A, B and C equally weighted or
should we weight A = B and AB = C? Rephrased biologically, the
problem is whether to relate species D with the homogeneous group
ABC, or with the stem AB-C, where C carries as much weight in
determining the relation with D as do A and B together. Although
in a simple case, such as the one described above the two alternatives
may not produce very different results, in a situation such as de-

1428 The University Science Bulletin

picted in fig. 3b species H might be weighted as % of the group A-H,
or J2, depending on the system adopted. Similarly species B would
be weighted Y% in the former case but only ^2 in the latter case.
When dealing with fairly large groups the second method would
therefore reduce the weight of the early admitted members and in-
crease the weight of those species admitted later.

The same problem is found in a situation such as shown in fig.
3c. By the first method species T is weighted %, by the second
method it is weighted only M.4. Neither of the two methods is en-
tirely satisfactory. By method one we are reducing the importance
of species H and Z in representing groups A-H and S-Z respectively.
If the relationship diagrams of figures 3b and 3c depict true phylo-
genetic relationships, then H and Z should represent half of their
respective lines regardless of subsequent diversification in the other
halves. On the other hand giving relatively greater weight to single
late arrivals also gives heavier weight to specialized features of such
species and thus would tend to distort the relational pattern, while
specializations in the diversified branch of the stem tend to cancel
each other, permitting a better average picture of the groups to
emerge. The optimal system of weighting would be one between
these two extremes, weighting each species according to its number
of generalized and specialized features. This is clearly impossible
without renewed introduction of a subjective element into our pro-
cedure. We therefore adopted the second method, i. e., the weight-
ing of new members as equal to the sum total of all old group mem-
bers, thinking it to be the less objectionable of the two. We feel
that this method will represent stems more correctly and that bias
introduced by specializations of late joiners will be kept down by
the large number of characters considered in our study.

We are reassured in our decision by the results of a comparative
study on the subgenera Ckilosima and Ashmeadiello. Figure 4
shows the results of a variable-group analysis of these subgenera by
weighting method one, while results by method two can be seen in
the left side of the diagram of fig. 1. General agreement as to re-
lationships and level of furcations is very good. The main difference
between the two diagrams is that in method one group 77-81 8 first
receives 79 before receiving group 67-72, 78 and 84, while in method
two it first receives group 67-72, then 78 and then 79 among others.

8. In the interest of brevity groups will be identified by their leftmost ( in the diagram )
and rightmost members with a dash separating the two. Thus 77-81 means group 77, 82,
83, 81. It clearly does not include all species ranging in number from 77 through 81,
and includes some beyond that numerical range.

Evaluating Systematic Relationships

1429

1000-1

64 75 66 68 71 72 82 81 78 74 85
63 69 65 67 70 76 77 83 79 84 80 73

900-

800-

700-

u

600 -J

Fig. 4. Diagram of relationships for the subgenera Chilosima (63-64) and
Ashmeadiella s. str. (65-85), obtained by the variable group procedure under
weighting method one, i. e., equal weights for all stems. Explanatory com-
ments as for figure 1.

Careful examination of the original correlation coefficients makes
the reasons for these differences clear. Group 77-81 is closer to 79
than to 78 except that 81 is closer to 78 than to 79. Also 81 is closer
to 67-72 than are 77, 82, and 83. Therefore in method two, where
81 receives as much weight as 77, 82 and 83 together, 67-72 joins
the nuclear group first. This is also partly due to the fact that un-
equal weighting of the species in the 67-72 group favors those close
to the 77-81 group. Since 78 is closer to 67-72 than 79, the latter,
while originally quite close to 77-81 is now temporarily delayed and
78 joins the combined group 67-81 before 79 does. These relations
are at too low a phyletic level to be included in the original diagram
of relationships drawn by C. D. M. who feels that there is little that
can be obtained from classical systematic studies of these species to
suggest whether method one or method two is preferable. In view
of the small over-all differences between the two methods and
especially in view of the fact that the lines concerned all join by
either method with a difference in correlation coefficients of less
than .06, it may well be that we have made too great an issue of the
matter. In any case we feel confident that weighting method two
will present us with a reasonably bias-free picture.

1430 The University Science Bulletin

The Weighted Variable Group Method

It was thought advisable to give a detailed account of our method
in order to enable readers to repeat the operations should they so
desire. The subgenera Chilosima and Ashmeadiella, which have
been used as a testing group before, will serve as an illustrative
example. These subgenera include species 63 through 85 (see
figure 1 ) .

Correlation coefficients among these 23 species are shown in table
4. All values are significant with probability values of less than one
percent. The highest correlation coefficient among these species is
.965 for 67 x 68.9 This is also the highest correlation involving either
of these two species. The next to enter group 67-68 is species 70
which has the greatest average correlation ( Ln = .892 ) with 67 and
68 since 67x70 = .896 and 68 x 70 = .889. No other species in the
study has as high an average correlation with 67-68, as can be
learned from a few trials. We established empirically, as a result
of numerous trials, that a drop in Ln of .030 gave a satisfactory limit
for groups; therefore 70 is not to be admitted to group 67-68 at this
particular time. Another high correlation involving species other
than 67 and 68 is 77 x 83 — .951. This is also the highest correlation
for the two species concerned. Next to join this nucleus is species
82 with an Ln value of .936. The drop is less than .030; therefore
82 is admitted. Next to join is species 79 with an Ln value against
77, 82 and 83 of .905. There is now a significant drop from the
previous L„ value and 79 is excluded for the time being. Drops in
Ln are always measured from the previous Ln, not from the initial
Ln. Our second group is therefore 77-83. In a similar manner we
established groups 63-64, 65-66 and 71-76, each consisting of only
two species.

So far only 11 species out of the 23 of the study have been placed
into groups. A systematic survey was then made of the remaining
12 species to see if any group had been missed. For example, ex-
amination of species 69 revealed that its highest correlation was
with species 72 ( 69 x 72 = .820 ) . However, this latter value was not
the highest correlation for 72, since 72x76 = .904. Thus 72 might
eventually join the group containing 76 and 69 might join the group
containing 72, both of which events came to pass at a later stage of
the analysis. At the present time, however, species 69 and 72 are
left unattached to any group. Similarly the remaining ten species
in the study were shown not to belong to any nuclear group. To

9. We shall use this symbolism in place of the more formal rar.es

Evaluating Systematic Relationships

1431

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1432 The University Science Bulletin

set up one of the latter we required a correlation coefficient which
was the highest one for both participating species (i. e., the re-
ciprocally highest correlation).

After the groups had been delimited a new correlation matrix
was computed considering the newly formed groups as single
variables, i. e., the previous matrix of 23 variables (matrix 1) was
reduced to one of 17 variables (matrix 2). It is self-evident that
the only correlation coefficients in need of recomputation were those
involving new groups. Correlations involving only species that had
remained single were not altered in any way. As a matter of fact,
a procedure was devised by means of which the correlations were
not even recopied, but variables joined into groups were crossed
out and the new group variables were entered along the margins of
the old matrix. The actual computational procedure is quite simple
and considerably less complicated than the computations for finding
the original correlation coefficients. It is described in the paper by
its originator (Spearman, 1913) and also by Holzinger and Harman
( 1941 ) . Let us illustrate this method by computing the correlation
between groups (63)x and (67)i10. The general formula for this
computation is

□ qQ •

r,.Q

V q + 2Aq V Q + 2AQ

where Q qQ is the sum of all correlations between members of one
group with the other group, Aq is the sum of all correlations be-
tween members of the first group, AQ is a similar sum between
members of the second group, q is the number of species in group
one and Q the number of species in group 2. Thus in this particular
case □ qQ equals (63x67) + (63x68) + (64x67) + (64x68)
= .692 + .682 + .681 + .672 '= 2.727; Aq in this case equals only
(63x64)— .908 while AQ equals 67jc_68 = .965, since each of these
groups consists of two species only. In cases where a group con-
sists of 3 species, for example, the A term consists of the sum of
(1x2) -f- (1x3) -f- (2x3). In the present case q = Q = 2 species.
Substituting into the formula given above:

2.727

(63)]x(67)1 = — — —————— = .704

V 2 + 2(.908) V 2 + 2(.965)

10. The notation (63)i, refers to the group of species formed in matrix 1, the lowest
numbered member of which is species 63, i. e., to group 63-64. Similarly (67)i, refers to
67-68, and (77),, to 77-82-83.

Evaluating Systematic Relationships 1433

These computations can be set up in a systematic manner and are
then neither particularly complicated nor time consuming. In the
special case where we wish to calculate the correlation coefficient
between a single species (x) and a new group (q), the formula is
amended as follows :

Si
X/ r„

LX-i[

V q + 2Aq

An illustration is the correlation of species 69 with group (77)^ 2rx.q
equals ( 69x77 ) + ( 69x82 ) + ( 69x83 ) = .764 + .788 + .738 = 2.290,
while Aq=( 77x82) + (77x83 ) + (82x83) =.925+.951+.948=2.824.

2.290
Then 69x(77)1 = -— = .779

V 3 + 2(2.824)
In such a manner a new 17 x 17 correlation matrix ( matrix 2 ) was
constituted. From this point on the species groups [(63)15 (65)^
(67)l5 (71)1? and (77) J were tested as though they were single
species.

Once matrix 2 had been computed the identical grouping pro-
cedure was followed. Group (71)x had a mutually highest correla-
tion with species 70 at .923. They were then joined by 72 and group
(67 )1 at Ln levels of .903 and .885 respectively. These affiliations of
72 and ( 67 ) j were also their highest correlations. The next prospec-
tive joiner was species 81 at Ln = .859, i. e. not quite the established
drop of .030. However species 81 had highest relations not with the
previous species but with group (77)x with a correlation of .923.
Therefore it was excluded from consideration as a candidate for the
earlier group and the runner up, species 78, used instead. The latter
gave an L„ value of .844, clearly a significant drop from .885. Species
81 meanwhile was used in a nucleus of a new group 77-81 [( 77 )2].
Situations such as the above were the exception. In general the re-
lations and choices were entirely straightforward and could be left
to the discretion of the computing assistants.

At the end of each grouping procedure the remaining single
variables were checked to avoid missing groups with low correla-
tions between members. With each grouping procedure the matrix
of correlation coefficients became smaller and the job of recomputa-
tion less. The weighting procedure adopted by us was automatic
in that all correlation coefficients used were from the previous matrix
and not the initial one. It took eleven matrices to obtain a single

1434 The University Science Bulletin

group out of the 23 species of the two subgenera. This amount of
work could have been reduced by raising the minimum recognized
difference in Ln level above .030 but there would have been a re-
sulting loss of detail in the diagram of relationships. Conversely,
however, reducing the recognized difference below .030 would not
have increased the meaningful detail, since even .030 was too small
to prevent the occasional reversal of r values discussed below.

Once computed, the relations were represented as diagrams of
relationships as in figure 1. The ordinate at the left of each diagram
is graduated in units of 1000 x r. The correlations between any join-
ing stems in the diagram can be read by measuring the level along
the ordinate of the horizontal line connecting the stems. Thus
species 63 and 64 are correlated at a level of .908, while group 63-64
is related to group 67-72 at .702. Furcations involving more than
three lines are shown by broken lines converging on the midpoint
of the horizontal line as in group 88-96 of the above figure. The
tops of the figures are at a level of 1000 (correlation of 1) since
obviously each species is perfectly correlated with itself.

In cases of groups of only two stems the L„ level corresponds to
the correlation coefficient of the two stems. When more than two
stems join to form a group the highest Ln level was graphed for all
group members. Thus while 77 x 83 equals .951, Ln for 82 against
77 and 83 equals .936. The group of these three species (77-83) is
shown related at levels .951. Occasionally the correlation coefficients
for the same group in successive matrices will rise a little. Thus in
this same figure groups 63-64, 69-85, 87-96, and species 86 are shown
joining at .702. The first three groups actually joined at .671, but
species 86 which joined their group at the next matrix did so at
level .702. This type of situation, which occurred infrequently, might
lead one to express concern about the validity of the method, since
regular decreases in levels of correlation coefficients and Ln values
are expected. However, it can be shown from Spearman's formula
for the correlation of sums of variables that slight increases in the
levels of correlation coefficients of the sums of variables above the
correlations of their component variables are possible. For ex-
ample, if A and B have formed the nucleus of a group at ra.b = .9
and C is about to join them, then by the rules of the variable group
method both ra.c and rb.P must <ra.b = .9. It can then be shown
that r(nb).c must be <.925. Thus rab.c, while it will usually be <ra.b,
could be slightly more than .9. Similar situations can be shown to
exist with larger-sized groups. The increases found by us were well

Evaluating Systematic Relationships

1435

below the mathematically possible limits. In all such cases the re-
lations were represented as multifid furcations of all the stems in-
volved in the reversal and at the highest of the several Ln levels con-
sidered.

In a successful method of studying relationships, the results of
the analysis should be relatively independent of the number of
species in the correlation matrix. If at least one species per species-
group is included in a matrix, the ideal method of analysis should
reproduce the diagram of relationships based on an earlier study of
a larger matrix. If the method can be shown to produce similar
results, the fact that our matrix contains only a sample from the
population of species can be ignored with greater assurance.

We tested this question by subjecting the odd-numbered species
in the entire genus Ashrneadiella to a weighted variable group anal-
ysis. This should give an adequate cross-section of relationships in
that genus. Since some trends might be lost by exclusion of the

69 65 71 83 79 85 89 93 97
63 75 67 77 81 73 87 91 95

900-

800-

700-

600 -J

Fig. 5. Diagram of relationships predicted for odd numbered species of the
genus Ashmeadiella on the basis of the relationship diagram of figure 1. Ex-
planatory comments as for figure 1.

1436

The University Science Bulletin

even-numbered species a special diagram of relationships was pre-
pared from figure 1 by using only odd-numbered species. Figure 5
shows this predicted diagram of relationships.

Figure 6 shows the results of the weighted variable group analysis
on the odd-numbered species. There is less structure in this dia-

69 65 71 83 79 85 89 93 97

63 75 67 77 81 73 87 91 95
1000-1

900-

800-

700-

\

1

./

J

600-

Fig. 6. Diagram of relationships obtained by an independent weighted
variable group analysis of the odd-numbered species of the genus Ashmeadiella.
Explanatory comments as for figure 1.

gram as compared with the predicted diagram of Figure 5. This
was to be expected since some of the structure was based on rela-
tions involving the missing even-numbered species. In general,
agreement between the two diagrams is very good, however, and
we therefore feel reassured that the species left out in our study
would not have changed greatly our diagrams of relationships.

Evaluating Systematic Relationships 1437

CONCLUDING COMMENTS

A detailed discussion of the comparisons of our findings with the
previous classifications of the four genera of bees is given by
Michener and Sokal ( 1957). It will suffice here to state that general
agreement was good but that a number of taxonomic re-evaluations
seemed necessary as an outcome of these analyses. It should be re-
membered that while diagrams such as figure 1 may suggest phylo-
genies, in reality they only indicate static relationships. As indi-
cated in the paper referred to, additional refinements were devised
to give diagrams of relationships which we believe more nearly ap-
proach phylogenetic trees.

In view of these results we are encouraged to believe that, since
the methods we have described are increasingly practical with the
growing availability of high speed computers, this or similar schemes
will be more widely utilized with different groups of organisms.
Although the method we have described is a first attempt and would
profit by either simplification or refinement, we believe it is a step
toward reducing the subjectivity of systematic work, and therefore
a step in the right direction.

LITERATURE CITED
Burt, C. L.

1937. Correlations between persons. Brit. Journ. Psychol., vol. 28, pp.
59-96.
Cattell, R. B.

1944. A note on correlation clusters and cluster search methods. Psy-

chometrika, vol. 9, pp. 169-184.
1952. Factor analysis. Harper & Brothers, New York, pp. xiii + 462.

1954. Growing points in factor analysis. Austral. Journ. Psychol., vol. 6,
pp. 105-140.

Holzinger, K. U., and H. H. Harman.

1941. Factor analysis. Univ. of Chicago Press, Chicago, pp xii + 417.
Howells, W. W.

1951. Factors of human physique. Amer. Journ. Phys. Anthrop., n. s., vol.
9, pp. 159-191.
McQuitty, L. L.

1955. A method of pattern analysis for isolating topological and dimen-
sional constructs, Research Report AFPTRC-TN-55-62, Air Force
Personnel and Training Center, Lackland Air Force Base, San
Antonio, Texas, v + 38 pp.

1438 The University Science Bulletin

Michener, C. D., and R. R. Sokal.

1957. A quantitative approach to a problem in classification. Evolution,
vol. 11, pp. 130-162.

Olson, E. C, and R. L. Miller.

1951. A mathematical model applied to a study of the evolution of species.
Evolution, vol. 5, pp. 325-338.

Sokal, R. R.

1952. Variation in a local population of Pemphigus. Evolution, vol. 6,
pp. 296-315.

1958. Quantification of systematic relationships and of phylogenetic
trends. Proc. Tenth Internat. Congress Entomology, Montreal,
(in press).

Spearman, C.

1913. Correlations of sums or differences. Rritish Journ. Psychology,
vol. 26, pp. 344-361.
Stephenson, W.

1936. The inverted factor technique. Brit. Journ. Psychol., vol. 26, pp.
344-361.
Stroud, C. P.

1953. An application of factor analysis to the systematics of Kalotermes.
Systematic Zool., vol. 2, pp. 76-92.

Sturtevant, A. H.

1942. The classification of the genus Drosophila with descriptions of nine
new species. Univ. Texas Publ., no. 4213, pp. 1-51.
Thomson, G. H.

1951. The factorial analysis of human ability. 5th ed. Houghton Mifflin
Company, New York, pp. xv + 383.
N» Tryon, R. C.

1939. Cluster analysis. Edwards Bros., Ann Arbor, Mich., pp. 1-122.

THE UNIVERSITY OP KANSAS

SCIENCE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 23

The Role of Tissue Spaces and Nerve Fibers in the Spread
of Tetanus Toxin in the Rat

BY

Alexander A. Fedinec * and Howard A. Matzke *

Abstract. The extensive review of the literature on all phases of the
tetanus problem reveals there is general agreement as to the etiology, patho-
genecity, immunology and treatment of the disease. However, the mode of
transport of the toxin in the body and its site of action are not fully under-
stood. The following four pathways have been suggested by various workers:
vascular stream, axons, lymphatics of peripheral nerves and tissue spaces of
peripheral nerves.

Intraneural or intramuscular injection of toxin results in local tetanus.
This is characterized by a segmental progressive envolvement of musculature
beginning with muscle groups in the injected extremity. Blood-borne tetanus
follows intravascular inoculation and always results in envolvement of the
facial and masticatory muscles first despite the cite of injection. Blood-borne
tetanus appears following intramuscular injection if the epineurium is removed
and the perineurium teased open in the nerves to those muscles. However,
if antitoxin is also administered intravenously no symptoms of tetanus de-
velop. Intravascular antitoxin does not prevent the onset of local tetanus if
the nerve sheaths are intact. Local tetanus results from injection of toxin
intramuscularly in the absence of nerve fibers to those muscles. However,
the epineural sheath must be intact. The results of these experiments suggest
that the primary path to the central nervous system for tetanus toxin is by way
of the tissue spaces of peripheral nerves.

TABLE OF CONTENTS

PAGE

Introduction 1440

Acknowledgment 1468

Material and Methods 1468

Results 1470

Discussion 1473

Summary and Conclusions 1475

Literature Cited 1479

Tables 1476-1478

* Department of Anatomy, University of Kansas, Lawrence, Kansas.

(1439)

1440 The University Science Bulletin

INTRODUCTION

The current literature reveals that considerable clinical and ex-
perimental work is still being carried out on tetanus, however, the
knowledge of its etiology has changed little since the time of Hip-
pocrates ( Singer, '28; Chadwich, '50 ) . It is probable that its char-
acteristic symptoms were known even prior to that time. The treat-
ment of tetanus in the early part of this century was empirical, and
to a certain extent remains so today (Adams, 1856; Rose, 1897; Abel
and Hampil, '35; Nielsen, '51; Melbourne Pediatric Society, '53;
Chackleton, '54; Drew, '54; Bucco, '54; Hampton, '54). No agree-
ment has ever been reached on all aspects of the etiology of tetanus
(Brunner, 1894; Rose, 1897; Meyer and Ransom, '03; Abel and
Hampil, '35; Doerr et al '35a, b; Penitshka, '53; Drew, '54; Wright,
G. P. '54).

The extensive literature prior to the discovery of Clostridium
tetani by Nicolaier in 1884 was partly summarized by Hutchinson
and Jackson (1861), Brunner (1894), Rose (1897) and von Behring
('04). Curling's attitude is representative of that era. He stated
that tetanus is a disease of the nervous system (Curling, 1836).
Objections were raised against this theory (Heiberg, 1861). (For
further discussion see review article of Abel and Hampil, '35.)
Simpson recognized that the poison is produced at the site of the
wound (Humphreys, '35). Carle and Rattone in 1884 successfully
transferred the infectious agent, and the same year Nicolaier identi-
fied the bacillus in mixed cultures. Pure cultures were obtained by
Kitasato ( a and b ) in 1889 whereas Faber in 1890 induced tetanus
with germ-free filtrates and immunity was demonstrated by von
Behring and Kitasato in 1890. They also discovered the antitoxin
(Kitasato, 1891; Kitasato, 1892; von Behring, 1892), and thus closed
the period (1884-1891) in which most of the bacteriological and
immunological factors were ascertained.

The new century began in which success in prophylaxsis (Abel
and Hampil, '35; Boyd, '46; Long and Sartwell, '47; Vit. Stat, of the
U. S., '50; D'Antona, '52; Melbourne Pediatrics Society, '53; Ramon,
'53; Lancaster, '53; Slome, '54 and Major, '54 ) , and improvement in
management of the disease (Lanfranchi, 1295; Humphrey, 1856;
Meltzer and Aver, '05-'06; Buxton and Kurman, '45; Torrens, Ed-
wards and Wood, '48; McDonald, '49; Mclntyre, '49; Nielsen, '51;
Raynaud and Wright, '53; Melbourne Pediatrics Society, '53; Chesni,
'53; Benbanaste, '54; Chackleton, '54; Bucco, '54; Schubert, '54;
Drew, '54) was achieved, however, the manner of distribution of

Tissue Spaces and Nerve Fibers in the Rat 1441

the toxin in the infected organism, its mode of action and the cause
of localized and generalized symptoms in tetanus are still not fully
understood.

A. Bacteriology, Immunology and Toxicity
Clostridium tetani is described as slightly motile in vegetative
forms, gram positive, anaerobic and a slender bacillus which pro-
duces highly resistant spores. The dimensions are 0.3 to 0.5[x in
width and 2 to 5[a in length, but vegetative filaments of much greater
length occur. The spores are spherical, terminal, and wider than
the bacillus, which has a characteristic drumstick shape. The
bacilli are arranged singly or occasionally in chains. Isolated
colonies in deep dextrose agar have a wooly appearance and may
either be flocculent or with an opaque center. Surface colonies are
flat, rhizoid, or even feathery, and frequently colonies on blood agar
exhibit hemolysis (Burrows, '50). The growth requirements of
Clostridium tetani and factors in toxin formation were recently ex-
tensively studied by Fildes ('25-'29) and Mueller and his co-work-
ers (Fisek, Feeney, Miller, '42-'54; Ehrismann, '37 and others).
Maclennan ('39) on the basis of specific flagellar antigens dif-
ferentiated by agglutination ten types of Clostridium tetani. Toxin
produced by all forms of Clostridium tetani is identical and is
neutralized by one antitoxin. Antibacterial sera contain agglutinins
and opsonins specific to each type.

Von Behring and Kitasato (1890) demonstrated that filtrates of
the cultures contained toxin, which may be separated from the
bacteria by filtering through Berkefeld and Chamberland filters.
The toxin diffuses from within the cell in the surrounding medium
(Stone, '54) and is probably protein in nature (Eaton, '36). In
purified form the protein minimum molecular weight is about 67,000
(Pillemer, Wittier, Burrell and Grossberg, '48); however, component
structure and qualitative differences were postulated (Friedemann
and Hollander, '43a; Haku, '39; Aristovsky et al '38 and others).
Imbriano ( '50a ) describes the tetanus toxin as a protein with neutral
reaction. It is composed of simple alpha amino acids such as di-
carboxylic amino acids (glutamic and aspartic) and basic amino
acids such as hystidine. It probably contains another amino acid
such as serin or threonine. Tetanus toxin does not have in its struc-
ture homocyclic amino acids such as phenylalanine and tyrosine or
heterocyclic amino acids such as tryptophane. Ehrlich in 1898
separated the tetanus endotoxin into two compounds, the teta-
nolysin and tetanospasmin.

1442 The University Science Bulletin

No specific characteristic action of the endotoxin is known, how-
ever, lethal doses produce the usual symptoms of endotoxin intoxi-
cation (Burrows, '50). Concentrated solutions of toxin show
hemolytic activity which is due either to enzymatic action on plasma
lecithin, producing lysolecithins in the first stage, or to esterase
and protease action on the lipoprotein membrane of the erythro-
cytes. Tetanus toxin loses its hemolytic activity in vitro and its
antihemolytic activity in vivo when it is transformed into toxoid.
Antisera, penicillin G, antilipases (quinine salts) and antioxidants
( nordihydroquaiaretic acid) inhibits or retards hemolysis (Im-
briano, '50b). The tetanolysin causes hemolysis of red blood cor-
puscles. It induces antihemolysin production and seems more
thermolabile than tetanospasmin (Zinsser and Bayne-Jones, '39).
The diffusion constant of tetanolysin is 0.037 (Porter, '47). Some
nonpathogenic strains produce tetanolysin (Kerrin, '30; Hall, '30).
Boux and Borrel (1898) demonstrated that tetanospasmin has a
strong affinity for nervous tissue. Meyer and Bansom ( '03 ) showed
that the toxin once fixed by the nervous tissue cannot be neutralized.
Abel, Hampil, Jonas, and Chalian ('38) found that when three
lethal doses of the toxin were injected intravenously, the irreversible
fixation of one lethal dose was completed in the relatively short
time of 5 to 7 hours. Becently Wright, E. A. ('54) demonstrated
the rapidity of fixation of the toxin in the guinea pig's medulla ob-
longata. He states that at least one fatal dose becomes fixed in
less than 30 minutes following injection of 100 lethal doses. The
high toxicity is apparent from the fact that as little as 0.000013
gamma of the crystalline toxin prepared by Pillemer, Wittier and
Grossberg in 1946 was able to produce tetanus in mice after intra-
muscular injection. The purified toxin contains about 6,400,000
lethal mouse doses per milligram. Using cold methanol Pillemer,
Wittier, Burrell and Grossberg ('48) prepared a toxin that is lethal
to mice in doses of 0.09 X 10~3 pt,g.

The toxin is destroyed by proteolytic enzymes, and an aqueous
solution of tetanus is highly unstable to heat and light (Dubos, '52).
It is inactivated by ultraviolet light (Courmont and Nogier, '09a, b),
exposure to Beta-rays (Baker, '35), hydrostatic pressure (Basset,
Machebouef and Wollman, '37) and in vitro by vitamin C (Junge-
blut, '37). (For further references concerning the stability of the
toxin see Porter, '47, or original papers.) Imbriano ('50c), study-
ing the enzymatic properties of tetanus toxin observed no carbo-
hydrasic, phosphatasic, oxidasic and peroxidasic properties, how-

Tissue Spaces and Nerve Fibers in the Rat 1443

ever, he observed proteasic action on gelatine and an esterasic,
lipasic and lecithinasic activity. The toxin hydrolyzes the fatty
acid fraction of lecithin but does not act on the glycerophosphoric
part. The esterasic activity is zero at 0°C. and increases progres-
sively with an increase in temperature (up to 40°C). The inocu-
lation of an incubated lecithin-tetanus toxin mixture does not pro-
duce tetanus in rats. The lecithin is hydrolyzed by the tetanus
toxin effectively destroying its toxicity.

Requirements for toxin production were described by Mueller
and associates ('42-'54), Fildes ('25-'29), Knight and Fildes ('30)
and Campbell and Fildes ( '31 ) . They pointed out the dependence
of spore germination on the oxidation-reduction potential of the
tissues. The genesis of tetanus in the body is broadly discussed
by Topley and Wilson (Wilson and Miles, '46). They conclude
that the presence of suitable accessory factors enable the spores
to germinate and multiply in the tissues. It seems probable that
this action is dependent on the formation of an area of necrosis in
the tissue with sufficiently low oxidation-reduction potential to per-
mit the spores to germinate. Pelloja and Tusini ('50) found that
the course of experimental infection appeared to be aggravated
by the presence of destructive lesions, especially of the muscles.
Histamine has no influence on the progress of experimental tetanus,
therefore, the symtoms of tetanus are not considered to be due to
the liberation of histamine by the injured tissues.

Von Behring and Kitasato (1890) in a series of ingenuous ex-
periments demonstrated active immunity by injecting animals with
modified toxin treated with a solution of iodine trichloride. Passive
immunity was obtained by injecting sera of actively immunized
animals ( Kitasato, 1889 a, b; von Behring and Kitasato, 1890; Kita-
sato and Weyl, 1890; Kitasato, 1891; Kitasato, 1892; von Behring,
1892 and Kitasato, 1893). Topley and Wilson (Wilson and Miles,
'46 ) described the antitoxin as being fairly stable in cold and in the
absence of light. Exposure to sunlight or diffuse daylight gradu-
ally destroys it. It resists heat at 60° C. for half an hour but is
partly destroyed at 65°C and completely at 68°C. (Tizzoni and
Cattani, 1890). The antitoxin is nondyalisable and is precipitated
by ammonium sulphate. Hydrochloric and lactic acids and caustic
potash destroy it. The antitoxin is standardized. The American
unit of antitoxin is defined as 10 times the least amount of serum
necessary to save the life of a 350 gram guinea pig for 96 hours
against the standard test dose of toxin. The standard dose of toxin

1444 The University Science Bulletin

is 100 MLD of a standard toxin preserved at the National Institute
of Health, Washington. The permanent Committee on Standard-
ization of the League of Nations (Prausnitz, '29) has suggested an
International Standard Unit equivalent to one half an American
unit and based on standard antitoxin kept at the Serum Institute in
Copenhagen (Dubos, '52).

The therapeutic an immunological value of immune serum has
been extensively studied. The immunity thus obtained is of short
duration ( McConkey and Homer, 17 ) , and of doubtful therapeutic
value ( Korganova-Myuller, '40), however, good results have been
reported following intrathecal administration in animals (Sherring-
ton, 17) and man (Yodth, '32). Others pointed out the danger of
intrathecal administration (Wainwright, '26; Spaeth, '41a, b and
others). The intravenous route was preferred by other groups
(Florey and Fildes, '27) especially after Abel's blood transport
theory was introduced in 1934. Bruce ('20) points out the effec-
tiveness of tetanus antitoxin therapy and prophylaxis during the
years 1914-1918. The incidence in British war wounded in 1914-
1918 was 1.17 per 1,000 while mortality in pre serum days was
about 85%. According to Major ('54), the records in the Surgeon
General's Office in Washington show only four deaths from tetanus
from April 17, 1917 to December 31, 1919 in the American Army
where prophylaxis was introduced. During the American Civil War,
less than two in 100,000 developed the disease. Bruce ('20) found
that the incubation period was prolonged and there were fewer
fatalities among persons who received antitoxin prophylaxis treat-
ment. Dosage requirements were higher and success depended on
the time of administration. Abel and Chalian ('38) pointed out
that neutralization is confined to the nonfixed but already absorbed
toxin in the nervous tissue. Russe ('50) presents a survey of the
records from 10,250 patients who had tetanus antitoxin for prophy-
lactic purposes. The data showed that 3.9% of the cases had serum
reactions. No fatalities resulted from these reactions. D'Antona
and Valensin ('37) and many other investigators pointed out the
greater value of active immunization over passive. Consequently,
the value of antitoxin serum prophylaxis became the subject of a
controversy. It was suggested it be used immediately with wound
treatment (Milko, '40) and in combination with tetanus toxoid
(Firor,'39).

Pettavel ('40) reviews the history of active immunization since
1917 when Bazy inoculated soldiers with iodine treated tetanus

Tissue Spaces and Nerve Fibers in the Rat 1445

toxin. D'Antona ('52) reviewed the important literature on prophy-
laxis and immunity during the period 1923-1951. Since the for-
maldehyde treated toxoid was prepared by Descombey in 1924 and
aluminium precipitated toxoid by Bergey (1934) and Jones and
Moss (1936), active immunization found extensive use in prophy-
laxis ( Ramon, '53 ) . The use of toxoid in treatment was also studied
(Raynaud and Wright, '53) and protection of short duration was
found. The lethal effect of toxin can be prevented or delayed by
large doses of tetanus toxoid alone given 1 or 2 days previous to the
administration of the toxin. The explanation suggested is a block-
ing by the toxoid of a hypothetical substance in the C. N. S. which
would otherwise react with the toxin and produce nerve cell dam-
age. Another explanation is that there is a competitive inhibition of
the action of the toxin. The British regulations for standardization
require that more than 9 guinea pigs must be injected with five
times the human dose, and that after 6 weeks at least two thirds of
the guinea pigs must contain a minimum of 0.1 A. U. per ml. of
blood serum (Topley and Wilson, see Wilson and Miles, '46).
Since no international standard exists for tetanus toxoid, the United
States Army specified that the toxin contain at least 10,000 guinea
pig MLD's per ml. and be detoxified with 0.4 per cent formalin. The
final preparation must be atoxic for guinea pigs in 5 ml. amounts,
and guinea pigs receiving 1 ml. as an immunizing dose must be able
to withstand 10 MLD of toxin six week later. ( Long, '43, cited by
Burrows, '50.) Surjan, Richter and Rethy ('54a, b) purified and
concentrated tetanus toxoid with trichloracetic acid and obtained
toxoid which contained a grade of purity: 250-940 Lf/mg. N., and
200-900 Lf/mg. N. Delsal and Mir ('54) also obtained high con-
centration and purity by freezing at -15° C. The toxoid was pre-
cipitated with trichloracetic acid at pH4. Ipsen ('54) concludes
that no correlation exists between susceptibility to toxin and im-
munizability to toxoid. Multiple inoculations were recommended
for high titer and long lasting antisera ( Kestermann, Schleining and
Voght, '39). Different methods were adopted in the British army
(Boyd, '38; Boyd and Maclennan, '42). Wolters and Fischoeder
('53) investigated the value of an aluminium-absorbed prophylactic
and found that after two injections at 4-week intervals a basal im-
munity lasting at least 10 years was produced. A later inoculation
of a booster dose resulted in the production of circulating antitoxins
even within the shortest incubation period of tetanus. Similar con-
clusions were previously drawn by Bigler ('51) and others. Com-

1446 The University Science Bulletin

bined inoculations of tetanus toxoid with pertussis and diphtheria
prophylactic (Faerber, '53), or with typhoid-paratyphoid vaccine
(Maclean and Holt, '40), are common. Inoculations at birth with
diphtheria, pertussis and tetanus was described by Chamberlain
and Bullock ('50). War records or reports from territories where
active immunization is obligatory, reveal that by proper immuniza-
tion tetanus can be completely prevented. For instance, among
16,000 wounded in the British Expeditionary Force in France no
cases of tetanus occurred (Bensted, '40). Long and Sartwell ('47)
state that in the period of 1942-45 only 12 cases of tetanus occurred
in the United States Army of which 6 were nonimmunized persons.
There were 4 cases in the United States Navy of which 3 were non-
immunized persons. Similar excellent results were obtained in the
civilian population where the inoculation with tetanus and diph-
theria prophylactic were made compulsory, e. g. France ( Bull. Of-
fice Internat, d'Hyg. Publique, 1940, 32:748). However, due to the
fact that preventive inoculation is not in common practice the inci-
dence of tetanus is still very great. The average incidence of tetanus
in the U. S. A. from 1947-1952 was 546 cases per year (Hampton,
'54) and in South Africa during the period of 1945-1950, 1019 fatal
cases were recorded. The annual death rate per million persons
(1946-48) was 16.1 (Slome, '54). A total of 237 deaths were re-
corded in Melbourne during twenty years representing a mortality
of 50% of clinical cases. There was no case which could be shown
to have occurred after a prophylactic injection of antitetanus serum
(Melbourne Pediatric Society, '53). It is evident from these few
records that the high mortality and frequent occurrence of tetanus
in cases without any prophylactic treatment form a great medical
problem which can be solved by obligatory prophylaxis and im-
proved treatment.

The distribution of tetanus bacillus is broad. It has been iso-
lated from superficial layers of cultivated earth (Nicolaier, 1884),
from animal (Fildes, '25) and human feces (Tulloch, '19 ). Also,
there are frequent reports that tetanus may be a secondary infec-
tion caused by materials in an unsterile condition, e. g. plaster
bandages (Murray and Danton, '49), catgut (Savolainen, '50), fol-
lowing tonsillectomy (Bajog, '53) and otitis media (Dave, '53).
Tetanus neonatorum, following umbilical infection, may occur.

It was revealed by the experiments of Sherrington (T7) and
others that there is present a great variability in susceptibility to
tetanus toxin in man and in animals. The most susceptible mam-

Tissue Spaces and Nerve Fibers in the Rat 1447

mals are the horse, guinea pig, monkey and mouse and the most
resistant the cat and dog (Fildes, '29a). Simic and Sahovic ('50)
found that eancerogenic tissue can also bind the tetanus toxin.
The sensitive cells have at least two binding groups. Aristovsky,
Kambrova and Firsova ('38) found that dogs are more sensitive
than mice to certain types of tetanus toxin. Haku ('39) performed
experiments on cold-blooded animals which normally are refractive
to tetanus toxin. He concludes that under higher temperatures a
third active component is present in the toxin ( in addition to tetan-
ospasmin and tetanolysin ) which acts upon the ganglion cells of the
heart.

Usually an incubation period intervenes between the inocula-
tion or entry of the infectious agent under the skin and appearance
of symptoms. The incubation period cannot be reduced to less than
eight hours with the usual toxic filtrate. Beyond that point the in-
cubation period is inversely related to the amount of toxin injected
(Burrows, '50). The usual incubation period in humans varies
from 3-21 days (Bruce, '20). Pillemer and Wartman ('47) pro-
duced tetanus symptoms in 30 minutes and death in an hour by in-
jecting 500,000 MLD of crystalline toxin intramuscularly. Pelloja
('50) concludes that the incubation period represents the time
required for the toxin to penetrate the neurons and the moment
of death does not depend solely on the action of the toxin but also
on factors such as the somatic constitution of the animal and the
decomposition of the toxin. Meyer and Ransom ('03) relate the
incubation period to the distance the toxin has to travel along the
nerves to reach the central nervous system. (For further dis-
cussion on the incubation period see Imbriano, '50d and D'Antona,
'51.)

B. Histopathology

Although pathologic changes occur in tetanus, it is generally
agreed that there is no set pattern of changes characteristic of the
disease. Most of the pathologic alterations reported in humans
are probably postmortem changes. Hassin ('48) emphasizes the
importance of differentiating the changes produced experimentally
in animals from those occurring in humans. Lesions in the ventral
horn cells were demonstrated by Nissl ( 1896 ) . In a review Barker
('01) pointed out the close resemblance between the changes pro-
duced by strychnine and those due to tetanus poisoning. Gold-
scheider and Flatau (1898) and Chantemesse and Marinesco
(1898) observed in their experimental animals gradual alterations

14—8050

1448 The University Science Bulletin

in the nucleolus and tigroid substance resulting in deformity of
nucleolus and complete chromatolysis from 2 to 24 hours after in-
jection of the toxin. Claude (1897) and Hassin (48) reported
demyelination associated with round cell infiltration, proliferation
of glia and preservation of axons. Recently Zaffiro ('50) observed
in guinea pigs that all the organs appeared grossly to be markedly
congested. There were small hemorrhages in the lungs, liver and
kidneys with areas of necrosis in the liver. Microscopically, the
brain showed vascular congestion, edema, proliferation of astrocytes
and neuronophagia. In the spinal cord there was congestion, de-
generation of motor neurons and neuronophagia. The alterations
in humans are similar to those found in experimental animals.

Curling (1936) stated that, "There are, therefore, no morbid
changes peculiar to tetanus, and by which it can be recognized."
Alterations in the anterior horn cells, medulla, and cortex were re-
ported by Halban ('00) and others. Baker ('42, '49) pointed out
that in man the brain is negative or reveals only scattered areas of
subarachnoid bleeding and intracerebral petechiae. Prior to the
third day no microscopic lesions were present. From the third to
the seventh day of illness neuronal changes appear involving scat-
tered cortical areas as well as specific cranial nerve nuclei. Neurons
are swollen and show a characteristic perinuclear chromatolysis.
From the seventh to the tenth day there appear focal areas of peri-
vascular demyelinization. If the patient survives beyond the tenth
day the glia proliferates to form scattered foci of small and large
glial nodules. All histologic lesions seem to have a vascular distri-
bution suggesting a hematogenous spread of the toxin. Kassay and
Mathe ('51) detected changes in the spinal cord, the nuclei of the
extrapyramidal tracts, the mesencephalic nucleus of the trigeminal
nerve, the hypothalamus and the pituitary gland. They consider
that these changes are a result of reflex irritation due to an excess
of acetylcholine, and that they represent a hyperfunction leading to
the death of the neurons concerned. Wechsler ('52) pointed out
that there are no consistent pathologic changes and Courville ('53)
states that there are no characteristic sites of inflammation. With
the exception of a general or moderate congestion in organs other
than the central nervous system, no pathological changes are ob-
served (Zinsser and Bayne-Jones, 1939).

Ranson and associates ('26-'29) made histological observations
of muscles which were altered to irreversible gel due to the action
of tetanus. Ranson and Sams ('28) studied the action of tetanus

Tissue Spaces and Nerve Fibers in the Rat 1449

upon muscles with regard to myostatic contracture of Moll ( 1886 ) .
The hypertonus in local tetanus is reflex in character and later the
myostatic contracture is superimposed upon it. (Similar conclu-
sions were reached by Klensch and Schlomer ('52) and others.)
After prolonged involvement of the muscles mild changes such as
an increase in the number of sarcolemmal nuclei and obscurity of
the cross-striations were observed. According to the authors the
rigidity was related to a change in contractility of the elastic ele-
ments in the muscle fibers resulting in shortening the muscles' rest-
ing length by 10 to 40 percent. Adams, Denny-Brown and Pearson
('53) were not convinced that the changes are necessarily in the
muscle fibers, and believe that a similar phenomenon is common
to all true elastic tissue. They feel the uneven intensity of staining
and the obscure and irregular striations reported by Davenport,
Ranson and Stevens ('29) fall within the limits of variability in-
herent in the process of fixation. Lesions similar to Zenker's hyaline
degeneration have been less often observed and muscle rupture may
occur from violent contraction in severe tetanus.

Tetanolysin causes destruction of red blood corpuscles. The en-
dotoxin does not have a specific action or result in a characteristic
alteration which differs from the general picture of endotoxin in-
toxication following administration of a lethal dose. The usual
alterations are damaged blood vessels resulting in degenerative
changes in the tissues supplied (Burrows, '50).

C. The Spread of Tetanus Toxin

Gumprecht (1893, 1895a, b) states that the first authors who
turned their attention to the problem of the spread of tetanus toxin
were von Brunner ( 1891-94 ) in German literature and Autokratow
(1892) and Courmont and Doyon (1892-93-99) in French litera-
ture. Brunner's experiments in 1894 were a confirmation and broad-
ening of results observed by Autokratow (1892), Courmont and
Doyon (1892) and Buschke and Oergel (1893), however, no sound
conclusions were drawn. In accordance with the attitude prevail-
ing at that time, their experiments were interpreted as providing
experimental evidence that tetanus toxin was carried from the
periphery to susceptible centers in the central nervous system by
some component of peripheral nerve trunks.

It was not until Marie (1897), Marie and Morax ('02-'03), Meyer
('01), Meyer and Ransom ('03), Loewi and Meyer ('08), Frohlich
and Meyer (16) and others performed a series of classical experi-
ments that the first concrete theory was formulated. This theory

1450 The University Science Bulletin

later became modified but essentially remained unaltered. Accord-
ing to the theory of Meyer and Ransom ('03) the following events
take place in tetanus. From the site of inoculation or natural in-
fection the toxin spreads through the tissue spaces. Some of it may
be absorbed directly by the naked terminals of motor nerves, how-
ever, Meyer and Ransom believe most of it is absorbed by the
lymphatic capillaries and circulated throughout the body by the
vascular stream. In this manner the toxin reaches all motor nerve
terminals through which it is absorbed. Then it is probably carried
in axis cylinders of the motor nerves to the spinal cord where it is
fixed by the motor neurons of the anterior homs. After a certain
amount of toxin has entered these ganglion cells there is induced a
state of increased irritability in these cells and this in turn causes
contraction of muscles innervated by these respective cells (homo-
lateral contraction). Excess toxin courses along the fibers of the
anterior commissure to the neurons of the other side of the spinal
cord producing tetanic contraction of the opposite extremity. After
enough toxin has accumulated in the spinal cord via carriage of
the toxin from the periphery in the motor nerve trunks, the toxin
is fixed by the tactile apparatus of the reflex arc resulting in an in-
crease in the reflexes of the involved extremity. Through further
spread of the nonfixed toxin in the direction of the brain new motor
centers and reflex arcs are affected.

Recently D'Antona ('49-'51) summed up the fundamental con-
cepts of the above described theory. Among others D'Antona sug-
gests that the motor nerves absorb the tetanus toxin at the motor
end plates which are covered only by the delicate sheath of Henle
( neurilemma ) . The myelin sheath, the membrane of Schwann and
the endoneural membrane of RufBni are absent at this point. The
sensory nerves do not absorb the toxin since they frequently termi-
nate in corpuscles (Golgi, Meissner, Pacinian etc.) which are
formed by structures impermeable to tetanus toxin or have a bio-
chemical constitution unadapted to fix the toxin. However, Zironi
('51) points to the fact that many motor endings are covered with
a neurilemma as they penetrate the sarcoplasm in muscle fibers, and
also there are a large number of free sensory endings that do not
possess myelin sheaths.

Meyer and Ransom ('03) produced tetanus dolorosus by injecting
the tetanus toxin into the spinal sensory areas. This never occurred
when the toxin was injected peripheral to the dorsal root ganglion.
To many workers these results suggested that tetanus toxin does

Tissue Spaces and Nerve Fibers in the Rat 1451

not pass beyond the dorsal root ganglion (Fleteher, '03 and Teale
and Embleton, '19-'20). Firor and Jonas ('38) produced reflex
tetanus by intraspinal injections without producing tetanus clo-
loros.us or muscular rigidity in the periphery. Zironi ('51) points
out that pain analogous to that occurring in tetanus dolorosiis re-
sults from sectioning sacral nerve roots independent of the introduc-
tion of toxin. Meyer and Ransom ('03) suggested that tetanus toxin
ascends along sensory fibers but much slower than in the motor
nerves. Wright, G. P. (53) explains the slow movement along
sensory nerves as due to the lack of a sufficient peripheral pressure
gradient in subcutaneous tissue fluid to propel the toxin along
sensory-cutaneous nerves to the central nervous system. On the
other hand the pressure gradient in muscles is considerably higher
as pointed out in Panomarew's experiments in 1928. This agrees
with the observations of Meyer and Ransom ('03), Zupnik ('05),
Sawamura ('09), D'Antona ('49-'52) and others who observed that
descending or general tetanus, but no local symptoms, resulted from
injections in sensory nerves, skin, testicles, etc.

In the years following Meyer and Ransom's ('03) experiments
numerous investigators presented evidence supporting their neural
theory: Permin (14), Friedemann et al ('39-'49), Buzello ('29),
Doerr et al ('35, '36), Bromeis ('38), Roofe ('47, '48), D'Antona
('49-'52), ZafBro and Edoardo ('50) and others. The theory was
also accepted by numerous clinicians (Jelliffe and White, '17;
Wechsler, '52 and others ) .

Although the above authors presented experimental evidence for
a theory which postulates a centripetal flow of axoplasm or some
other means by .which substances are carried centrally along the
axons, not all results of work related to the structure of the axon
support such a theory. For further discussion on the structural
aspects of the problem see review articles of Parker ( '29 ) , De Renyi
('32) and the more recent electron microscopic observations of De
Robertis ('54), Meyer ('55) and Young ('56).

The Marie-Meyer central neural theory became a subject of
controversy shortly after its formulation. According to some authors
it failed to account in a satisfactory manner for many of the features
of the disease (Abel and Hampil, '35). New theories and modifi-
cations were proposed. The variations of Marie-Meyer theory are,
( a ) the carriage of toxin in the lymphatic vessels of the peripheral
nerves to the cerebrospinal fluid, thus affecting the centers in the
central nervous system, and (b) the tissue space theory which

1452 The University Science Bulletin

proposes a centripetal movement of the toxin in perineural spaces
of peripheral nerve trunks toward the spinal cord and a continuous
upward movement in the cord to the vital centers in the medulla
oblongata.

The lymphatic spread of tetanus toxin in the peripheral nerve
trunks towards the spinal cord and cerebrospinal fluid was first
suggested by Gumprecht (1895 a, b). Field ('06-'07) was of the
opinion that tetanus toxin does not travel up the nerve by reason
of specific attraction to nervous tissues, but because the lymphatic
flow in the nerve is from the periphery towards the center. Teale
and Embleton (20) strongly supported this theory although they
did not exclude the possibility of axonal carriage. Golla ('17) also
believed in some degree in the perineural lymphatic spread. Partial
involvement was proposed by many others (Stinzing, 1898, and
Robertson, '16). Wischnewsky ('28) traced methylene blue cen-
trally in nerve trunks and Horster and Whitman ( '31 ) detected dye
in the cerebrospinal fluid following injection of trypan blue and
tetanus toxin intraneurally. These results were interpreted as
further support for the theory that tetanus toxin is carried to the
central nervous system via the lymphatics of peripheral nerves.

The tissue space theory was first formulated by Panomarew ('28)
and his associates. These authors demonstrated a direct connection
between the perineural tissue spaces and subarachnoid spaces.
They noticed that the rate of centripetal movement of the toxin
along the nerve trunk is dependent on the subarachnoid pressure
gradient in cerebrospinal fluid and on the pressure in muscles in
the periphery. Abel, Evans, Hampil and Lee ('35) pointed out
that the tissue pressure of muscles is not of sufficient magnitude
to move solutions through narrow tissue spaces and to overcome
the pressure in the cerebrospinal fluid. Abel's ('34) measurements
of such a pressure were recently criticized by Wright, G. P. ('53).
Wright along with Drinker and Field ('33) and Barcroft and Dorn-
horst ( '48 ) point out that during the mildest exercise, such as walk-
ing, an adequate pressure gradient is produced several times a
minute. Wright, E. A., Wright, G. P., Baylis, Ambache, Davies
and their associates ('48-'54) attempted to disprove the axis cylinder
as a means of transportation of tetanus toxin. In a series of experi-
ments they present strong evidence demonstrating centripetal move-
ment of toxin in perineural tissue spaces of peripheral nerve trunks
and the spinal cord.

Indirect support was given to the endoneural and perineural lym-

Tissue Spaces and Nerve Fibers in the Rat 1453

phatic spread or tissue space spread theories by the anatomical find-
ings of Key and Retzius (1875), Funaoka and Yamada ('29), Clark
('29), Mantell ('32), Sullivan and Mortensen ('34), French, Strain
and Jones ('48), Brierley and Field ('49), Moore ('53), Moore,
Hain, Ward and Bridenbaugh ('54) and others. These authors used
intraneural or intrathecal injections of dye, radiopaque materials,
radioactive phosphorus, drugs and other substances, and detected
its appearance in cerebrospinal fluid or along the peripheral nerve
trunks.

Others argue that the many substances and techniques used for
the above studies actually dissect a connection between the cerebro-
spinal fluid and the peripheral nerve. Following an extensive re-
view of the literature Elman ('23) concluded a communication did
not exist.

Following injection of tinted radiopaque media into a peripheral
nerve, French, Strain and Jones ('48) reported free flow of the
medium within the fasciculi, however, lateral diffusion was pre-
vented by the endoneurial connective tissue planes. In the distal
portion of the nerve the medium was confined to the periaxonal
connective tissue. Adjacent to the dura it assumed a position in
the perineurial interstices and finally in the perineurium itself.
From this position the medium entered the subdural space and
finally broke through into the subarachnoid space at or near the
arachnoid cuff.

Moore ('53) followed the course of colored efocaine in the periph-
eral nerve. The material remained in the epineurium and entered
the parenchyma of the spinal cord directly. The dorsal root gan-
glion acted as a temporary block to the central flow of the medium.
After passing up and down in the spinal cord, some material dif-
fused through the pia to enter the subarachnoid space. Elman
('23) reported that although intravital injections of India ink in-
troduced into cerebrospinal fluid escaped from the cord into lym-
phatic channels of the region, he could not demonstrate the existence
of a fluid pathway physiologically connecting subarachnoid cavity
with perineural spaces. Weed ( '14-15 ) , Iwanow and Romodanow-
sky ('27), Zhdanow ('31), Kurdiimow ('32) and Defrise ('30)
pointed out that there is no direct connection between perineural
and interfascicular lymph vessels and the central nervous system
and cerebrospinal fluid, but that lymph from peripheral nerve trunks
reaches lymph nodes adjacent to the vertebral column and eventu-
ally the principal lymphatic vessels. Similar conclusions were drawn

1454 The University Science Bulletin

by Abel, Evans, Hampil and Lee ('35), Abel, Hampil and Jonas
( '35 ) and others.

Although it is commonly accepted that epineural lymphatics do
not communicate with the cerebrospinal fluid it is interesting to
note the experiments of Brierley and Field ('48) and Field and
Brierley ('48). These authors found that finely divided India ink
particles injected into the subarachnoid space may be traced along
emerging nerve roots to the prevertebral lymphatics. The flow is
dependent on the pressure gradient between the subarachnoid
space and abdominal lymphatics. The flow can be reversed by in-
creasing pressure in the abdominal lymphatics to 15 mm Hg.

Causey ('48) applied pressure to nerve trunks and subsequently
measured the fibers and nerve trunk diameter. He concluded that
perineural tissue spaces filled with tissue fluid actually exist. Birren
and Wall ('56) studied the proportion of nerve fibers, connective
tissue spaces, and blood vessels in the sciatic nerve of the rat,
preparing the tissue by the freezing-drying method. They found in
animals of 50-250 days of age the median volume of the nerve oc-
cupied by nerve fibers was 60.9 percent, the connective tissue spaces
amounted to 36.8 percent and blood vessels 1.2 percent.

The following arguments have been presented by various authors
in support of the general concept that tetanus toxin reaches the
central nervous system through peripheral nerve trunks:

1. To the present time numerous investigators (Baylis, et al, '52;
Wright, et al, '51 ) have demonstrated the presence of toxin in nerves
nearest the point of inoculation. Meyer and Ransom ('03) de-
tected toxin in the nerve one and one-half hours after subcutaneous
injection. Furthermore, Riggs and Matzke ( unpublished ) by utiliz-
ing a fluorescent antibody technique showed that the toxin reaches
the sciatic nerve within six hours following inoculation into the
gastrocnemius muscle of the rat.

2. If antitoxin is injected into the nerve trunk or spinal cord
segment proximal to the point of inoculation of the toxin, tetanus
fails to develop in that extremity (Meyer and Ransom '03; Permin,
14; Teale and Embleton, 19-'20; Roofe, '47 and others ) . Antitoxin
injected into the sciatic nerve of the left leg delays or prevents the
onset of tetanus in the right leg into which toxin was inoculated
(Wright, et al, '51).

3. Various blocking agents have been introduced into the sciatic
nerve in an attempt to prevent the spread of toxin. Teale and
Embleton (19-'20) using tincture of iodine, egg albumin and horse

Tissue Spaces and Nerve Fibers in the Rat 1455

serum successfully delayed the onset of symptoms. Wright et al
('51) and Baylis, Mackintosh, Morgan and Wright, G. P. ('52) in-
jected ethanolamine oleate and quinine urethane into the sciatic
nerve. These materials had the effect of stimulating the deposition
of connective tissue without interfering with the function of the
axons. The severity of the symptoms resulting from injection of
toxin distal to the sclerosed area was greatly reduced.

4. Following intraneural injections a wide variety of materials
have been traced along peripheral nerves to the central nervous
system by a number of techniques. Recently Brierley and Field
('49) with India ink and Wright et al ('51) with radioactive mate-
rials claim that some element of the peripheral nerve constituted
the pathway to the central nervous system for these various mate-
rials. Perdran ('37) suggested the axis-cylinder as a pathway for
dyes and salts in solution. Marinesco and Draganesco ('23) felt
that the tissue spaces of peripheral nerves constituted the pathway
for herpes virus, whereas Howe and Bodian ('42) felt that the axis-
cylinders constituted the route to the central nervous system for
the virus of poliomyelitis. All of these experiments indicate some
element of the peripheral nerve is involved, however, there is no
agreement as to whether this may be the axis-cylinder, the tissue
spaces or lymphatics.

5. Results of experiments where the peripheral nerves or spinal
cord were transected also indicate that the peripheral nerve serves
as a pathway for the toxin to the central nervous system. Meyer
and Ransom ('03), Permin (T4), Firor, Lamont and Shumacker
('40), Friedemann et al ('41), Baylis, Joseph, Mackintosh, Morgan
and Wright, G. P. ('52) and others were able to avert fatal intoxica-
tion if the spinal cord was sectioned at a site between the point of
inoculation and the brain stem. By sectioning the peripheral nerve
von Brunner (1894), Courmont and Doyon (1899), Tizzoni and
Cattani (1890), Ranson ('28) and others, were able to prevent the
development of tetanus in the muscle innervated by that nerve.
By freezing a segment of the sciatic nerve and allowing 16 days
for recovery of the blood supply and connective tissue Roofe ('47,
'48) was able to prevent the development of local tetanus. Zaffiro
and Edoardo ('50) prevented the development of local tetanus
and markedly delayed the onset of symptoms of general tetanus
by longitudinally sectioning the peripheral nerve trunk.

6. Meyer and Ransom ( '03 ) injected toxin directly into the spinal
cord. The first musculature involved was that innervated by nerves

1456 The University Science Bulletin

arising from the injected segment. Sawamura ('09) observed
tetanic involvement of the muscles in the leg opposite to that which
was injected with toxin. Wright, E. A. et al ('51) was able to
trace radioactive materials and tetanus toxin into the nerve of the
leg opposite to the injected extremity and also up the spinal cord
to the brain stem. In other experiments these authors ('50) were
able to bring about salivation, strabismus, torticollis and brady-
cardia following injection of the toxin into cranial nerves VII,
X, XII. Although the peripheral structures involved are widely
separated anatomically, they all receive their motor innervation from
nuclei which are in proximity, one to another, in the brain stem.
Furthermore, electromyograms indicate that the muscles which are
first and most severely involved are those innervated by the spinal
cord segment innervating the inoculated area and not necessarily
those muscles which are adjacent to this area (Davies et al '54).
Rocchi ( '40 ) observed a markedly increased electrical sensitivity in
muscles injected with toxin and showing tetanic symptoms as com-
pared with the rest of the musculature in the body.

7. Meyer and Ransom ('03) suggested a hematoencephalic
barrier existed for tetanus toxin. This theory has been supported
by the extensive work of Friedemann and associates (Zuger, Hol-
lander and Tarlow, '39-'41). These authors were able to demon-
strate both local and general tetanus following intramuscular inocu-
lations even though all circulating toxin was neutralized by an
antitoxin. Local tetanus was produced in actively and passively im-
munized animals following intramuscular injection of toxin ( Meyer
and Ransom, '03 and Permin, '14).

8. Firor, Lamont and Shumacker ('40) and Wright, E. A. ('53)
demonstrated that the lethal dose of toxin decreases progressively
and steeply when inoculations are made into the blood stream,
limb musculature, nerve trunks, spinal cord, and medulla oblongata
respectively.

9. Bromeis ('38) correlated the toxin content of a nerve at dif-
ferent levels with the time of inoculation. Permin ('14) injected
toxin intramuscularly in the hind leg, antitoxin in the blood and cut
the nerve after the injection at different times. Tetanus was not
prevented if the nerve was cut five hours after injection. From
that he concluded that tetanus toxin spreads along nerve fibers
rather rapidly. On the basis of time of toxin application and time
of onset of tetanus symptoms, Roofe ('47) determined that the
toxin travels along the sciatic nerve at the rate of 3.35 mm/hour.

Tissue Spaces and Nerve Fibers in the Rat 1457

Descombey and Robin ('30) pointed out the importance of the
site of inoculation to the time of development of the symptoms.
The closer the site of inoculation is to the central nervous system,
the shorter the distance the toxin needs to travel and thus, the
shorter the incubation period.

10. Friedemann, Hollander and Tarlov (41) and D'Antona
('49a, b, '51) point out that it takes 20 to 80 times more antitoxin
to neutralize intramuscularly injected toxin than to neutralize the
same amount injected intravenously. If, following intramuscular
injection of toxin, the nerves supplying that extremity are sectioned
the amount of antitoxin necessary to neutralize this toxin is the
same as that required to neutralize the same amount of intrave-
nously injected toxin (Friedemann, Zuger and Hollander, '39b and
D'Antona, '49a, b, '51). Friedemann and Traub ('49) pointed
out that the quantity of antitoxin needed to save an animal inocu-
lated subcutaneously with tetanus toxin is considerably reduced if
the antitoxin is introduced into the spinal cord rather than into a
vein.

11. The use of parabiotic animals in studying the spread of
tetanus toxin was first suggested by von Sauerbruch and Heyde in
1908. Ranzi and Ehrlich ('09) injected toxin into one member
of a pair 5 days after they were placed in parabiosis and noted
that the uninjected parabiont failed to develop any symptoms of
tetanus. Rocchi ('40) utilized the technique of crossed sciatic
nerve regeneration in parabiotic rats. In one pair with 20 days re-
generation time toxin was injected into the extremity into which
the sciatic nerve was regenerating from the opposite animal. Tet-
anus developed in the opposite animal presumably by coursing
along the crossed regenerated nerve. Eight hours after inocula-
tion of this animal toxin could be detected in the blood and certain
organs of the uninjected animal. Richter and Schreckenbach ('54)
produced general tetanus in both parabionts united for 126 days by
injecting toxin intravenously in only one. Utilizing a larger series
of parabiotic animals, some with and others without nerve cross.
Matzke and Fedinec ('55) concluded that toxin will travel rapidly
along nerve trunks, however, it may also be disseminated by the
blood. These findings that toxin may spread by both the peripheral
nerves and vascular stream support those of von Brunner, 1894 and
Albert, 18.

A number of investigators deny the importance of the peripheral
nerve in the transport of tetanus toxin to the central nervous system

1458 The University Science Bulletin

(Zupnik, '0Q-'08; Albert, '18). In a series of nine papers between
1934 and 1938 Abel and his associates put forth a number of argu-
ments and experimental results supporting the theory that the
vascular stream is the main route for the toxin. This theory is sup-
ported by a number of other workers particularly Zironi ( '51 ) and
Penitschka ('53). The principal arguments cited to support this
theory are as follows:

1. Zironi ( '51 ) points out that if the toxin travels by way of the
nerve it should then be possible to detect it within the nerve, how-
ever, if a nerve from an animal infected with the toxin is homo-
genized and the homogenate injected into another animal, the
injected animal fails to develop tetanus. He therefore concluded
that the toxin was not present in the nerve.

It has been demonstrated numerous times that tetanus toxin
has a strong affinity for nervous tissue. Despite this fact homo-
genates of brains of animals infected with toxin did not produce
tetanus when injected into other animals (Wassermann and Takaki,
1898). Imbriano ('50c) demonstrated a strong lecithinase activity
of the toxin. Riggs and Matzke (unpublished) were able to demon-
strate the presence of the toxin in the myelin sheaths of peripheral
nerves utilizing a fluorescent antibody technique. This latter may
explain the findings of Pochhammer ( '09 ) who suggested that toxin
causes local alterations in the myelin sheaths. All of these experi-
ments would indicate that toxin has a strong affinity for nervous
tissue and is probably bound by this tissue. In this state it is un-
able to infect other animals.

2. It is generally agreed by most supporters of the nerve trans-
port theory that the toxin travels more rapidly along motor nerves
than sensory. Zironi ( '51 ) argues that the termination of motor
fibers are well protected by Henle's membrane and the sarcolemma.
This constitutes a barrier as great as that around the free sensory
endings, therefore, the toxin should penetrate both equally well.
Wright, G. P. ( '53 ) argues that the more rapid movement of toxin
along motor fibers is not dependent entirely upon the protection at
the motor end plates but upon a pressure gradient between the
periphery, namely the muscles, and the central nervous system.
This pressure gradient is not as great along sensory nerves.

3. Vaillard and Vincent (1891) and Zupnik (1900-'08) showed
that local tetanus may be entirely avoided if the toxin is injected
into the subcutaneous tissues of the tip of the toe, the end of the tail,
the dorsum of the foot or in the neighborhood of certain joints that

Tissue Spaces and Nerve Fibers in the Rat 1459

are not in contact with the body of a muscle. These authors felt
that this supports the blood dissemination theory and local action
on muscles directly. This opinion differs markedly from conclusions
drawn from similar experiments by Meyer and Ransom ('03),
D'Antona ('49-'52), Wright, E. A. et al ('51) and Wright, G. P.
('53).

4. As cited above, if a peripheral nerve is sectioned and the
toxin injected distal to this point, the animal fails to develop tetanus.
Zironi ('51) feels that this is due to the trophic effect of the sec-
tioned nerves on the peripheral tissues altering the permeability of
the vessels and the diameter of the lumen. The toxin, therefore,
does not penetrate the vessels as readily as in an animal with an
intact nerve. Ranson ('28) observed the development of tetanus
following section of "the sensory fibers only, indicating that the
trophic effects had no influence on the spread of the toxin.

5. Lenormant, Lanari and Devoto, and Pelloja ('51), (quoted by
Zironi, '51 ) sectioned the motor nerve to a muscle in tetanus and
noted that the local tetanus did not disappear. Ranson and Sams
('28) believe that local tetanus is first reflex in character and later
myostatic contracture is superimposed upon it. Klensch and
Schlomer ('52) essentially agree with this conclusion stating there
are two phases in local tetanus: a.) the phase of typical tetanus
when the muscle contraction is due to an increased discharge of im-
pulses reaching the muscle; and b. ) a phase of prolonged con-
tracture due to changes within the muscle itself. This latter phase
is not altered by sectioning of the motor nerve to the muscle.

6. Harvey ('39) has shown that if tetanus toxin is injected intra-
neurally soon after the nerve has been cut, local tetanus appears.
He further states that local tetanus cannot be produced after the
motor end plates have degenerated. No other workers have been
able to repeat his experiments.

7. Abel et al ('35) failed to prevent the development of local
tetanus by injecting antitoxin into the nerve proximal to the site of
inoculation of toxin. Furthermore, they were unable to produce
local tetanus by intraneural injections of tetanus toxin. This work
is supported by that of Penitschka ('53). He injected Mo MLD of
tetanus toxin intraneurally in the hind leg of a dog and Hoo MLD
intramuscularly into the front leg. Local tetanus appeared in the
front leg several weeks before its appearance in the hind leg. This
was despite the fact that a larger dosage was injected intraneurally
in the hind leg.

1460 The University Science Bulletin

No workers have been able to repeat the experiments of Abel
and his associates. Most of them have been able to block the course
of toxin in the nerve with intrasciatic injection of antitoxin. D'An-
tona ('51) feels that toxin is absorbed at the motor end plates and
passes to the central nervous system more rapidly than when it is
injected directly into the nerve trunk. This suggests an explanation
of Penitschka's results, namely that the intramuscular injection was
more effective.

8. Zironi ('51) believes that antitoxin injected into the vascular
stream is less effective because it is diluted and thus there is less
chance for it to combine with the specific antigen. Injections into
the cerebrospinal fluid are thus more effective since the toxin upon
passing through the blood-brain barrier is immediately neutralized
by the antitoxin. Many experiments were performed to ascertain
which method of antitoxin administration is the most effective.
The intravenous and intraspinal are commonly believed to be the
most desirable. D'Antona ('51) suggests that toxin or antitoxin in-
jected into the cerebrospinal fluid is absorbed into blood and is
transported to peripheral nerves which represent the port of entry
for the toxin.

From the above discussion it is evident that a commonly ac-
cepted theory as to the mode of transport of tetanus toxin is lacking.
Much of the work already done must be repeated and supple-
mented with results obtained by utilizing new and more refined
techniques.

D. The Site and Mode of Action of Tetanus Toxin

It is generally agreed that tetanus toxin attacks neurons of the
central nervous system, however, there is considerable disagreement
as to the site and mode of action of the toxin in local and general
tetanus. The difficulty is due principally to the failure to reach
universal agreement as to the route the toxin takes from the site of
injection or infection to the susceptible organs. Since there is no
generally accepted theory as to the mode of spread of tetanus
toxin it is not surprising that there is no universal agreement as
to the action of tetanus toxin.

Concerning the mode of action of tetanus toxin, Courmont and
Doyon suggested as early as 1893 that the enzymatic action of the
toxin resulted in the production of a secondary toxic substance in
the muscle. Coltorti and Villari ('54) observed that the enzymatic
dephosphorization of ATP ( sodium salt ) in muscle tissue increased
proportionately with the degree of tetanic spasm in the muscle.

Tissue Spaces and Nerve Fibers in the Rat 1461

They suggested that the spasm was due to nervous stimulation re-
sulting from toxin fixed in the central nervous system. Schulze
('49) found that tetanized muscle produces more lactic acid per
unit of tension than does normal muscle, and the solubility of pro-
teins increased in the rigid muscle. Firor, Lamont and Shumacker
('40) and Firor ('47) suggested that a secondary substance was
produced in the spinal cord. This substance is either an altered
form of tetanus toxin or a new material produced by the action of
the toxin on the nervous system. This secondary material is then
transported to the vital centers where it has its lethal effect.

Shintaro Asakura and associates ('50) studied the effect of tetanus
intoxication on the metabolism of the central nervous system.
They were able to recover from the cerebrospinal fluid and blood
certain metabolites such as reducing sugars, acetone bodies, pyru-
vate and alpha-ketoglutarate. This work is interesting in light of
the findings of Firor and associates ('40) working with cross-circu-
lated dogs, and the observations of Matzke and Fedinec ('55) on
parabiotic rats. The latter injected one parabiont and noticed
that the uninjected animal died suddenly without showing any of
the typical symptoms of tetanus. Time of death of both animals
was approximately the same. Zironi ('51) states that some rabbits
injected subcutaneously with toxin die without showing typical
symptoms of tetanus. On the other hand, Friedemann, Hollander
and Tarlov ('41) deny the production of a secondary substance.

Klemm (1889) suggested that an actual alteration in the molec-
ular configuration of motor nerves resulted in a lowering of the
resistance in these nerves to efferent impulses. This in turn
brought about an increased contraction of the musculature of the
affected side. Pochhammer ('08-'09) suggested that focal altera-
tions in the myelin sheaths of both afferent and efferent fibers of the
motor nerve trunk result in "short circuiting." He felt this theory
could account for the symptoms of local tetanus. It is interesting
to note that by using the fluorescent-antibody technique it can be
shown that toxin is present in the myelin sheaths of peripheral
nerves and the protoplasm of the nerve cell body (Matzke and
Riggs, unpublished.)

Harvey ('39) suggested that tetanus toxin affects the metabo-
lism of acetylocholine and cholinesterase content of the muscles
involved. Stevenson ('55) reviewing this problem states that
tetanus toxin is irreversibly fixed at cholinergic endings or at a
point on the unmyelinated terminal fibrils proximal to the acetyl-
choline release region. He favors the latter site as the most likely

1462 The University Science Bulletin

since the toxin does not appear to depress the acetylation of choline
in vitro. There is a decrease in the release of acetylcholine which
Stevenson feels is secondary to a block in transmission of the im-
pulse proximal to the acetylcholine release region. He further
points out that the physiological disturbances resulting from fixa-
tion of the toxin in the region resemble those which follow the post-
traumatic degeneration of a nerve. The period required for com-
plete recovery in local tetanus is within the range of time required
for the repair of a crushed nerve.

Hajek, Godbey and Hines ('47) studied the functional changes in
muscle and nerve resulting from prolonged shortening. They
observed that the function of a muscle was decreased after tenotomy
or tetanus injection. This effect was reversible. In the case of
tetanus the normal functional state was re-established 50 days after
injection.

Before the discovery of the tetanus bacillus it was commonly
held that the seat of action of tetanus was the spinal cord ( Curling,
1836). This opinion was held by many of the early writers such
as Galen, Fernelius, Willis, Hoffman and Lieutaud (Abel and
Hampil, '35). Finally in 1903 Meyer and Ransom formulated a
theory which clearly stated that the site of action was the cell bodies
within the central nervous system. Heiberg, who wrote in 1861,
was one of the earlier writers who felt that the disease involved
the blood, resulting in blood poisoning, the effects of which were
localized in the muscles.

The fact that both local rigidity and general involvement may be
present in tetanus led to the conclusion that the toxin may have a
two-fold action, one on the central nervous system and the other
on the muscles or motor end plates. This view was supported
by Klemm (1889), Veillard and Vincent (1891), von Brunner
(1894), Zupnik ('00-'08) and by many later workers. Abel and
associates ( '34-'38 ) and Penitschka ( '53 ) support this theory. They
feel that the toxin exhibits both a central and a peripheral action
each independent of the other. The motor nerve cells of the spinal
cord, medulla and pons are attacked by the toxin whereas the un-
remitting rigidity of voluntary muscles results from the fixation of
the toxin by the motor end plates. Zironi ('51) presents a similar
view which states that tetanospasmin is fixed by the muscle and
this in turn excites the motor and sensory nerves which facilitates
the fixation of the circulating toxin by the neurons and the cor-
responding segments of the spinal cord.

Tissue Spaces and Nerve Fibers in the Rat 1463

Wright, G. P. ('54) states that although there is general agree-
ment that local tetanus occurs only when the proprioceptive reflex
mechanism is intact, the point or points in this reflex pathway
which are attacked by the toxin have not yet been determined.
He suggests three possible sites of action of the toxin: (1.) the
myoneural junction, (2.) the proprioceptive sensory nerve-endings,
and ( 3. ) the central synapse.

1. Myoneural junction. Harvey ('39) following sectioning of
the sciatic nerve, injected tetanus toxin into the tibialis anticus
muscle of the cat. He noted that upon applying a single shock
to the distal end of the cut nerve repetitive action potentials in the
muscle could be recorded. From this he postulated that the tetanus
toxin brought about changes at the myoneural junction which had
the effect of converting a single excitation into a brief tetanus.
Gopfort and Schaefer ('40), Acheson, Ratnoff and Schoenbach
('42) and Perdrup ('46) could not confirm Harvey's findings.

2. The proprioceptive sensory nerve-endings. Sherrington ('06)
drew attention to the similarities between tetanus and decerebrate
rigidity. This led a number of workers to suggest that local tetanus
is brought about by pathological exaggeration of the sensitivity of
proprioceptive end organs (Liljestrand and Magnus, '19; Ranson,
'28; Schaefer, '44; Perdrup, '46). Klensch and Schlomer ('52) be-
lieve that to demonstrate local tetanus a central excitation by the
toxin and a peripheral nonspecific stimulus is necessary. Vrbova
('54) demonstrated that if the limb of a rat is fixed in an extended
position the effects of local tetanus are exaggerated, whereas if
the limb is fixed in a flexed position the rate of involvement of
tetanus is decreased. She concluded from this that the propriocep-
tors are important in the development of tetanus.

3. Spinal cord synapse. Gumprecht (1893) suggested that tet-
anus acts on the central nervous system resulting in local rigidity.
This is also an important portion of the theory formulated by Meyer
and Ransom ('03). Sherrington ('06) observed that the effects
of strychnine poisoning are similar to those appearing in tetanus.
A large number of workers contributed evidence strengthening this
theory that the action of tetanus toxin is on the spinal synapse
(Friedemann, Hollander and Tarlov, '41; Friedemann and Traub,
'49; Davies, Morgan, Wright, E. A. and Wright, G. P., '54). Vejss
and Kozesnik ('54) lowered the body temperature of animals by
cooling or by injecting anesthetic drugs such as lobeline and choral
hydrate. They noted that the development of tetanus was markedly

1464 The University Science Bulletin

hindered. They felt this indicated that the locus of action of tetanus
toxin is the central nervous system. Doerr and Seidenberg ('35)
injected a sublethal dose of toxin into the lower extremity bring-
ing about a condition of local tetanus. Following this he injected
a lethal dose into the upper extremity. He noted that the symptoms
of local tetanus in the lower extremity were exaggerated by what
he concluded was a flow of impulses from the cervical region of
the cord, brought about by complete poisoning of this region by
the toxin. He referred to this as his dynamic activation theory.
Brooks, Curtis, and Eccles ('55) confirmed the findings of Sherring-
ton that the action of tetanus toxin in the spinal cord closely re-
sembles that of strychnine. They felt that the toxin acts by sup-
pressing all types of synaptic inhibition rather than converting this
inhibition to excitation. The symptoms of general tetanus may be
similarly explained in that the synaptic inhibitory action is sup-
pressed as the toxin spreads through the central nervous system.

E. Treatment of Tetanus

In clinical literature and textbooks many forms of tetanus are de-
scribed. Thus we find descriptions of cephalic tetanus, ascending
tetanus, descending tetanus, tetanus neonatarum in which the
organism enters the umbilicus, puerperal tetanus in which the
mother is infected during delivery, postoperative tetanus from
surgical wounds, and recurrent tetanus in which the symptoms seem
to disappear only to recur (Nielsen, '51). Idiopathic or medical
tetanus is probably nonexistent as such, but is a form of wound
tetanus which develops very late after the original infection has
been overlooked or forgotten (Wechsler, '52). Courtois-Suffit and
Giroux in 1918 presented a broad description of different forms and
variations of tetanus. Generally there are two forms of the disease,
the generalized and the localized. The characteristic manifestation
of local tetanus is a persistent rigidity of a group of muscles. This
state may develop into general tetanus but cases of recovery without
further involvement are also reported. The general clinical picture
in man usually begins with stiffness of the jaw and neck, diffi-
culty in swallowing, stiffness of the legs or arms, restlessness, irri-
tability, headache and convulsions. Fever or chilliness is occasion-
ally the initial evidence of the disease. Later, trismus develops and
the face assumes the characteristic expression, the so-called sardonic
smile (risus sardonicus) . The abdominal and lumbar muscles may
assume a completely rigid state. The contraction of the back
muscles is often so great that episthotonos occurs. The most char-

Tissue Spaces and Nerve Fibers in the Rat 1465

acteristic feature of general tetanus is the development of painful
convulsions which are elicited by the slightest stimuli. Profuse
sweating accompanies the seizures and the viselike contractions of
the chest muscles renders the patient unable to cry out. Dyspha-
gia, cyanosis and asphyxia sometimes appear. The pulse and respira-
tory rates are increased and the temperature commonly rises 2° to 5°
F. Death sometimes occurs from spasm of the respiratory muscles.
More frequently, however, the immediate cause of death is not ap-
parent (Firor, '47). The basal metabolic rate in tetanus is moder-
ately high and exceeds the normal by 114f (Holmdahl and Thoren,
'54). The factors which are important in determining the course
of development of the disease are described by Cole ('40) as: age
and general physique, sex, type and severity of the wound, site of
the wound, incubation period, rate of onset of symptoms, time at
which antitoxin is given, and whether or not prophylaxis antitoxin
has been given. Similarly Firor ('40) points to these factors as
those which can influence the therapeutic result. Cole ( '35a ) classi-
fies clinical tetanus into four groups: 1) local tetanus, only; 2)
generalized rigidity with slight or absent reflex spasm and irri-
tability; 3) generalized rigidity with marked reflex spasm and irri-
tability followed by slowly subsiding generalized rigidity; and 4)
marked and continuous spasms with hyperpyrexia and death from
cardiac or respiratory failure.

The treatment of tetanus is closely allied with the prevailing
theory or theories regarding the etiology of the disease. It seems
impossible to present a standard description of treatment due to
the immense number of clinical reports which, prior to the dis-
covery of antitoxin, merely recommend an empirical treatment
which was based on a meager knowledge of the disease (viz. reports
published in Index Cat. of Surg. Gen. Wash. 1893). Even the
present day treatment is a matter of controversy dependent on new
experimental and clinical evaluations. The fact remains that cer-
tain aspects of the etiology of tetanus are still not satisfactorily ex-
plained. Chackleton ('54) suggests the following steps in the treat-
ment of tetanus: 1) prevention of the absorption of further toxin
by the administration of antitoxin and surgery of the wound; 2) con-
trol of reflex spasm; 3 ) prevention of intercurrent pulmonary infec-
tion; 4) control of fluid and electrolyte balance; and 5) maintenance
of strength. See Mollerfeld ('55), Smith ("55) and Drew ('54) for
review and modern treatment of tetanus.

Treatment with low molecular polyvinylpyrrolidone (Schubert,
'54) has been described. Some authors obtained good results with

1466 The University Science Bulletin

ACTH (Lewis, Satoscar, Joag, Dave and Patel, '54) whereas others
obtained little or no effect ( Greene, Ambrus and Gershenfeld, '53 ) .
Also the method of artificial hibernation has not yet been standard-
ized (Chesni, '53; Weisse, '55). The more commonly used thera-
peutic agents for treatment of tetanus are curare, myenesin, peni-
cillin, sedatives and antisera.

The prophylactic value of antisera and the controversy regard-
ing its therapeutic effect and amount and route of administration
have already been pointed out. Treatment of the wound has to be
conservative. Radical surgical intervention such as amputation
(Lanfranchi, 1295; Rose, 1897; Clemmey, 1898; Smith, '05-'06) does
not alter the ensuing development of tetanus. (McClintock and
Hutchings, '13; Coleman, '30 and Abel and Hampil, '35.) Trache-
otomy has been used as early as 1837 by Curling and 1856 by
Humphrey. Inasmuch as respiratory complications from spasmatic
contraction and respiratory failure are common in severe tetanus
(Turner and Galloway, '49) or in oversedation, clinicians still rec-
ommend its use (Herzon, Killian and Pearlman, '51 and others).

Eccles, Katz and Kuffler ('41); Eccles and Kuffler ('41); Kuffler
( '42a, b, '43 ) described the action of curare on the myoneural junc-
tion as a differential screen, permitting normal impulses to pass while
blocking frequencies outside the normal range of duration. Mclntyre
('49) presented a broad review On curare; its history, nature and
clinical use. According to him Sewell, a veterinarian, first used
curare in animal tetanus. According to Drew ('54) Burrall in 1858
was the first to use this drug in human tetanus. Drew also pointed
out that the modern management of clinical tetanus with curare
began with the work of Florey, Harding and Fildes ('34), Cole
('34), Mitchell ('35) and West ('36). The great value of the
aqueous or longer-acting depot form is emphasized by many workers,
and different forms of application are described ( Cole, '35b; Schles-
inger, '46; Deters and Briggs, '52; Siguier et al '54 and others). The
danger of curare overdosage is discussed by Foregger ('50) and
Pearlstein and Weinglass ('44).

Since Berger and Bradley ('46, '47) introduced it, myenesin has
been widely used in the management of tetanus ( Diaz-Rivera, Trilla
and Pons, '54 ) . A number of theories have been suggested as to its
site of action; namely, diencephalon and brain stem (Stephen, '47),
brain stem and spinal cord ( Schlessinger, Drew and Wood, '48),
selective action on subcortical nuclei, cord and myoneural junction
progressively (Gammon and Churchil, '49), and subcortical efferent
pathways ( Finkelman, '49 ) .

Tissue Spaces and Nerve Fibers in the Rat 1467

Sedation is widely used, mostly in combination with muscle re-
laxant drugs (McDonald, '49). Antibiotics, especially penicillin, is
highly recommended for control of symbiotic organisms. (Diaz-
Rivera, Deliz, Berio-Suarez, '48; Diaz-Rivera, Ramirez, Pons and
Torregrosa, '51).

F. Summary

1. There is general agreement as to the bacteriological and im-
munological characteristics of tetanus. However, many questions
concerning the chemical structure of the toxin and its action in vivo
remain to be answered.

2. Pathologic changes resulting from tetanus apparently do not
follow a set pattern from individual to individual. This is further
complicated by the presence of post-mortem changes in most fatal
cases of human tetanus, however, it is apparent that following re-
covery from a sublethal dose of toxin, no permanent alterations in
the central nervous system occur.

3. The treatment of tetanus has been empirical and aimed at re-
ducing the circulating toxin by administering antitoxin, preventing
secondary invasion, eliminating muscle spasms and treating the site
of infection.

4. Considerable controversy exists over the route followed by the
toxin from the site of injection or infection to the central nervous
system. There are a number of supporters for each of the following
pathways:

a. the nerve fibers

b. the perineural tissue spaces

c. the perineural lymphatics

d. the vascular stream

e. a combination of any two or more of the above.

5. Disagreement also exists as to the mode and site of action of
the toxin. The following are some of the theories that have been
proposed :

a. Direct action on the central nervous system.

b. Peripheral action on muscles, motor end plates or sensory end-
ings.

c. Dual action on the periphery and central nervous system.

d. Acts as an enzyme altering the activity of the cell.

e. The toxin is transformed or results in the production of a sec-
ondary substance which in turn is toxic.

It is apparent from the above discussion that a great deal of ex-
perimental work must be done to answer the many questions still

1468 The University Science Bulletin

remaining in all phases of the tetanus problem. The objective of the
following experiments is to help clarify some of the problems related
to the spread of tetanus toxin in the body.

ACKNOWLEDGMENTS
This investigation was supported by a Research Grant from
the National Institute of Neurological Diseases and Blindness of
the Public Health Service. Toxin and antitoxin was supplied
through the courtesy of Eli Lilly & Company, Indianapolis. Evipal
was supplied through the courtesy of Winthrop Chemical Company,
Inc., New York.

MATERIALS AND METHODS

Rats from a highly inbred Sprague-Dawley strain were used in
all the experiments. An attempt was made to utilize littermates
for each experimental group. If this was not possible, animals of
approximately the same age and weight were selected. Both sexes
were represented in each experimental group. Diet, temperature
and other environmental conditions were carefully controlled.

The tetanus toxin utilized in the experiments had a potency of
640,000 MLD/cc and the antitoxin 4,800 American units per cc.
Evipal Soluble injected intraperitoneally was used as the anesthetic.
Immediately before using, all these substances were diluted with
triple distilled water. The toxin and antitoxin were injected intra-
muscularly, intravenously or intraneurally with a micrometer syringe
to which a 30 gauge hypodermic needle was mounted.

Experimental Series I. In order to determine the dosage of toxin
necessary to kill fifty percent of the animals injected (LD 50),
eight groups of ten animals each were inoculated intramuscularly
with equal volumes of the following dilutions of the toxin: 1:300,
1:400, 1:450, 1:475, 1:500, 1:550, 1:575, 1:600. The volume de-
livered in each case was 0.0015 ml/100 gr. body weight.

Experimental Series II. The relationship of the time and pattern
of development of symptoms and time of death to the site of in-
oculation was studied in three groups of ten animals each. Each
animal of the first group received 2 MLD of toxin in the right
gastrocnemius muscle. In animals of the second group, the right
sciatic nerve was exposed and 2 MLD of toxin was injected peri-
pherally into the nerve. Care was taken to prevent leakage and
backflow from the nerve. The procedure was carried out with the
aid of a binocular microscope. In animals of the third group 2
MLD of toxin were injected into the right saphenous vein of each.

Tissue Spaces and Nerve Fibers in the Rat 1469

The vein was exposed and the inoculation observed with a binocular
microscope. The type of symptom and time of appearance was
recorded as well as the time of death.

Experimental Series III. At 21 days of age the right sciatic nerve
was sectioned and resutured in each of 31 rats. From 200 to 375
days were allowed for regeneration. The degree of functional
recovery was determined by noting the reaction to nociceptive
stimuli and direct electrical stimulation of the sciatic nerve proximal
to the suture. If die nerve fiber plays a role in the transport of the
toxin, the rate of passage of the toxin along the small regenerated
fibers should be slower than that along the normal nerve fibers.
Two MLD of toxin were injected into the nerve distal to the suture.
The time of appearance of symptoms was noted. The animals
were sacrificed and the number of regenerating fibers distal to the
suture determined.

Experimental Series IV. To determine whether the number of
fibers in the sciatic nerve influenced the rate of passage of the toxin
six groups of five animals each were used. Varying numbers of
nerve fibers were destroyed with an electro-coagulator. One of the
following strengths of current were applied to the sciatic nerves
of members of each group for the indicated time: 0.5 mamp, 15 sec;
1 mamp, 10 sec; 1 mamp, 15 sec; 1 mamp, 20 sec; 1 mamp, 25 sec;
1 mamp, 30 sec. Ten days were allowed for regeneration of the
connective tissue and blood vessels. Two MLD of toxin were in-
jected into the nerve distal to the lesion. Immediately after death
the sciatic nerve was removed and sections distal to the lesion
were stained using the silver method of Romanes ('50). The num-
ber of intact fibers was compared to that found in the normal left
sciatic nerve.

Experimental Series V. The effect of complete destruction of the
nerve fibers on the spread of tetanus toxin was studied in 29 rats.
The right sciatic nerve was crushed over a distance of 4 mm with
a flat forceps. Between one and 100 hours after crushing 2 MLD
of toxin were injected into the right sciatic nerve below the crushed
area in 14 animals. Eight rats were inoculated at intervals between
100 and 200 hours and 7 between 200 and 300 hours after crushing.

Experimental Series VI. In order to determine the role of the
tissue spaces of peripheral nerves in the spread of tetanus toxin 80
rats were divided into seven groups. Five controls were inoculated
in the right gastrocnemius with 2 MLD of toxin. Fifteen other con-
trols were inoculated similarly with 2 to 5 MLD 48 hours after the

1470 The University Science Bulletin

genitofemoral, obturator, femoral, lateral cutaneous of the thigh,
posterior cutaneous of the thigh, and hamstring branches of the
sciatic nerves were sectioned.

In twenty animals the above nerves were sectioned and the
epineurium removed from the sciatic nerve in the middle of the thigh
for a distance of approximately 1 cm. This was accomplished by
exposing the nerve and carrying out the following procedure with
the aid of the binocular microscope. A shallow circular incision
through the epineurium was made with an iridectomy knife. The
cut end of the epineurium was secured with a fine forceps, peeled dis-
tally for the required distance and dissected away. The following
day the sensitivity of the skin in the distribution of the sciatic nerve
was tested. Those animals that gave a good response were injected
with 2-5 MLD of toxin in the gastrocnemius of the operated side. In
another group of ten animals the above procedures were repeated,
however, in addition, the perineurium was carefully teased open with
a blunt hook.

In another group of fifteen animals the right lower extremity was
denervated and a segment of the epineurium removed from the
sciatic nerve perineurium and opened as described above. Twenty-
four hours later 5 MLD of toxin were injected into the right gastroc-
nemius muscle. Two hours after this 28.8 times the amount of anti-
toxin necessary to neutralize the injected toxin was inoculated into
the left saphenous vein. Animals which survived this procedure and
showed no signs of tetanus were reinjected with 5 MLD of toxin in
the sciatic nerve proximal to the area from which the epineurium was
removed.

In a group of five animals the right lower extremity was dener-
vated and a segment of the epineurium removed from the sciatic
nerve. Five MLD of toxin were injected into the nerve proximal to
the bare area and antitoxin into the left saphenous vein. Another
group of ten animals was treated similarly except that the epineurium
was left intact and the toxin injected into the gastrocnemius muscle.
In all the animals of this experiment the time and pattern of develop-
ment of symptoms were recorded.

RESULTS

Experimental Series I. The results of the first experiment are
recorded in Table 1. The data reveal that the least amount of toxin
necessary to kill 100 percent of the animals was 0.0015 ml/100 gr.
body wt. of the 1:400 dilution. This is referred to as the minimum
lethal dosage (MLD). The medium lethal dosage (LD 50) is that

Tissue Spaces and Nerve Fibers in the Rat 1471

amount of toxin which kills 50 percent of the animals. In this ex-
periment this dosage was found to be 0.0015 ml/100 gr. of body wt.
of the 1:475 concentration. The same dosage of the 1:550 dilution
failed to kill any of the animals although all developed symptoms of
tetanus. This is referred to as the dosage of maximal tolerance
(LDO).

Experimental Series II. Following injection of the toxin into the
gastrocnemius or sciatic nerve the first symptoms appeared in 18-26
hours. The first sign was that of hypersensitivity of the inoculated
extremity which was held in a flexed position to prevent contact with
objects. This was gradually replaced by spastic extension at all joints
of that extremity. A few hours later the tail became rigid arching
toward the side of the involved extremity which was moderately ab-
ducted. Within 40-50 hours following injection, lumbar scoliosis to-
ward the affected side appeared. At this time the animal may become
extremely hypersensitive responding to the slightest stimulus with
convulsive seizures. These seizures may result in injury with blood
appearing in the mouth, nostrils, and eyes. The scoliosis disappeared
when the opposite extremity became involved. However, pleuro-
thotonos directed ventrally resulting from spasticity of the thoracic
musculature appeared at 60 hours. There followed in rapid order,
spasticity of the forelimbs, trismus, contraction of the facial muscula-
ture, immobility of the vibrissae, and depression of the ears. Death
occurred in 68-74 hours following inoculation.

It is apparent from the above description that intramuscular or
intraneural injection of toxin results in a progressive involvement
of the musculature of the body in a segmental manner. This is re-
ferred to as local tetanus which may be either ascending ( as in the
above case) or descending, depending on the site of injection. This
is in contrast to blood-borne tetanus which follows the intravenous
injection of toxin.

When 2 MLD of toxin is injected intravenously the first symptoms
appeared in 54-62 hours. The first signs involved the facial, masti-
catory, and some of the axial musculature. A few hours later
pleurothotonos appeared due to the spastic contraction of the back
and neck muscles. Death occurred in 64-72 hours. At this time the
animal may or may not show involvement of the extremities. In
contrast to local tetanus the site of intravenous injection had no bear-
ing on the sequence of involvement of muscular groups, however, in
both cases there was a definite period of time between injection and
appearance of the first symptom. This is referred to as the latent
period. This period was two to three times longer in blood-borne

1472 The University Science Bulletin

tetanus than in local tetanus although the time of death was about
the same in both groups. The latent period may be shortened in
both groups by increasing the dosage and, in addition, in local tet-
anus, by injecting closer to the central nervous system. At the time
of death in ascending local tetanus, virtually all the musculature of
the body was involved; this is referred to as general tetanus. This
condition rarely occurs in descending and blood-borne tetanus.

Experimental Series III. The degree of functional nerve fiber re-
covery, following 200 to 375 days regeneration time, varied con-
siderably; however, all animals responded to nociceptive stimuli and
direct stimulation of the regenerated sciatic nerve. The number of
regenerating fibers averaged 6,289, the range being from 3,800 to
7,867. This is to be compared with an average of 5,958 and range
of 5,302 to 6,564 in the normal intact sciatic nerve. The diameter
of the regenerating fibers was less than that of normal axons.

Following intraneural injection of toxin distal to the suture, the
animals developed the first symptoms of local tetanus in 30 to 38
hours. These appeared in the injected extremity. Typical ascend-
ing tetanus followed; however, the time sequence was considerably
longer than in the controls. Death occurred at 116 to 154 hours.
There was no correlation between the time of appearance of symp-
toms and death, on the one hand, and regenerating time and number
of fibers, on the other.

Experimental Series IV. When variable numbers of fibers of the
sciatic nerve were destroyed by electrocautery, the response to
nociceptive stimuli and direct electrical stimulation decreased with
an increase in the area of the nerve damaged. Histologic sections
of the nerve distal to the lesion revealed that the number of intact
fibers varied from 635 to 5,435, however, in no instance did the num-
ber of intact fibers approach that found in a normal nerve. Unlike
the condition found in regenerating nerves, the diameter and gen-
eral appearance of the intact fibers was comparable to that of those
found in the normal nerve. Another important point of comparison
with regenerating nerves is that relatively little connective tissue was
deposited at the site of the lesion produced electrically as compared
with that found at the suture in regenerating nerves.

When toxin was injected into the sciatic nerve, distal to the
lesion, local tetanus appeared in the injected extremity in 22 to 36
hours. Typical ascending tetanus followed with death occurring at
133 to 210 hours. Table 2 indicates there is no relationship between
the above time sequence and strength of current used to produce
the lesion. Furthermore, there was no correlation between the

Tissue Spaces and Nerve Fibers in the Rat 1473

number of intact fibers and the time sequence of development of
symptoms.

Experimental Series V. The results of this experiment are re-
corded in Table 3. Histologic sections of the nerve distal to the
crushed area revealed only degenerating fibers. Despite this fact,
local tetanus appeared in the injected extremity in 23 to 30 hours,
following inoculation of the sciatic nerve distal to the crushed area.
Ascending tetanus, followed by death, occurred at 175 to 245 hours.

Experimental Series VI. The data from this series are recorded
in Table 4. In control animals with an intact sciatic nerve and con-
nective tissue sheaths, intramuscular injection of toxin resulted in
local ascending tetanus. When the epineurium is removed from the
sciatic nerve and the other nerves to the same extremity cut, blood-
borne tetanus resulted in 15 of the 20 animals injected with toxin in
the gastrocnemius. The remaining five developed local tetanus.
Typical ascending tetanus did not proceed beyond the lumbar seg-
ments. At this time blood-borne tetanus was superimposed upon
this condition.

If, in addition to removing the epineurium, the perineural septa
were teased open, all animals developed blood-borne tetanus fol-
lowing intramuscular injection. In these animals tetanus could be
prevented if antitoxin was administered intravenously within two
hours after injection of the toxin. Antitoxin did not prevent the
occurrence of local tetanus if the epineural sheath was intact or if
the toxin was injected into the nerve proximal to the bare area, how-
ever, the resulting ascending tetanus was arrested by the time
pleurothotonus appeared. The animals survived although spastic
contractions of the involved musculature persisted for several
months.

In the animals where the epineurium had been removed the re-
sponse to nociceptive stimuli applied to the area of distribution of
the sciatic nerve was normal. Histologic sections through the bare
area of the nerve revealed that the great majority of the axons were
intact and the epineurium absent.

DISCUSSION

Basing his conclusions on the results of others as well as his
own work, D'Antona ('51) formulated a theory on the mode of
spread of tetanus toxin. He stated that at the site of injection or
production, the toxin is absorbed by both the nerve endings at the
motor end plates and the blood capillaries. From the former the
toxin passes into the central nervous system via the efferent nerve

1474 The University Science Bulletin

fibers. The motor neurons in the segments innervating the inocu-
lated extremity are the first to be attacked by the toxin. This results
in spastic contraction of the muscles innervated by these neurons.
The toxin then follows axons within the central nervous system
to the opposite side as well as up and or down depending on the
site of injection. This results in a progressive involvement of neu-
rons of adjacent segments and contraction of the muscles innervated
by the corresponding neurons. Death results when the vital centers
of the brain stem are reached by the toxin.

From the capillaries the toxin is carried throughout the body in
the vascular stream. Naked nerve endings of motor end plates
throughout the body absorb the toxin which is then transported
to the central nervous system by all somatic efferent fibers. The
first musculature to be affected will be that with the shortest ef-
ferent fibers. Death results from involvement of the vital centers.

The latent period (the time elapsing between injection and the
appearance of the first symptoms) is two to three times longer in
blood-borne tetanus than in local tetanus, however, the time of
death is approximately the same.

The results of the experiments reported here support many as-
pects of D'Antona's theory. The time sequence and pattern of ap-
pearance of symptoms following intramuscular and intravascular
injection of toxin agrees with the above descriptions. Tetanus re-
sulting from intramuscular injection of toxin is referred to as local
tetanus, whereas, intravascular administration results in blood-
borne tetanus. These results therefore support the hypothesis that
the toxin reaches the central nervous system via peripheral nerve
trunks.

The principal point of disagreement concerns the element of the
peripheral nerve trunk responsible for the transport of the toxin.
The results of these experiments do not support D'Antona's axon-
transport theory. Toxin reaches the central nervous system when
all axons have degenerated and only the connective tissue is intact.
In these experiments the intravascular toxin may be eliminated by
the intravenous administration of antitoxin. Furthermore, the rate
of passage of the toxin to the central nervous system appears to be
independent of fiber size or number.

Symptoms of local tetanus can be prevented only when the con-
nective tissue of the epineurium is removed and the perineural
septa teased open. In this case blood-borne tetanus results follow-
ing intramuscular injection of toxin unless antitoxin is administered
intravenously.

Tissue Spaces and Nerve Fibers in the Rat 1475

Baylis et al ('52) concluded that the toxin reaches the central
nervous system via the tissue spaces of peripheral nerves. Utilizing
substances which stimulated the deposition of connective tissue in
the nerve thus blocking the tissue spaces, they were able to prevent
the appearance of symptoms of local tetanus following injection of
toxin into the nerve distal to the sclerosed area. The results of the
experiments reported here support this conclusion. The removal
of the epineurium removes or opens the tissue spaces so that the
toxin is prevented from passing along the nerve. It is absorbed
into the vascular stream either at the site of injection or it infiltrates
the tissue spaces surrounding the bare area of the nerve and is
there absorbed by the blood. Deposition of scar tissue at the site
of crushing or suturing a nerve partially blocks the tissue spaces
and delays the onset of symptoms of local tetanus.

Abel and Iris coworkers ('34-'38) concluded that the only way
tetanus toxin could reach the central nervous system was directly
from the vascular stream. Friedemann and associates ( '39a, b, '41 )
state that the blood-brain barrier is impermeable to the toxin. The
results of the experiments reported here do not directly eliminate
the vascular stream as an alternate route, however, toxin can reach
the central nervous system even though all circulating toxin is
neutralized by antitoxin. The pattern of development of symptoms
of blood-borne tetanus suggests that the toxin passes in via peri-
pheral nerves since the first muscles involved are those with the
shortest axons. Wright, G. P. ('53) also points out that the first
musculature involved is the most active. He believes that activity
increases the pressure gradient between the periphery and central
nervous system and thus the rate of flow of the toxin in the tissue
spaces.

Further research is necessary to determine the role of the blood-
brain barrier in the passage of tetanus toxin into the central nervous
system. Furthermore, the present work points out the necessity of
determining more accurately the morphologic relationship of the
tissue spaces of the peripheral nerves to the peripheral and central
tissue spaces and cerebrospinal fluid chambers. Investigations de-
signed to clarify these relationships are being carried out.

SUMMARY AND CONCLUSIONS

1. The following three forms of tetanus were described:

a. Local tetanus: A localized group of muscles are first in-
volved unilaterally. The condition spreads segmentally in
a caudal and cephalic direction as well as contralateral!}'

1476

The University Science Bulletin

4.

5.

6.

until death ensues. This form results from intramuscular or
intraneural injection of toxin.

b. Blood-borne tetanus: This condition follows intravascular
injection of toxin. The masticatory, facial and mid-line mus-
culature are first involved. Death resulting from respiratory
or circulatory collapse occurs before the muscles of the ex-
tremities show any signs of tetanus.

c. General tetanus: This term is used to describe the condition
when all the muscles of the body are involved. It usually
appears as the terminal stage of ascending local tetanus. It
rarely follows blood-borne or descending local tetanus.

The time of appearance of the symptoms of local tetanus is in-
dependent of the number or size of the nerve fibers to the ex-
tremity injected.

Local tetanus will occur in the absence of nerve fibers to the
injected extremity even though toxin in the vascular stream is
neutralized by antitoxin.

The symptoms of local tetanus are delayed when scar tissue
forms on the nerve as a result of crushing or suturing.
Blood-borne tetanus appears when the connective tissue is re-
moved from the nerve to the injected extremity, however, this
can be prevented if antitoxin is administered intravenously.
It is suggested that the only route to the central nervous system
for tetanus toxin is by way of the tissue spaces in peripheral
nerve trunks.

Table 1 — Determination of MLD, LD50 and LDO

Number of animals

Dilution

of

toxin

Number
developing
symptoms

Number

dead in

144 hours

10

1:300
1:400
1 :450
1:475
1:500
1 :550
1 :575
1:600

oooooooo

10

10

10

10. . .

7

10

10

5
2

10. . .

0

10

0

10. .

0

Dosage— 0.0015 ml/100 gr. body wt.

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Tissue Spaces and Nerve Fibers in the Rat 1479

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THE UNIVERSITY OF KANSAS

SGIENGE BULLETIN

Vol. XXXVIII, Pt. II] March 20, 1958 [No. 24

The Family Succineidae (Gastropoda: Pulmonata)

in Kansas

BY

Charles D. Miles
TABLE OF CONTENTS

PAGE

Introduction 1499

Acknowledgments 1502

Review of Literature 1502

Methods 1503

Characteristics of the Family Succineidae 1505

Jaw 1505

Shell 1506

Genitalia 1506

Key to the Species of Family Succineidae in Kansas 1510

Accounts of Species 1511

Succinea ovalis Say 1511

Succinea concordialis Gould 1514

Succinea vaginacontorta Lee 1517

Succinea pseudavara Webb 1519

Oxyloma return (Lea) 1521

Quickella vagans (Pilsbry) 1525

Quickella ivandae Webb 1529

Origin of Species of Succineidae in Kansas 1530

Summary 1531

Literature Cited 1532

Arstract. This study consists of a review of the snails of the Family
Succineidae in Kansas. Dissections of the soft parts have been made and
used with the shells as a basis for taxonomy. Seven species are recognized.
The literature is reviewed.

INTRODUCTION

This study was undertaken in an attempt to determine the kinds
and distribution of species of the Family Succineidae (Pulmonata
== Gastropoda) occurring within the State of Kansas. Prior to 1954,

(1499)

1500 The University Science Bulletin

at which time this problem was selected, nine species of living suc-
cineids had beben reported within the State. With two exceptions,
the identification of these species was based upon characters of
shells, without consideration being given to the soft anatomy of the
animals.

Until relatively few years ago, all land snails in North America
were treated systematically by considering the details of the shell.
Tryon (1866) does not mention the soft anatomy of most species
of land snails in his monograph, including the members of the
Family Succineidae. Three years later, Binney and Bland (1869:
265) characterized the soft parts of Succinea as follows: "Animal
resembling the animal of Helix, but shorter. Eye-peduncles short,
expanded at their base or conic; tentacles very short and small.
Respiratory foramen in the mantle, in the angle at the posterior part
of the aperture of the shell." In their discussion of the various
species of Succinea in their monograph, mention of the living animal
is made only in connection with Succinea ovalis Gould ( = Oxyloma
decampi gouldi ) ( op. cit. ) . Only a terse description of the external
appearance of the animal is included in the characters given for this
species.

In many families of snails, the shell seems to be sufficient for
accurate diagnosis of the various genera and species. The Family
Pupillidae may be cited as an example, for the numerous species,
some of which are widely distributed in North America, and else-
where, are separated satisfactorily by such features as size of the
umbilicus of the shell, presence or absence of lamellae in the
aperture, number and position of lamellae ( if present ) , and similar
characters of the shells ( Franzen and Leonard, 1947 ) . In addition,
color and markings on the shell and number of whorls present are
useful taxonomic tools in many families of land snails.

In the Family Succineidae, the shells are not diagnostic; they
are simple, lack ornamentation, lamellae, and other characters use-
ful for purposes of identification. Also, shells vary according to
age of the animal and according to the environment. Although
Succinea ovalis Say is readily identified at all ages by the shell, the
shells of the younger animals of some other species are similar
enough to create confusion. This confusion, due to similarities of
shells, exists between genera, as well as species. Separating young
Quickella vagans and young Succinea concordialis, by characters of
their shells, is a perplexing task, especially as these two species are
commonly found inhabiting the same locality.

Family Succineidae (Gastropoda: Pulmonata) 1501

Several workers have observed that accurate identification of al-
most every species of succineid requires examination of the genitalia,
especially of the male organs (Pilsbry, 1948; Lee, 1951a). Because
of definite differences in the male genitalia of Succineidae, three
genera, Succinea, Oxyloma, and Quickella, are currently recognized
in North America; not many years ago, all members of this family
were placed in the nominate genus, Succinea. The distinguishing
features of the three genera are discussed in a later section of this
report.

The soft anatomy of members of the Family Succineidae was not
utilized for taxonomic purposes to any great extent until Quick
(1933) published results of a study of European members of the
family. His systematic treatment of the family was based mostly
on anatomical characters, particularly of the genitalia.

Using the genitalia of the snails as the basis for his study, Lee
(1951a) reduced the number of species of succineids recognized in
Michigan from fourteen, which had been described solely on the
basis of shells, to three species. Lee reported that the appearance
of the shell is too variable to be of taxonomic use in most species of
the Family Succineidae in Michigan. The present study tends to
support this view, although there are several species of succineids
in Kansas that seemingly do not occur in Michigan.

It seems paradoxical that the Family Succineidae has been rela-
tively neglected in North America, for species are numerous in
this country. Where succineids occur, they are usually far more
numerous in total number than all other snails in the locality. I
have visited localities in Kansas where the only limit to the number
of succineids collected, perhaps consisting of one or two species,
was the time factor and the energy of the collector. As cases in
point: at the artesian marsh, 1% miles south of Muscotah, Atchison
County, in July, 1955, the supply of Oxyloma retusa was seemingly
inexhaustible; roadside ditches along Kansas Highway 7, near Iowa
Point, Doniphan County, Kansas, harbored O. retusa in incredible
numbers in April, 1956; at the University of Kansas Natural History
Reservation, several miles northeast of Lawrence, Douglas County,
great quantities of Succinea concordialis were along South Creek in
May, 1956; at any one of a number of localities near Lawrence, such
as "Haskell Bottoms," Quickella vagans was found to be abundant.

In spite of the abundance of Succineidae, few papers of any
significance concerning the family have been published in North
America; there is one unpublished doctorate dissertation concerning

1502 The University Science Bulletin

this family (Lee, 1951a). Numerous locality records pertaining to
various species of the family are found in the literature, but the
validity of many of these reports is questionable because, in most
instances, identifications were based on the shell. Likewise, certain
reports concerning the ecology of succineids must be taken ad-
visedly, because of uncertainties of identification.

The primary objective of my study was to report the species of
the Family Succineidae existing in Kansas, on the basis of details
of their soft anatomy, especially their genitalia. The shells of some
species included in this report are so variable as to be of little
taxonomic use; this is especially true of Quickella vagans, the shells
exhibiting considerable variation over the State.

Investigation has failed to verify the occurrence of Succinea avara
Say and S. grosvenori Lea; S. avara has heretofore been considered
the most prevalent species of Succineidae in Kansas. Perhaps the
most significant finding of my study is the widespread occurrence
in Kansas of Quickella vagans; its shells have been assigned to one
or several species of the genus Succinea by previous workers.

Future diligent collecting may reveal the presence of additional
species of the Family Succineidae to be added to the list of mollusks
of Kansas. Detailed ecological studies of all species will be desir-
able, for most succineids in the State seem capable of inhabiting
more than one distinct kind of habitat. It is hoped that additional
attention will be focused upon the Family Succineidae in Kansas
and elsewhere in North America.

ACKNOWLEDGMENTS

I am grateful for guidance and suggestions furnished by Professor
A. Byron Leonard; he suggested that I study the Succineidae and
accompanied me on several collecting trips. The shells shown in
Plate I were photographed by him. Professors E. Raymond Hall
and John A. Weir read the manuscript and offered valuable sug-
gestions, as did Thane S. Robinson also. My wife, Virginia, col-
lected many specimens, typed the manuscript and assisted other-
wise.

REVIEW OF LITERATURE

In Kansas, living gastropods, in general, have been relatively
little studied, considering their abundance in species and in num-
bers of individuals. The earliest report of species of the Family
Succineidae is by Binney and Gray (1885:55); they quote F. W.
Cragin and J. B. Quintard on the occurrence of Succinea lineata
Binney (= S. grosvenori Lea) near Topeka, Kansas, and at Silver

Family Succineidae (Gastropoda: Pulmonata) 1503

Lake Township, Shawnee County. In addition, they listed S. avara
Say as a "widely distributed species" and S. haydeni Binney ( =
Oxyloma) from Silver Lake Township as "rather common." A year
later, Call (18S6:202) reported S. obliq.ua Say (=S. ovalis Say)
from Neosho County, Kansas. Hanna (1909:96) in a list of the
mollusks of Douglas County, Kansas, reported the occurrence of
S. avara Say ("not common"), S. grosvenori Lea, S. rusticana Gould,
and S. stretchiana Bland. Hanna considered S. stretchiana "our
common Succinea" (loc. cit.). In the same number of the Nautilus
as Hanna's paper, Baker (1909:94) noted the occurrence of S. avara
from Anthony, Harper County, Kansas.

Nothing further was contributed to the literature concerning the
Succineidae in Kansas until 1942, when Franzen and Leonard
(1942:339) reported on the mollusks of Kingman County. They
listed S. concordialis Gould and S. grosvenori Lea as occurring
within that county. The following year, Alice E. Leonard (1943:
240) reported S. concordialis and S. haydeni (= Oxyloma) from
Meade County, Kansas. Dorothea Franzen (1944:272), in her
paper on mollusks previously unreported from Kansas, mentioned
that Succinea haydeni ( = Oxyloma ) inhabited the artesian marsh
south of Muscotah, in Atchison County, in association with various
other gastropods. Two years later, Leonard and Leonard (1946:
120) recorded S. avara as living "along streams in grass and sedges"
in Greenwood County, Kansas. Leonard and Goble (1952:1036)
in a paper on the mollusks inhabiting the University of Kansas
Natural History Reservation, situated 5/2 miles northeast of Law-
rence, in Douglas County, reported S. avara and S. concordialis.

In all of the identifications noted above, succineids were identi-
fied only by characteristics of the shells. S. vaginacontorta Lee
(1951b) and Quickella wandae Webb (1953) were described from
Kansas and both authors followed the modern procedure of bas-
ing their diagnoses on characteristics of the genitalia of the animals.
Leonard (1950:23) has discussed the fossil record of various mem-
bers of the Family Succineidae in Kansas, particularly pertaining to
the Pleistocene. He has described one new fossil species, Oxyloma
navarrei ( loc. cit. ) from Kansas, in deposits of Kansan ( Mid-Pleisto-
cene) age.

METHODS

Primary attention has been centered upon the soft anatomy of
the animals collected in the course of this study. Living speci-
mens of snails were collected from numerous localities over much
of the state of Kansas from April, 1955, until August, 1956. Two

1504 The University Science Bulletin

rather extensive collecting trips were made possible by the State
Biological Survey of Kansas.

Early in this study, difficulty was encountered in keeping alive
snails of certain species until they could be examined in the
laboratory. This was especially true of Oxyloma retusa. Trans-
porting living snails from the field to the laboratory frequently en-
tailed a high mortality rate, which was partly overcome by placing
snails in airtight containers. Undoubtedly, the loss of moisture
from perforated jars contributed to the death rate of the animals.
As an added precaution, living snails were stored in a refrigerator
in the laboratory, which aided considerably in maintaining them
until they were examined. The use of a portable ice chest when in
the field prevented the great majority of snails from dying; this
chest was used successfully on a five day trip over the State in
June, 1956.

In the field, snails from a single locality were placed in a container
and labeled with a field number, locality, and date. At the same
time, pertinent information was entered in a field book on a page
bearing the field number, locality, date, and additional information
pertaining to ecology, weather conditions, associated species of
snails and relative abundance of the snails.

Prior to dissection, snails were killed by immersion in hot water
after first allowing the animals to extend themselves on a spoon
or section-lifter. The temperature of the water should be at, or
near, the boiling point. Seven seconds, or less, in hot water is
sufficient to relax the columellar muscle. The animal is then easily
removed from its shell by inserting a pin in the foot and gently
pulling the animal out.

In the early stages of this study, the snails were dissected in water
after removal from the shell. A liquid medium prevents desicca-
tion, and aids immeasurably in differentiating the various internal
organs. Later, the technique of dissecting in 70 percent alcohol,
suggested by Abdel-Malek (1954:104), was followed. Abdel-
Malek reported that "dissection in water was found to be inadvis-
able because various organs of the snail (Helisoma trivolvis Say)
. . . and the genitalia particularly tend to absorb considerable
water" (loc. tit.). Dissected soft parts were preserved for future
study in small glass vials containing 70 percent alcohol, bearing a
label with the field number, locality, specific name, and date of col-
lection; shells of the snails were placed in cardboard containers,
bearing a corresponding label.

Family Succineldae (Gastropoda: Pulmonata) 1505

Genitalia of representatives of the various species of Succineidae
examined in this work have been mounted on slides for a permanent
record. For study purposes, storage in alcohol is preferable.

Although not given critical attention in this study, because of
their questionable taxonomic use, four measurements of the shells
of living specimens were made: greatest length (L), greatest diam-
eter (D), length of aperture (LA), and diameter of aperture
(DA). From these four measurements, three ratios are obtained:
the ratio of diameter of shell to length of shell (D/L); ratio of
length of aperture to length of shell ( LA/L ) and ratio of diameter
of aperture to length of aperture ( DA/LA ) .

Measurements were made to 0.1 mm by means of a vernier
caliper. Although these ratios have not been treated statistically,
the average ratios obtained from a series of shells may be com-
pared with the average ratios of shells from other series. Certain
trends are thus brought to light, which will be discussed in the
treatment of the several species. In their present form, nevertheless,
these ratios have little taxonomic value, because specimens of
different species may yield similar ratios.

The standard reference work used in my study is Vol. 2, part 2,
of Pilsbry's classic monograph of The Land Snails of North America
(1948), in which the anatomy of most known species of North
American Succineidae are considered.

The genitalia of the several species were drawn by a technique
that conveniently results in correct proportions. Intact reproduc-
tive systems were mounted on glass slides and projected on white
paper by means of a photographic enlarger; the outlines were traced
in ink. This procedure was followed in all cases except that, for
technical reasons, the outlines for Figures 4, 7, and 9 were drawn
freehand. The jaws, Figures 10a and b, were outlined by means
of camera lucida.

CHARACTERISTICS OF THE FAMILY SUCCINEIDAE

The Family Succineidae includes a large group of terrestrial
pulmonate snails that are world-wide in distribution. Species are
most numerous in North America, southern Asia, and Hawaii (Pils-
bry, 1948:772). Because of several distinctive characters, dis-
cussed in detail by Pilsbry (loc. cit.), the family is not considered
to be closely related to any other family of pulmonate gastropods.

Jaw (Fig: 10a, b) : Perhaps the most distinctive character, found
only in the Family Succineidae, is the large, squarish accessory plate

1506 The University Science Bulletin

dorsal to the cutting edge of the jaw, which ordinarily possesses a
single median projection on the cutting edge. In several species of
Succinea, ribs are present on the anterior surface of the jaw, but
in Kansas, only Succinea ovalis possesses such ribs.

Shell ( plate I ) : The shell, which is similar in general form in all
species in Kansas, has been primarily responsible for the present
confused state of taxonomy in this family. The shell is simple and
lacks distinctive color patterns or ornamentation which might be
useful taxonomically. It is thin, transparent, and imperforate. The
spire is short, and there are up to four whorls in specimens from
Kansas. A characteristic feature of the shell is the relatively large
aperture, ranging from 50 percent to over 80 percent of the length
of shell.

The color of the shell varies among members of a single popula-
tion, but in general, is some shade of yellow or brown. The family
has long been referred to as "the amber snails" ( Lee, 1951b ) . The
color of the shells of any species of the Family Succineidae should
not be primarily relied upon in identifying species. Although the
shells of Succinea concordialis have a ruddy or reddish apex, the
same condition has been observed in the shells of other species.
Furthermore, the color of the shell often changes after death of the
animal.

Genitalia (Figs. 2-9): As noted earlier, the reproductive system
furnishes the most accurate means of distinguishing species of Suc-
cineidae. The genitalia of all species of this family may be termed
simple, in that accessory organs, such as dart sacs, muciparous
glands, and flagellae, present in some other families of snails, are
lacking in the Succineidae.

The succineids are monoecious, as are most pulmonate gastropods.
The male and female reproductive systems open to the outside of
the animal through a common genital atrium, which is situated im-
mediately behind the right eyestalk. Since male and female systems
join at a common atrium, the intact reproductive system may be
removed from the animal with comparative ease. A characteristic
feature shared by all species of Succineidae is that the darkly pig-
mented retractor muscle of the right eyestalk passes between the
junction of penis and vagina.

Since detailed anatomical and functional studies of the reproduc-
tive organs of Succineidae are not available for most species, it is
necessary to resort to interpretations based on studies of snails be-
longing to other families. The unreliability of such extrapolations
is obvious.

Family Succineidae (Gastropoda: Pulmonata) 1507

The hermaphrodite gland (ovotestis) is at the extreme posterior
end of the animal and occupies the apex of the shell in the living
state. Both ova and sperm are formed in this gland. Hickman
(1931:243) studied spermiogenesis in Succinea ovalis and dis-
cussed this organ in some detail. He characterized the hermaphro-
dite gland as being composed of many acini "pressed closely to-
gether, each opening into an atrium which communicates with the
hermaphrodite duct." Hickman (op. cit.) referred to Succinea
ovalis as a "simultaneous hermaphrodite" because male and female
germ cells develop at the same time from an indifferent germinal
epithelium. The color of the hermaphrodite gland is usually
whitish-gray, with the periphery deeply pigmented dark gray. The
gland is invested by the digestive gland on all but the surface to
which the hermaphrodite duct attaches. The surrounding digestive
gland may be teased away rather easily from the hermaphrodite
gland.

The hermaphrodite duct originates on the free surface of the
hermaphrodite gland. The size and shape of this duct varies ac-
cording to the age of the animal. In small, immature specimens,
the duct is thin and often unconvoluted; as the animal increases in
size, the duct becomes thicker and convoluted to a greater degree.
The degree of pigmentation on the duct has been used in the diag-
noses of some species of Succineidae by Pilsbry (1948). Degree
of pigmentation is of no taxonomic value, because most series of all
species considered in the present paper have shown extreme varia-
tion in this character. The hermaphrodite ducts of different indi-
viduals vary from deeply pigmented to light-colored and immacu-
late. It is worth noting that pigmentation of any internal organ
is too variable to be of taxonomic importance. Furthermore, there
is seemingly no correlation between age of the animal and degree
of pigmentation of any structure.

The hermaphrodite duct terminates at the fertilization sac,
situated at the base of the seminal vesicles. This sac sometimes
is and sometimes is not the same color as the seminal vesicles.

The seminal vesicles of all individuals examined are paired; Pils-
bry (op. cit.) reported this to be the prevalent condition among
species of Succineidae. The two structures are occasionally equal
in size, but generally unequal; in a series of any species in Kansas,
the relative sizes of the seminal vesicles varies considerably.

The albumin gland is often the largest organ of the genitalia,
especially in Quickella. The ventral proximal portion of the gland
contains the fertilization sac and seminal vesicles. Albumin for the

1508 The University Science Bulletin

ova is supposedly supplied by this gland, which in Helisoma trivolvis
opens through a duct into the fertilization sac (Abdel-Malek 1954:
114). The albumin gland is usually pale, although it may be pig-
mented to varying degrees.

The term oviduct is here used in preference to uterus, because all
members of the Family Succineidae are oviparous; the term "uterus"
should be reserved for use in animals that are viviparous or ovovivi-
parous.

The size of the oviduct varies with the stage of sexual develop-
ment of the animal. In immature animals the oviduct is smaller in
diameter and more compact; its bulk may be exceeded by that of
the prostate gland. In sexually mature animals the distended ovi-
duct may obscure the penis, vagina, and seminal receptacle. This
distention is especially obvious in large individuals of Succinea
concordialis.

The term seminal receptacle is synonymous with "spermatheca";
both words seem equally applicable. Sperm introduced during
mating are stored in this organ (Abdel-Malek, op. cit.). it is usually
globular, although it may be elongate-oval in Quickella vagans.
The size of the structure varies widely in some series of animals.
The duct of the seminal receptacle originates on the oviduct at
varying positions, according to the species. The duct is long and
slender in Succinea concordialis, and short and thick in S. ovalis.
Intermediate stages are found in other species. The portion of the
female tract below the origin of the duct of the seminal receptacle
is termed the vagina.

A nearly infallible rule seems to be that the seminal receptacle
is associated with the left salivary gland, for in most individuals,
the seminal receptacle or its duct is connected with this salivary
gland by connective tissue. The seminal receptacle and intestine
may be in actual contact, or they may be separated and joined by
connective tissue.

The relative length of the vagina is constant within a species, but
varies among species of the Family Succineidae. In Succinea ovalis
and Oxyloma retusa the vagina is nearly as long as the penis; it is
much shorter than the penis in Quickella vagans, Q. wandae, Suc-
cinea concordialis, S. vaginacontorta, and S. pseudavara. A peculiar
torsion in the vagina is the primary distinguishing character of S.
vaginacontorta ( Lee 1951b ) .

The prostate gland is relatively large and conspicuous in Quickella,
Oxyloma, and Succinea ovalis. In the remaining species of sue-

Family Succineidae (Gastropoda: Pulmonata) 1509

cineids occurring in Kansas, this gland is smaller. Contrary to the
description by Pilsbry (1948:835), in which the prostate gland of
Succinea concordialis is termed large, this gland is relatively small
in this species in Kansas. In young Quickella and Oxyloma, the size
of the prostate gland frequently exceeds that of the oviduct.

The term vas deferens is applied to the sperm duct from the point
at which it leaves the prostate gland until it inserts on the distal end
of the penis; its length and diameter are variable. It is long and
relatively thin in Oxyloma retusa and Succinea ovalis, but shorter
and thicker in S. concordialis. Intermediate conditions prevail in
other species. Usually, the distal portion of the vas deferens is dif-
ferentiated into an epiphallus, which in turn inserts on the penis.
The epiphallus reaches its greatest differentiation in Oxyloma and
is completely contained within the penial sheath in that genus. In
Succinea, the epiphallus may or may not be completely contained
within the sheath. An epiphallus is not distinctly differentiated in
Quickella, although the vas deferens thickens as it enters the penis.
In this study, a short epiphallus is considered to be present in
Quickella beginning at, or near, the insertion of the penial retractor
muscle on the sperm duct. The penis has been used extensively as
a diagnostic feature in species of the Family Succineidae. In two
genera, Oxyloma and Succinea, the penis is surrounded by a sheath,
which varies in thickness and other characters among the various
species. In Quickella, a penial sheath is not present.

The penis of Oxyloma possesses a structure termed an appendix,
a small protuberance situated approximately at the point where
the epiphallus inserts. The size, shape, and position of this ap-
pendix has been given considerable taxonomic weight in diagnosing
species of Oxyloma ( Pilsbry, 1948 ) ; several species have been diag-
nosed almost exclusively upon the appearance of the appendix.
It has become evident to me, however, that the appendix varies
within a population, from a small, blunt structure, scarcely evident,
to one that is longer and tapering. It seems obvious that variations
in the appendix are not specifically diagnostic. All populations of
Oxyloma discovered in Kansas are considered to be Oxyloma retusa.

Quickella is unique among genera of Succineidae in Kansas in
lacking a penial sheath. The penis possesses an appendix that is
usually larger than the penis. Lee (1951a) believed that the terms
penis and appendix are not applicable in Quickella, because he con-
sidered these structures to be elaborations of the vas deferens.
However, in the present discussion, the standard interpretation of
penis and appendix is retained.

1510

The University Science Bulletin

100

39

38

■Li

wMIUl
=)

Map of Kansas sho.7in£ counties in which species
of Succineidae were collected during this study*
I

39

38

100

97

Succinea ova lis M
jccinea concordialis Q

Succinea pce-idavara Q)
Succinea vaginacontorta

CVicfrella varans Q*
Quickella w^ndae @

Oxyloraa retusa A,

Fig. 1. Map of Kansas showing counties in which the seven species of
Succineidae were collected in the course of this study. For each of four
species, each symbol represents more than one record-station of occurrence; for
example, although only one symbol for Quickella vagans is shown in Douglas
County, collections of this species were made from several widely spaced local-
ities within the county. For Succinea pseudavara, each symbol represents one
collection. For Succinea vaginacontorta and for Quickella xvandae, each symbol
represents one specimen.

l.A.

B.

2. A.

B.

3. A.
B.

4. A.

B.
5. A.

B.

Key to the Species of the Family Succineidae in Kansas

Penial sheath not present; large appendix present which equals, or

exceeds, length of penis 2

Penial sheath present. Appendix, if present, smaller than penis ... 3
Appendix approximately equal to, or slightly longer, than penis,

Quickella vagans
Appendix much longer than penis due to presence of outgrowth from
distal end; this outgrowth may be thrust "papilla-like" into appendix,

Quickella wandae

Appendix present on distal end of penis Oxyloma retusa

Appendix not present on penis 4

Vagina having prominent torsion just above entrance to genital

atrium Succinea vaginacontorta

Vagina lacking torsion 5

Vagina nearly as long as penis; duct of seminal receptacle sur-
rounded by loop of oviduct Succinea ovalis

Vagina much shorter than penis; duct of seminal receptacle not sur-
rounded by loop of oviduct 6

Family Succineidae (Gastropoda: Pulmonata) 1511

6. A. Vas deferens flattened, wide, becoming incorporated with sheath of
penis soon after crossing junction of penis and vagina; foot of animal
possesses prominent dark gray flecks over entire surface

Succinea concordialis

B. Vas deferens not flattened, thin; becomes attached to sheath of penis

distally; foot of animal with pigmentation concentrated primarily

anteriorly and on posterior tip Succinea pseudavara

ACCOUNTS OF SPECIES

Succinea ovalis Say
(Plate I B, K, L; figures 2, 10a)

Succinea ovalis Say, 1817, Jour. Acad. Nat. Sci. Philadelphia, vol. 1, p. 15;

DeKay, in part, 1844, Nat. Hist. New York, Mollusca, p. 53, pi. 4, fig. 52;

Pilsbry, 1908, Proc. Acad. Nat. Sci. Philadelphia, p. 45, with var. optima,

p. 46, and var. chittenangoensis, p. 49; Walker, 1928, Terrestrial Mollusca

of Alabama, p. 167; Walker, 1904, Sixth Rep. Michigan Acad. Sci., p. 187;

Ingram, 1943, Nautilus, vol. 56, p. 92; Pilsbry, 1948, Land Mollusca of

North America, vol. 2, p. 801, figs. 429-432.
Helix (Cochlohydra) ovalis Say, Ferussac, 1822, Tabl. Syst. Limagons, p. 26;

Hist. Nat. Moll., pi. 11a, fig. 1.
Succinea ovatis Leidy, 1851, in Terr. Moll., vol. 1, pp. 213, 231, pi. 13, figs.

I-III.
Succinea ohliqua Say, 1824, in Appendix to Keating's Narrative Exped. source

St. Peter's River, etc., Major Long's Second Expedition, vol. 2, p. 260, pi.

15, fig. 7; W. G. Binney, 1S78, Terr. Moll., vol. 5, p. 423, and of most

authors of the last century.
Succinea campestris Say, Gould, 1841, Invertebrates of Massachusetts, p. 195,

fig. 126; DeKay, t. c, p. 53, pi. 4, fig. 54. Not Succinea campestris Say.
Succinea toUcniana Lea, 1841, Trans. Amer. Philos. Soc, vol. 2, p. 32; W. G.

Binney, 1878, Terr. Moll., vol. 5, p. 425, pi. 67b, fig. 2.
Helix {Cochlohydra) putris, Ferussac, Tableau Syst. fam. Limagons, p. 26, no.

9; Hist. Nat. Moll., pi. 11 A, fig. 9.
Succinea putris L., Cockerell, Nautilus, vol. 6, p. 30.

Type locality: Philadelphia, Pennsylvania.

Shell: "oval, inflated, thin, translucent, of a greenish-yellow tint,
the summit paler or reddish; glossy; lightly marked with wrinkles
of growth. Whorls two and a half, strongly convex, the last in-
flated, convex throughout. The aperture is ovate, about three-
fourths the length of shell" (Pilsbry, 1948:802). The shells of Kan-
sas specimens agree closely with this description, excepting the
ratio of length of aperture to the length of shell, which averages
somewhat less than three fourths ( see below ) . The shell is usually
the same color over its entire surface; the "reddish summit" present
in the living state, is due to coloration of the brown digestive gland
in the apex. Because of its distinct appearance at all stages of
growth and its limited distribution in Kansas, the shell is seldom,
if ever, mistaken for that of any other species of succineids in the
State.

16—8050

1512 The University Science Bulletin

Table 1. Measurements of Shells of Succinea ovalis Say

length of diameter of

length diameter aperture aperture

18.8 mm. 10.8 mm. 12.0 mm. 8.6 mm.

11.6 6.9 8.8 6.0

17.4 9.4 11.7 7.4

15.2 8.6 10.4 6.8

9.8 5.7 7.1 4.5

From shells of 47 specimens, collected on 9 April, 1955, near Iowa
Point, Doniphan County, the following average ratios were ob-
tained:

Diameter of shell

Length of shell

Length of aperture
Length of shell

Diameter of aperture
Length of aperture

= 59%, ranging from 55% to 63%
= 70%, ranging from 59% to 75%
68%, ranging from 62% to 76%

Soft anatomy: The external appearance of the living animal varies
considerably, as is true with most species of Succineidae in Kansas.
The ground color of the foot is usually pale yellow; the sole is
always yellow-orange, the periphery possessing darker orange color
than the center. The sole is seemingly never pigmented.

Although some specimens are nearly immaculate, some pigmenta-
tion usually is present. Pigmentation varies from a faint gray
stippling anteriorly on the sides of the head and foot, to dark gray-
vertical bars extending the entire length of the foot. In some ex-
amples, there is whitish stippling on the sides of the foot and under
the collar, giving a glistening appearance to the animal when
viewed under a dissecting microscope. The top of the head is in-
variably pigmented. The majority of specimens possess dark gray
pigmentation evenly distributed over the dorsal surface of the head,
in the area bordered laterally by the tentacular retractor muscles,
which are visible through the epidermis of the snail.

The pattern of markings of the mantle over the lung varies
markedly; the color in this species is ordinarily some shade of
brown.

The jaw of Succinea ovalis is unique among Kansas succineids
in possessing accessory ribs, in addition to the usual median pro-
jection (Fig. 10a).

Genitalia (Fig. 2): The genitalia of Succinea ovalis are distinc-
tive among the species under consideration in several respects. The
base of the duct of the seminal receptacle is as large in diameter,
frequently larger, than the oviduct at its point of origin, but narrows

Family Succineidae (Gastropoda: Pulmonata) 1513

before reaching the globular .seminal receptacle. The oviduct de-
scribes a loop around the duct of the seminal receptacle. This
feature is diagnostic of Succinea ovalis in Kansas. The penis is
large, completely filling the sheath. The vagina is nearly as long
as the penis.

Genitalia are white except that the hermaphrodite duct, prostate
gland, and seminal vesicles are invariably pigmented to some
degree. The prostate gland is large and usually brown. The vas
deferens is long and sinuous and differentiated into a distinctly
thicker epiphallus beyond the point of insertion of the penial re-
tractor muscle. The epiphallus is invariably free from the penial
sheath, the epiphallus usually producing a sigmoid flexure. The
penial retractor muscle is relatively thick and of equal diameter
throughout its length.

The appearance of the free, looped epiphallus varies from speci-
men to specimen, but the loop is always present.

Distribution: Succinea ovalis barely enters northeastern Kansas
in Doniphan County, inhabiting bluffs along the Missouri River.
Numerous specimens have been collected from these bluffs along
Kansas Highway 7, from Iowa Point to White Cloud (Doniphan
County). Baker (1902:219) gives its distribution as eastern and
central parts of northern United States, and west to Manitoba,
Canada, and south to Arkansas and Georgia. This would place
the western limit of its range in Kansas.

In Kansas, Succinea ovalis was generally found in shaded, damp
situations along the base of bluffs, sometimes at considerable dis-
tance from the river. Invariably, the snails were on the ground
under a protective cover of grass, leaves and twigs. A series of ap-
proximately fifty living specimens was collected on 9 April, 1955,
from a low, wooded hill overlooking the Missouri River at White
Cloud. On most other occasions, fewer specimens were found at
any given time, generally six to twelve, but in April, 1956, seventeen
specimens, ranging from immature to adult, were found in a small
ravine filled to a depth of two feet with dead leaves. The snails,
some of which were hibernating, were clinging to leaves.

General remarks: Strandine (1941:86) studied the ecology of a
population of Succinea ovalis on a flood plain in Illinois. He found
that the available leaf-mold seemingly influences population density,
which "increases in the spring and early summer when the available
leaf-mold is greatest." Strandine further found that "fluctuations
in the density of the Succinea ovalis population coincides with the

1514 The University Science Bulletin

fluctuations in the soil moisture, the organic matter, and the pH;
the snail population increasing when the pH is increasing and the
soil moisture and organic matter are decreasing."

Shimek (1935:6) found Succinea ovalis generally distributed
throughout Iowa in wooded areas, especially in low alluvial bottom-
lands. He noted a population twenty-five to seventy-five feet above
the river, entirely above the highest flood level.

Parasites: Ingram and Hewitt (1943:92) reported the occurrence
of the sporocyst of a trematode (Leucochloridiion) in Succinea
ovalis in New York State. The sporocyst is banded with shades of
brown, white, and green. Sporocysts, presumably of Leucochlo-
ridium, have been observed in Kansas in three individuals of S.
ovalis, intertwined among the organs of the reproductive system.

Ellis (1926:140) reported that the sporocyst (Leucocliloridium)
is a stage in the life cycle of the fluke Distomum; the brightly
colored branches in the tentacles of the snail (Succinea putris)
attract the attention of birds, which peck off the tentacles and
infect themselves. According to Ellis, the snail then grows a new
tentacle!

Succinea concordialis Gould
(Place I F, I; figures 3, 10b)

Succinea concordialis Gould, 1848, Proc. Boston Soe. Nat. Hist., vol. 3, p. 38

(Lake Concordia); 1851, in Binney, Terr. Moll., vol. 2, p. 82, pi. 67a, fig. 2;

W. G. Binnev, 1878, Terr. Moll., vol. 5, p. 418; 1885, Manual American

Land Shells, p. 441; Pilsbry and Ferriss, 1906, Proc. Acad. Nat. Sci. Phila.,

p. 159, figs. 11, 12; F. C. Baker, 1939, Fieldbook Illinois Land Snails, p.

123, figs.; Pilsbry, 1948, Land Mollusca North America, vol. 2, p. 833, figs.

452-454.
Succinea munita Binney, 1851, Terr. Moll., vol. 1, p. 128 (nomen nudum),

teste W. G. Binney.
Succinea forshet/i Lea, 1864, Proc. Acad. Nat. Sci. Philadelphia, p. 109; Jour.

Acad. Nat. Sci. Philadelphia, ser. (2), vol. 6, p. 178, pi. 24, fig. 107.
Succinea haleana Lea, 1864, Proc. Acad. Nat. Sci. Philadelphia, p. 109.
Succinea halei Lea, Jour. Acad. Nat. Sci. Philadelphia, ser. (2), vol. 6, p. 180,

pi. 24, fig. 110.
Succinea witteri Shimek, 1913, Nat. Hist. Bull. State Univ. Iowa, vol. 6, p. 31,

pi. 1, figs. I-IV.

Shell: "obliquely ovate, elongate, reflexed, apex acute, thin but
firm, transparent, shining, feebly striated lengthwise and spirally,
color pale honey yellow, with the tip ruddy; whorls three and some-
what more, very oblique . . .; aperture ample, not less than
two-thirds the length of the shell, well-rounded at the base; . . .
a broad, thin callus covers the left margin, which is slightly detached
anteriorly, so as to form the rudiment of an umbilicus" (Pilsbry,
1948: 833).

Family Succineidae (Gastropoda: Pulmonata) 1515

Tahle 2. Measurements of Shells of Succinea concordialis Gould

length of diameter of

length diameter aperture aperture

I 1 .8 nun. 6.5 mm. 8.0 mm. 4.8 mm.

13.4 7.8 9.7 6.0

13.7 7.3 9.6 5.5

12.7 7.2 9.0 5,3

11.0 6.0 8.0 4.6

9.1 4.9 6.1 3.6

In all series of Succinea concordialis examined from Kansas, the
ratio of the length of shell to length of aperture approximates 70
per cent; this same ratio is present in Succinea ovalis. However, the
shells of the two species differ in the average ratio of the diameter
of aperture to length of aperture (60 per cent in S. concordialis, 70
per cent in S. ovalis).

Table 3. Average Ratios Derived from Dimensions of Shells of
S uccinea concordialis

D LA DA

Locality No. shells Date collected L L LA

Lone Star Lake, Douglas Co 13 16 Aug., 1955 .55 .70 .59

Sharon, Barber Co 9 3 Apr., 1956 .57 .65 .61

Lawrence, Douglas Co 8 16 May, 1956 .56 .70 .62

State Lake, Meade Co 8 3 June, 1956 .56 .68 .62

Alma, Wabaunsee Co 12 12 June, 1956 .54 .70 .59

The shells of adult specimens of Succinea concordialis are not
easily confused with other species in Kansas. The shining surface
and ruddy apex is characteristic. However, shells of immature
specimens are often without the ruddy apex and may be confused
with those of Quickella vagans.

Type locality: Lake Concordia, Louisiana.

Soft anatomy: In the living state, the animal is easily identified
by the characteristic black or dark gray, white-spotted, mantle, which
is seen through the transparent shell. Pilsbry (1948:834) states
that the spots are yellow in this species, but all specimens seen by
me from Kansas possessed white spots, which appeared yellow when
viewed through the shell. The foot is light gray, flecked with dark
gray. This pigmentation is more or less evenly distributed over the
exposed portion of the body when the animal is active. The dark
maculations are larger than those on any other species of Succineidae
in Kansas. Pigmentation on the posterior tip of the foot and dor-
sally on the head is arranged in a pattern of vertical bars; this pat-
tern is not exhibited on the sides of the foot. The sole is gray, some-
times tinged with yellow. A few specimens possess faint stippling
on the sole of the foot.

Jaw (Fig. 10b).

Genitalia (Fig. 3): The genitalia of Succinea concordialis vary

1516 The University Science Bulletin

little in populations in the state. The species in Kansas differs from
the description given by Pilsbry ( 1948:835) primarily in one respect:
the prostate gland is always relatively small in comparison to the
oviduct and other organs of the genitalia. Pilsbry ( loc. cit. ) states
that the prostate gland of this species is comparatively large; his
accompanying illustration confirms his statement. It should be
noted, however, that Pilsbry's description is based on rather small
specimens from one locality ( Del Rio, Texas ) . More than one hun-
dred specimens have been examined from widely scattered localities
in Kansas; the shells of these specimens ranged from less than 6 mm.
in length ( immature ) to more than 14 mm. ( adults ) .

The duct of the seminal receptacle is unusually slender in Swo
cinea concordialis and originates low on the oviduct at approximately
the level where the vas deferens crosses the junction of penis and
vagina. The seminal receptacle is globular. The vas deferens is
diagnostically short, wide, and somewhat flattened, becoming at-
tached to the sheath of the penis immediately after crossing the
junction of penis and vagina. The penial sheath is small in rela-
tion to the size of the complete genitalia; this size relationship is
especially noticeable in large snails. The sheath is thin, semi-
transparent, enclosing a relatively thin penis. A distal opening in
the sheath, allowing the epiphallus partially to emerge, has been
seen in a few small specimens, but in the great majority of specimens,
the sheath is entire. The penial retractor muscle inserts on the dis-
tal end of the sheath. Penis and sheath may be dark gray, although
they are usually immaculate.

The hermaphrodite duct is usually some shade of gray, never
brown as in some other species of Succineidae in Kansas. The duct
becomes tightly convoluted soon after leaving the hermaphrodite
gland, then straightens before inserting at the fertilization sac. Ap-
proximately two-thirds of the duct is convoluted.

Distribution: Succinea concordialis seemingly occurs through-
out Kansas (Map, Fig. 1); in numbers it is exceeded only by Quick-
ella vagans. Pilsbry (1948:834) cites the following states in which
this species has been reported: Texas, Louisiana, Arkansas, Mis-
souri, Illinois, Iowa, Tennessee, Alabama, and Florida. Webb
(1954:10) reported this species from Marshall County, Oklahoma.
Franzen and Leonard (1942:339) noted its occurrence in Kingman
County, Kansas.

Succinea concordialis has been found in Kansas in two distinct
habitats. Generally, it occurs near the edges of streams in wet

Family Succineidae (Gastropoda: Pulmonata) 1517

situations, clinging to vegetation near or in water, or living in the
wet leaf mold at the edge of the water.

Elsewhere, the species inhabits comparatively dry environments.
In Meade County State Park, a number were found near the lake,
in a low area overgrown by thick vegetation, but several isolated
places were sandy and barren. The sand was only slightly damp.
S. concordialis was found here, inhabiting the damper places in
the sand. Quickella vagans was also present in approximately equal
numbers. The only shade was provided by a dead bush.

On a few other occasions, Succinea concordialis was found in
drier situations, and the species is observed in the open, exposed
to the sun, more than other species of Succineidae in Kansas.

General Remarks: Succinea concordialis was reluctantly placed
in the section Calcisuccinea by Pilsbry (1948:826) because the geni-
talia resemble superficially those of S. campestris Say, S. luteola
Gould, and S. /. floridana Pilsbry. Webb (1954:10) has recently
removed S. concordialis from Calcisuccinea and placed it in a new
section, Desmosuccinea, characterized by the fusion of the pore in
the distal end of penial sheath, which prevents the epiphallus from
emerging in a loop as it does in species of Calcisuccinea. I have
observed several immature specimens of S. concordialis that pos-
sessed a pore which allowed the epiphallus to emerge in a slight
loop. Nevertheless, Webb's action of placing S. concordialis in a
new section seems justified, because the pore in the penial sheath
is seemingly always obliterated in the adult animal.

Parasites: One specimen of S. concordialis from Lone Star Lake,
Douglas County, was infested by a fly larva, seemingly identical
with those in several specimens of Quickella vagans.

Succinea vaginacontorta Lee

(Plate I C; figure 4)

Succinea vaginacontorta Lee, 1951, Occas. Pap. Mus. Zool.; Univ. Michigan, no.
533, p. 1, pis. 2, figs. 7.

"Shell is dextral, fully developed. Whorls 3, with suture only
moderately impressed. Whorls which increase proportionately from
the apex to the body whorl with no marked discontinuity in size.
In life shell is a dull, translucent, light, horn yellow with greenish
tones. Striae present and well raised. Epidermis of upper whorl
eroded away revealing dull white deeper layers of shell, parietal
wall with well developed callus, covering base of slightly curved
columella. Aperture obliquely ovate with a slight flare at base of

1518 The University Science Bulletin

outer lip. Interior of aperture with a thin shiny transparent sheen"
(Lee, 1951b: 2, description of holotype).

Shell of holotype: Length, 9.8 mm; width, 5.9 mm; aperture
length, 5.7 mm; aperture width, 4.2 mm.

Type locality: Southwest corner of Sec. 18, T. 33S, R. 28W, ap-
proximately 8 miles south and 4 miles west of Meade, Meade County,
Kansas.

I found only one specimen of this species; it was at Meade County
State Park, 3 June, 1956, on a bank of an impoundment pond near
the lake. The specimen was associated with numerous individuals
of Quickella vagans. The animal remained unrecognized until dis-
sected. Measurements of its shell are: length, 8.4 mm; diameter,
4.9 mm; length of aperture, 5.2 mm; diameter of aperture, 3.6 mm.

This shell generally fits the original description; the callus is not
well developed, but the striae are quite pronounced.

Externally, "the animal is a dull, grayish white with the anterior
part finely peppered with black pigment . . . the surface of this
hermaphroditic animal is sufficiently transparent in the region of the
genital furrow to expose the outlines of both the vagina and penis
. . . the edge of the mantle may have a few to many concentra-
tions of black pigment. Some appear gray. The rest of the mantle
is finely re'iculated" (Lee, 1951b:3).

Genitalia (Fig. 4) : Succinea vaginacontorta is distinguished from
all other known species of the Succineidae by the presence of a
prominent torsion in the vagina, occurring slightly posterior to the
junction of the vagina with the common genital atrium.

In the single specimen examined, the hermaphrodite duct is light
gray. The unequal seminal vesicles are somewhat darker. The
albumin gland, oviduct, prostate gland, and vagina are white. The
penial sheath is relatively larger than in any other succineid exam-
ined and the epiphallus emerges from the penial sheath in a flattened
loop. Distally, the penial sheath and vas deferens are peppered with
gray. The prostate gland is small, and the vas deferens thick. The
latter structure approaches the junction of penis and vagina and then
rises to run parallel to the penial sheath until the vas deferens enters
the sheath distally. The penial retractor muscle is thin and inserts
on the penial sheath at the base of the free loop of the epiphallus.
The seminal receptacle is small and globular, the duct originating
immediately above the twisted portion of the vagina. The duct is
long and slender.

The penial sheath is thick and resistant to cutting, more so than
in any other species of the family known to occur in Kansas. As

Family Succineidae (Gastropoda: Pulmonata) 1519

Lee (1951b:5) indicated in his original description, the penis is
enclosed in a sheath "of tough, fibrous, fuzzy, connective tissue".
In other species of Succinea examined, the sheath is thin and usually
semi-transparent. The penis is relatively thin in S. vaginacontorta.

General remarks: On the basis of the genitalia, Lee considered
that the affinities of Succinea vaginacontorta were with the section
Calcisuccinea Pilsbry, which includes S. campestris Say, S. luteola
Gould, and S. /. fioridana Pilsbry.

Webb (1953:216) has placed S. vaginacontorta in a separate sec-
tion, Heysuccinea. He considered the distinctive torsion in the
vagina, and the vas deferens being free from the junction of penis
and vagina, sufficient justification to warrant this.

Distribution: Succinea vaginacontorta has been reported to occur
only in Kansas. Webb ( op. cit. ) found the species at the following
localities: Lawrence, Douglas County; south of Kingman, Kingman
County; 6 miles north of Meade, Meade County; 3-4 miles north of
Herington, Dickinson County.

Succinea pseudavara Webb
(Plate ID; figure 5)
Succinea pseudavara Webb, 1954, Gastropodia, vol. 1, no. 2, p. 10, figs. 8.

Shell: Amber-colored, rather polished, frequently coated with
mud as in Quickella vagans; these two species may easily be con-
fused. Transverse striae are less regular and riblike than in Q.
vagans. The spire is approximately as long as the aperture. The
apex is pink-brown, darker than the remaining whorls. The shell
lacks the filelike papillae of Q. vagans and the revolving striae of
S. concordialis.

Identification of S. pseudavara probably cannot be made from
the shell alone, as Webb pointed out in his original description. He
did not include measurements of shells. Several shells of this
species from the bank of the Arkansas River in the City Park at
Cimarron, Gray County, Kansas, measured as follows:

Table 4. Measurements of Shells of Succinea pseudavara Webb

length of
length diameter aperture

7.2 mm. 4.3 mm. 5.0 mm.

8.1 4.7 5.4

8.2 4.9 5.5
7.0 4.0 4.5

Table 5. Average Ratios Derived from Dimensions of Shells
of Succinea pseudavara

Locality No. shells Date collected

Cimarron, Gray Co. 8 3 lune, 1956

Medicine Lodge, Barber Co. . . . 7 4 June, 1956

diamt

aper

■ter of
ture

2.9 mm.
3.3
3.2
2.6

is of Shells

D
L

LA
L

DA
LA

.59
.53

.66
.62

.59

.57

1520 The University Science Bulletin

Because specimens of S. pseudavara were found at only four
localities, two of which yielded four snails, no generalities can be
made from ratios derived from measurements.

Type locality: "About 1% miles east of the University of Okla-
homa Biological Station, at Lake Texoma, near Willis, Marshall
County, Oklahoma" (Webb, 1954:18).

External appearance of animal: The color of the foot is grayish-
white and lacks the brownish tint usually present in Quickella
vagans, with which it may be confused. A small amount of gray
stippling is present anteriorly on sides of foot and on top of head.
The posterior tip of the animal also is obviously pigmented, in a
vague pattern of vertical bars. The sole of the foot was pigmented
in only two individuals. The pigmentation on the head and foot
is much finer than the flecks present on Saccinea concordialis, and
is concentrated on the anterior and posterior parts of the animal.
In S. concordialis, the pigmentation tends to be evenly distributed.

The pattern of the mantle resembles that of Quickella vagans.
The ground color is usually translucent gray with darker gray
present in the form of a reticulum. This darker gray is sometimes
arranged in peripheral blotches, simulating the usual condition
found in Quickella. A brownish tint on the mantle has been ob-
served in a few specimens. The rim of the mantle is white. The
kidney usually is not obvious when viewed through the shell.

The eye-stalks are lightly pigmented and there is a narrow
stripe of dark gray extending medially between their bases. The
dorsal portion of the head and foot, when extended, is evenly
pigmented gray on the surface bordered by the dark colored
tentacular retractor muscles.

Genitalia (Fig. 5): Succinea pseudavara is obviously related to
Succinea concordialis in details of its genitalia. The penial sheath
of the former is large and ovate, enclosing a penis similar in ap-
pearance to that of S. concordialis. The penis is relatively thick
distally, tapers as it approaches the genital atrium and increases
abruptly in diameter before reaching the atrium. Most specimens
of S. pseudavara examined by me agreed with Webb's description
(1954:18) in lacking a pore on the distal end of penial sheath, a
loop of the epiphallus, therefore, not appearing. In two specimens,
however, a pore was present and the epiphallus emerged in a
loop. Approximately thirty specimens were dissected.

The seminal receptacle is globular; its duct is long and slender.
The duct originates low on the oviduct; the vagina is therefore
short. The prostate gland is similar in appearance to that of S.

Family Succineidae (Gastropoda: Pulmonata) 1521

concordidis, but is relatively larger. The vas deferens is somewhat
thinner than in S. concordidis and is not flattened, which is one of
the best methods of distinguishing the genitalia of the two species.
The vas deferens reaches the junction of penis and vagina, then
rises to extend along the penial sheath before becoming incor-
porated distally in the sheath and penial retractor muscle. The
penial retractor muscle is thin. The seminal vesicles are usually
darker than the hermaphrodite duct and were relatively small in
the specimens examined.

The entire reproductive system is colored varying shades of gray;
the brownish color frequently seen in species of Quickella has not
been observed in S. pseudavara.

Distribution: This species previously has been reported only
from Oklahoma ( Webb : 1954 ) . In die course of my study, speci-
mens were collected from four counties in Kansas: Osborne, Gray,
Barber, and Sumner.

Succinea pseudavara was found along the edges of streams in Kan-
sas, associated with Quickella vagans and S. concordialis. The shade
provided by bridges seems to attract this species, although speci-
mens were also collected in more exposed places, under the con-
cealment afforded by grass and other vegetation. The shell may be
coated with mud.

General remarks: I agree with Webb (op. cit.) that S. pseudavara
is related to S. concordialis. The genitalia of the two species are
similar in several significant respects : ( 1 ) the penes of both species
are morphologically similar; ( 2 ) the duct of the seminal receptacle
originates at approximately the same position on the oviduct in both
species; (3) the epiphallus does not ordinarily appear in a free loop
from the sheath of the penis; (4) the size and insertion of the penial
retractor muscle are alike in both species; and (5) the vas deferens
reaches the base of the junction of the penis and vagina in both
species.

Webb placed S. pseudavara and S. concordialis in a new section,
Desniosuccinea. The striking similarities in the genitalia of the two
species certainly warrants this action.

Oxyloma retusa (Lea)
(Plate I G, H; figures 6, 7)

Succinea retusa Lea, 1834, Trans. Amer. Philos. Soc., vol. 5, p. 117, pi. 19, fig.

86; (Ohio, near Cincinnati).
Succinea higginsi "Bland, Nov. spec.", Tryon, July, 1866, Amer. Jour. Conch.,

vol. 2, p. 237, pi. 17 (2), fig. 24; Bland, October, 1866, same volume,

p. 373, pi. 17, fig. 24; W. G. Binney, 1885, Manual American Land Shells,

p. 198, fig. 206.

1522 The University Science Bulletin

Succinea retusa magister Pilsbry, 1899, Nautilus, vol. 12, p. 103; 19:109, fig. 2;

Walker, 1906, Mollusca of Michigan, p. 502, fig. 110; F. C. Baker,

1939, Fieldbook Illinois Land Snails, p. 125, fig. B.
Succinea calumetensis Calkins, 1878, Valley Naturalist, vol. 1, no. 11, p. 57,

text fig.; cf. Walker, 1906, Mollusca of Michigan, p. 502, fig. 108.
Oxyloma retusa (Lea), Pilsbrv, 1948, Land Mollusca of North America, vol. 2,

p. 785, figs. 421, 422C.

Shell: "ovately oblong, very thin, pellucid; spire short; whorls 3;
aperture below dilate and drawn back. Diam. 0.3, length 0.7 of an
inch. ... It differs so much from any of the described species
in the dilation and retraction of the inferior part of the aperture that
I have not hesitated to consider it new" (Lea's description, quoted
by Pilsbry, 1948:786).

The shell of this species is an intense yellow-horn color. Whorls
vary from 2 to 3, depending on the age of the animal. The suture
between body whorl and apex is not prominent. The whorls are
less convex than in any other known species of the Succineidae in
Kansas. The aperture is elongate, occupying 70 percent of the total
length of the shell. In some populations, individual shells vary in
greatest diameter to a marked extent. Lee (1951a) has pointed out
that the parietal area and outer margin of the peristome tends to be
straight, producing in outline an aperture resembling an elongate
teardrop.

Type locality: Ohio, near Cincinnati.

Table 6. Measurements of Shells of Oxyloma retusa ( Lea )

length of diameter of

Length diameter aperture aperture

12.6 mm. 6.8 mm. 9.2 mm. 5.6 mm.

11.7 6.5 8.5 5.4
10.3 5.9 7.3 4.7
14.1 7.0 10.1 6.2

9.0 5.0 6.8 4.2

Table 7. Average Ratios Derived from Dimensions of Shells
of Oxyloma retusa

D LA DA

Locality No. shells Date collected L L LA

Muscotah, Atchison Co 16 5 Apr., 1955 .55 .73 .64

Iowa Point, Doniphan Co 10 4 Sept., 1955 .47 .68 .60

Iowa Point, Doniphan Co 12 21 Apr., 1956 .54 .70 .60

The diameter of the shell is approximately 50 percent of the
length of shell in Oxyloma retusa. A few series were collected
in which the diameter of shells averaged less than 50 percent of
the length.

External appearance: Baker (1939:125) reported that Oxyloma
retusa appears in two color phases: dark in spring, and light in
summer and autumn. Both color phases occur in Kansas. In the
light phase, the foot of the animal is light gray, or yellow, with

Family Succineidae (Gastropoda: Pulmonata) 1523

clumps of fine stippling present laterally and dorsally on the head
and foot. Pigmentation is more profuse anteriorly, in some speci-
mens disappearing toward the posterior half of foot. The sole of
the foot may be pigmented. In the darker color phase, considerably
more pigment is present on the head, mantle, and foot. The
lateral and anterior pigmentation on the foot of a few specimens
may be arranged in vertical rows similar to the pattern exhibited
in Quickella. The underside of the rim of the mantle is usually
pigmented, the pigment being concentrated in the vicinity of the
opening to the lung.

The mantle over the lung is stippled with fine dark gray dots,
present on a gray or tan background. The stippling is uniformly
distributed anteriorly on the mantle, but becomes dispersed pos-
teriorly, often appearing in isolated groups on the posterior portion
of the animal. Sparse stippling is often present on the rim of the
mantle. As in Quickella, the pigmentation on the mantle may be
concentrated peripherally into blotches, from which faint streaks
radiate toward the kidney.

Genitalia (Figs: 6, 7): Oxyloma retusa is characterized by the
presence of a small appendix on the penis, at about the position
where the epiphallus inserts (Fig. 7). The appendix is extremely
variable in size and shape within the species; this variability also
occurs within a population. In large series of animals the appendix
may vary from a blunt, scarcely developed protuberance, to a
slender, tapering structure. The size of the appendix seemingly
is not correlated with the size of the animal. The epiphallus is
completely contained within the penial sheath and may be coiled
or relatively straight. The penis is large and may be straight
within the sheath, but frequently is bent or coiled in a half turn.
When the sheath is opened, the position of the appendix depends
upon the degree of coiling of penis and epiphallus (Fig. 7).

The seminal receptacle is globular; its duct is short and of
approximately the same diameter throughout its length. The duct
originates high on the oviduct, resulting in a vagina the length of
which is equal to that of the penis. The prostate gland is large
and usually speckled with gray. The vas deferens is long, reach-
ing to the junction of penis and vagina, and then rising to insert
on the distal end of the epiphallus after entering the penial sheath.
The albumin gland is relatively small in comparison with oviduct
and prostate gland. As in all species of Succineidae in Kansas, the
degree of pigmentation of the genitalia varies from animal to
animal.

1524 The University Science Bulletin

Distribution: Oxyloma retusa was collected only in Atchison
and Doniphan counties in northeast Kansas. In Atchison County,
a large population exists in a marsh maintained by artesian flow,
1/2 miles south of Muscotah. Although the snails were abundant
there in 1955, the population has since declined.

In Doniphan County, specimens were taken in considerable
numbers throughout the course of this investigation. A favorable
habitat for Oxyloma retusa is roadside ditches along Kansas High-
way 7, in the vicinity of Iowa Point. Succinea ovalis inhabits the
bluffs above these ditches. O. retusa may be seen on vegetation
several feet from the ground. Specimens were taken also from
dead vegetation on the ground and from the wet banks near water.

Elsewhere in North America, O. retusa has been reported from
Ohio, Illinois, Iowa, Minnesota, North Dakota, and Montana
(Pilsbry 1948:786).

General remarks: Alice E. Leonard (1943:240) reported Suc-
cinea haydeni Binney (=Oxyloma haydeni) at Meade County
State Park and noted that "this snail thrives on the moist marshes
and borders of the pools in the artesian basin, where it is frequently
found on watercress." A series of shells in the University of Kansas
Museum of Natural History, collected by her from Meade County,
are large, ranging from 15 mm. to 17 mm. in length. The shells
generally agree in characters with those determined to be Oxy-
loma retusa from Atchison and Doniphan counties. The average
ratios of the shells from Meade County are given below, along with
average ratios obtained from a series of O. retusa from Doniphan
and Atchison counties.

Table 8. Table of Average Ratios of Oxyloma

Locality No. shells Species

Meade County State Park .... 7 Oxyloma haydeni?

Muscotah, Atchison Co 16 Oxyloma retusa

Iowa Point, Doniphan Co 10 Oxyloma retusa

In June, 1956, I visited Meade County State Park and searched
in vain for specimens of Oxyloma. A resident of Meade County
informed me that several years ago, probably in 1953, considerable
grading was done in the artesian marshes by the use of bulldozers.
Perhaps the population of this species was destroyed in the process;
at any rate, neither shells nor live snails were discovered in June
of 1956. Because of the appearance of the shells mentioned above,
however, I believe that the species of Oxyloma from Meade County

D

LA

DA

L

L

LA

49

.72

.59

55

.73

.64

50

.70

.59

Family Succineidae (Gastropoda: Pulmonata) 1525

is 0. rctusa, although final judgment concerning the identification
should be reserved until the animal is found and examined ana-
tomically.

Qiiickclla vagans (Pilsbry)

(Plate I E, J; figure 8)

Sucdnea campcstris vagans Pilsbry, 1900, Nautilus, vol. 14, p. 74.

Quickella vagans (Pilsbrv), 1948, Land Mollusca of North America, vol. 2,

p. 843, fig. 456.

Shell: "Globosely oval shape, very convex whorls and rather dull
or weakly glossy surface, which is irregularly and often in places
somewhat coarsely wrinkled. It is small, fragile, with only %'i to
2% whorls in the type lot, in southern specimens as many as three
whorls; subtranslucent ecru-olive, without the opaque whitish
streaks or the whitish inner layer of campestris" (Pilsbry, 1948:843).

The shell of this species varies strikingly within Kansas. The
number of whorls ranges from 2/2, in small individuals, to nearly
4. The color varies from light yellow to reddish brown. In some
populations, the whorls tend to become flattened, giving the shell
an elongate appearance. Striae are not well developed, although
mature shells are never so glossy as those of Succinea concordialis.
The shells of living snails are frequently coated with mud and
debris.

The shells of young specimens may be confused with those of
young S. concordialis.

Table 9. Measurements of Shells of Quickella vagans (Pilsbry)

length of diameter of

length diameter aperture aperture

12.6 mm. 6.3 mm. 7.3 mm. 4.6 mm.

11.5 6.2 7.1 4.7

10.0 5.8 6.6 4.4

9.0 5.0 5.4 3.3

6.7 4.2 4.3 3.0

Table 10. Average Ratios Derived from Dimensions of Shells
of Quickella vagans

D LA DA

Locality No. shells Date collected L L LA

Lone Star Lake, Douglas Co 16 28 July, 1955 .56 .61 .66

Lawrence, Douglas Co 30 4 Aug., 1955 .55 .64 .63

Sterling, Rice Co 6 1 Apr., 1956 .61 .62 .70

Sharon, Barber Co 6 3 Apr., 1956 .57 .60 .67

State Park, Miami Co 10 12 May, 1956 .57 .57 .71

Burlingame, Osage Co 10 12 June, 1956 .53 .61 .65

Type locality: Cape May Point, New Jersey.

External appearance of animal: Quickella vagans exists in 2 color
phases, light and dark. In the dark phase, there is a characteristic

1526 The University Science Bulletin

pattern of vertical bars completely around the foot. These bars
always are interrupted ventrally, then continue a short distance to
the sole. Anteriorly, four or five bars of pigment are present on the
head, extending from the base of the eyestalks ventrally to near the
mouth. These anterior bars tend to slant medially as they approach
the mouth.

In the light phase, the pigmentation is scanty, or absent. If traces
of the vertical bars persist, they invariably are on the anterior part
of the animal.

The ground color is variable. It may be cream white or gray,
or brownish-orange. There is nearly always a tinge of brown pres-
ent on the animal, either laterally on the foot and/or dorsally on the
head. The color of the sole of the foot is similar to the remainder of
the foot, usually with varying degrees of brown. In some specimens
the sole is stippled, but ordinarily it is immaculate.

There is seemingly no correlation between size of the animal and
the color phase; both phases and intermediates usually are repre-
sented within a population.

The mantle of QuickeUa vagans also varies considerably although
some modification of a basic pattern is present in most specimens.
This pattern consists of streaks of pigmentation radiating inward
toward the kidney from the margin of the mantle. Peripherally, the
pigmentation is concentrated in blotches, from which the radiating
streaks originate. The color varies from gray to brown, and in some
specimens may be so intense as to obscure the pattern. The kidney
usually is bright yellow or orange, easily visible through the mantle,
but this organ may be nearly invisible because of intense pigmenta-
tion on the mantle. The ground color of mantle over the lung is
usually translucent gray, but may be brown. The color of mantle
and foot are independent of one another; a specimen with a lightly
pigmented foot may possess a dark mantle.

Genitalia (Fig. 8) : The distinguishing characteristic of Q. vagans
is the penis-appendix complex. The two structures are variable in
their relative sizes, because the appendix is capable of infolding.
The size of the appendix may be approximately equal to that of the
penis, but is usually somewhat larger. There is no sheath investing
the penis. Longitudinal ridges occur within the penis and appendix.
From the distal end of the penis the epiphallus rises slightly for in-
sertion of the penial retractor muscle, then descends, as the vas
deferens, along the posterior surface of penis-appendix to the junc-
tion of penis and vagina. The penial retractor muscle inserts on the

Family Succineidae (Gastropoda: Pulmonata) 1527

epiphallus and sends a primary branch to the appendix; the penis
proper is free of the retractor muscle, except for minor connectives.

The duct of the seminal receptacle originates low on the oviduct,
somewhat as in Succinea concordialis and S. pseudavara. The length
of this duct varies and is relatively much thicker than in the two
species mentioned above. The seminal receptacle is sometimes
globular, but is often elongate-oval. The vagina is short.

Distribution: Quickella vagans is generally distributed over the
State and is by far the most abundant representative of the Suc-
cineidae in Kansas. Pilsbry (1948:844) records this species from
New Jersey and North Carolina. Kagan (1949:38) was the first to
discover that the genus Quickella occurs as far west as Michigan.
Lee ( 1951a) later identified the species in Michigan. Webb ( 1953)
reported it in Kansas. This species is undoubtedly distributed over
much of North America, but has been, and is, confused with species
of Succinea.

Q. vagans inhabits a variety of habitats. The majority of speci-
mens collected by me were found near margins of streams and lakes.
In such places, individuals generally occur on the ground under a
protec'ive cover of grass, leaves, and twigs, but some were above the
ground attached to blades of grass and vegetation. An examination
of the undersides of smooth logs and sticks lying on banks of streams
often reveal specimens.

This species inhabits also dryer localities, sometimes at con-
siderable distances from standing water. Specimens have been
collected on the banks of dry streams in shady, deciduous woods.
In June, 1956, large individuals were numerous in low, sandy,
areas, barren of vegetation, in Meade County State Park of western
Kansas. The sand was damp beneath small dead branches, scat-
tered over the sand. Several large specimens of Succinea con-
cordialis were associating with the QuickeUas.

At Atwood Lake in Rawlins County, of northwestern Kansas,
several examples of Q. vagans and S. concordialis were found in
the dry bed of a stream draining into the lake. The snails were
under logs and boards on the bed of the stream, shaded by trees.
There was no standing water in sight; Atwood Lake was com-
pletely dry at the time.

In Johnson County, of eastern Kansas, a large population lived
in a heavily wooded area, along the base of a small hill, under
grass and brush, approximately lA mile north of De Soto. This
locality is more than one hundred yards from the Kansas River.

Q. vagans invariably lives beneath logs or other vegetation and

1528 The University Science Bulletin

has not been observed moving in places exposed to the rays of the
sun; whereas, S. concordialis often is active in hot, dry places
exposed to direct rays of the sun.

General remarks: As mentioned earlier, S. avara and S. gros-
venori previously have been considered to be abundant in Kansas.
However, collections studied by me from many localities in Kansas
have failed to reveal either species, as well as other species of
Succinea previously reported in this state. It seems that the vari-
able shell of Quickella vagans, and perhaps that of Q. wandae,
have been confused with species of Succinea. Additional collecting
in Kansas may, of course, reveal the presence of additional species
of Succinea, but I surmise that most shells previously regarded as
S. avara and S. grosvenori were, in fact, Quickella. This statement
is qualified because the distribution and abundance of Succinea
vaginacontorta and S. pseudavara are imperfectly known; both
species resemble Quickella and associate with it.

Parasites: Two kinds of parasites have been found to infest Q.
vagans in Kansas. The least abundant of these parasites is a fly
larva observed in five snails, four of which were collected at Lone
Star Lake in Douglas County. Another infected snail was found
near Coffeyville, Montgomery County. The larva lies partially
against the reproductive system. The genitalia of some infected
snails were relatively underdeveloped, when compared with unin-
fected snails of approximately the same size and from the same
population. The external appearance of the snail is not altered by
the presence of the parasite.

Dr. H. B. Hungerford of the Department of Entomology, Univer-
sity of Kansas, examined a larva and reported the larva probably
to be a first instar of a muscoidean fly, species undetermined.

Dr. Clifford Berg, associated with the Cornell University Agri-
cultural Experiment Station, and a specialist on fly larvae parasitic
in snails, has reported the occurrence of fly larvae (Dyctya ex-
pansa) in Oxyloma decampi gouldi in Alaska (1953:631).

The second parasite found to infest Q. vagans is a small nematode.
There are invariably many individuals of this parasite in an infected
snail; the incidence of infection in Q. vagans is high. In one popu-
lation from Miami County State Park, more than 50 percent of
the snails were infected.

In Michigan, Kagan (1949:38) discovered that Quickella was
infested by sporocysts of the trematode, Leucochloridiiin}, but in
my studies of several hundred Q. vagans from Kansas, no Leuco-
chloridium was found.

Family Succineidae (Gastropoda: Pulmonata) 1529
Quickella wandae Webb

(Plate I A; figure 9)

Quickella wandae Webb, 1953, Jour. Tennessee Acad. Sci., vol. 28, p. 216, pis.
3, 4.

Shell: "very similar to that of Quickella vogons (Pilsbry), green-
ish yellow, epidermis brownish where thick, apex less obtuse than
in Quickella uagans, size less. Umbilicus closed on the lower half
of the columella, rarely partly unsealed" (Webb, 1953:216).

I found only one specimen of Q. wandae. It was obtained along
with several specimens of Q. vagans on 8 April, 1956, from the bank
of a small stream in Franklin County, Kansas.

The one shell examined by me possesses 2% whorls. The color is
greenish yellow. Striae are well developed on the body whorl, but
are scarcely evident on the apex. A parietal callus is present, but is
not prominent.

Webb did not include measurements of shells with his original
description of this species. The one shell examined by me measured:
length, 5.4 mm; diameter, 3.5 mm; length of aperture, 3.4 mm; diam-
eter of aperture, 2.4 mm.

Type locality: 8/2 miles south and )'i mile east of Alma, Wabaunsee
County, Kansas.

External appearance of animal: Webb's original description,
which referred to a drowned specimen, is as follows: "Rim of the
mantle at aperture and exertile body flecked with brown; liver very
light brown; grooves of the foot whitish; regions between the grooves
blackish, fading posteriorly on the foot. Body anterior to the vis-
ceral pillar grayish, marked dorso-laterally on each side by an un-
pigmented line. Sole whitish, densely orange-flecked. Mantle of
the lung whitish with about nine interrupted streaks of dash-like
markings" (Webb, 1953:216).

In the one specimen examined by me, the lateral vertical bars
are well developed around the entire foot. They are interrupted
ventrally, as in Quickella vagans. The mantle pattern differs from
the typical pattern of Q. vagans; instead, the dashlike markings
mentioned in the description by Webb are present. The ground
color of the mantle over the lung is light translucent gray.

Genitalia (Fig. 9): The diagnostic feature which serves to dis-
tinguish the two known species of Quickella in Kansas, is a peculiar
white, smooth, extension on the appendix in Q. wandae, which
lengthens this organ considerably more than the penis. Webb ( loc.
cit. ) referred to this extension as a "glandular outgrowth" on the ap-
pendix.

1530 The University Science Bulletin

The relationship of penial retractor muscle to the penis-appendix
complex may offer an additional diagnostic character. In Q. vagans,
the muscle inserts on the epiphallus, sending a primary branch to
the appendix; the penis is nearly free of any contact with the muscle.
In the one specimen of Q. wondae examined, the muscle inserts on
the epiphallus and the primary branch attaches to the penis near the
junction of penis and appendix; the appendix is free of the muscle.
Whether or not this character is specific for Q. wandae is unknown,
because of the lack of material.

The remainder of the genitalia of the one animal observed does
not differ significantly from the same organs in Quickella vagans.
The hermaphrodite duct is gold-colored, not pigmented. The sem-
inal vesicles are of unequal size and lightly flecked with brown. The
seminal receptacle and duct appear as they do in Q. vagans.

Distribution: Q. wandae has been reported only from Kansas.
Webb collected specimens in four counties: Douglas, Leavenworth,
Wabaunsee, and Anderson. The one specimen collected by me adds
Franklin County to the known range. All specimens collected by
Webb were found in rather shady, deciduous woods on well-drained,
damp terraces.

General remarks: It seems that the shells of Q. wandae and Q.
vagans are indistinguishable.

Webb (1953:216) states that the outgrowth or extension of the
appendix of Q. wandae may be thrust "papilla-like" into the lumen
of the appendix.

ORIGIN OF SPECIES OF SUCCINEIDAE IN KANSAS

The distribution of most of the Recent species of Succineidae
known to live in Kansas now is too imperfectly known to ascertain
their probable origin. Succinea concordialis has been reported
from Texas, Louisiana, Arkansas, Missouri, Illinois, Kansas, Iowa,
Tennessee, Oklahoma, Alabama, and Florida, but the species may
range farther west than is presently known; I collected numerous
specimens only 12 miles or so east of the Colorado line (banks of
the Republican River, St. Francis, Cheyenne County, Kansas).

Oxyloma retusa, which has been reported to inhabit the north-
central and northeastern United States (Ohio, Illinois, Minnesota,
North Dakota, and Kansas), is in a genus the taxonomic status
of which is in a confused state. In Kansas, the shells of O. retusa
vary between different populations and probably this is true else-
where. The distribution of this species probably is incompletely
known.

Family Succineidae (Gastropoda: Pulmonata) 1531

Quickella Dagans has been recorded from New Jersey, North
Carolina, Michigan, and Kansas. Probably it occurs more widely
and farther west than the mentioned occurrences indicate; several
alleged species of Succinea from the western United States may
belong to the genus Quickella.

S. Daginacontorta and Q. wandae are known only from Kansas,
but probably occur elsewhere. S. pseudavara is known only from
Kansas and Oklahoma; it also probably is more widely distributed
than is known at present.

S. avails is the only species of the Succineidae for which the
distribution in North America is well known. This species ranges
throughout the northeastern two-thirds of that continent, and enters
Kansas only in the extreme northeast part of Doniphan County,
which is the northeasternmost county and at about the western
limit of the range of the species. Succinea ovalis has entered Kan-
sas probably from a northern or eastern route of dispersal.

SUMMARY

1. The Family Succineidae is represented in Kansas by three
genera and seven known species. Shells of most species lack dis-
tinguishing characters and cannot be relied upon for purposes of
identification.

2. The genitalia of these seven species are distinctive and fur-
nish the most reliable means of distinguishing one species from
another.

3. The four species of Succinea have in common a penis that
lacks an appendix and that is contained within a sheath. Suc-
cinea ovalis is easily diagnosed because the duct of the seminal
receptacle is surrounded by a loop of the oviduct. S. vaginacontorta
possesses a large, thick penial sheath and is unique in that the
vagina is twisted to a striking degree. The genitalia of S. con-
cordialis and S. pseudavara are similar, but may be separated by
the wide, flattened vas deferens possessed by the former species.

4. Oxyloma retusa is the only known species of Succineidae in
Kansas that possesses a small appendix on the distal end of the
penis and an epiphallus completely enclosed by the penial sheath.

5. The two species of Quickella lack a penial sheath and possess
a large appendix which equals, or exceeds, the size of the penis.
Quickella wandae is distinguished from Q. vagans by the long out-
growth from the distal end of the appendix that is present in the
former species.

1532 The University Science Bulletin

6. S. pseud avara is reported for the first time to occur in Kansas.
It was found in four counties within the state. Additional collect-
ing will be necessary to determine the abundance and distribution
of this species.

7. S. vaginacontorta was collected only in Meade County, where
it lived in association with Q. vagans. The former species has been
reported in eastern Kansas.

8. S. ovalis barely enters the northeast corner of Kansas, occur-
ring in Doniphan County.

9. S. concordialis is exceeded in abundance of individuals only
by Q. vagans. The first species is distinctive in the living state by
the intensely pigmented mantle and foot.

10. Oxyloma retusa occurs in Atchison and Doniphan counties,
in northeastern Kansas. An Oxyloma has been reported from
Meade County, but my search there failed to uncover it. Com-
parison of shells of Oxyloma from Meade, Atchison, and Doniphan
counties suggests that O. retusa is the only species of Oxyloma
occurring in Kansas.

11. S. avara, S. grosvenori, and S. stretchiana have been reported
from Kansas by other workers, but I have not found these species
and suspect that the specimens so identified were Q. vagans, or
some other one of the species reported in the present account.

12. Q. wandae is known to occur in various counties in eastern
Kansas. Only one specimen was collected in my study, from
Franklin County.

13. The Family Succineidae seems adaptable to various ecological
conditions. Several of the species have been found to inhabit wet
situations and also dryer habitats.

14. Three kinds of parasites have been observed to infest the
Succineidae in Kansas: a stage in the life cycle of a trematode
(presumably Leucochloridium) in Succinea ovalis; a fly larva, ap-
parently the first instar stage, in S. concordialis and in Quickclla
vagans; and a nematode, often infesting a large percentage of
specimens from a single population, in Q. vagans.

LITERATURE CITED
Abdel-Malek, E. T.

1954. Morphological studies on the family Planorbidae ( Mollusca:Pul-
rnonata). 1. Genital organs of Helisoma trivolvis (Say). Trans.
Amer. Micros. Soc, vol. 73, pp. 103-124, 4 pis.
Baker, F. C.

1902. The Mollusca of the Chicago area. Bull. Ill, Chicago Acad. Sci.,
Nat. Hist. Survey. (Pt. 2, The Gastropoda): 137-410, 9 pis., 125
text-figs.

Family Succineidae (Gastropoda: Pulmonata) 1533

1909. Mollusks from Kansas and Oklahoma. Nautilus, vol. 23, pp. 91-94.
1939. Fieldbook of Illinois land snails. Illinois Nat. Hist. Survey, Man.
2, xi + 166 pp., illus.
Berg, C. O.

1953. Sciomyzid larvae (Diptera) that feed on snails. Jour. Parasit., vol.
39, pp. 630-636.
Binney, W. G., and Bland, T.

1869. Land and fresh water shells of North America. Smithsonian Miscl.
Coll., 194, pt. 1, xii + 316 pp., 544 figs.
Binney, W. G., and Gray, A. F.

1885. Notes on the land shells of Kansas. Bull. Washburn Coll. Lab. Nat.
Hist., vol. 1, pp. 55-56.

Call, R. E.

1886. Notes on the land Mollusca of the Washburn College Biological Sur-
vey of Kansas. Bull. Washburn Coll. Lab. Nat. Hist., vol. 1,
pp. 201-205, 1 fig.

Ellis, A. E.

1926. British Snails. A guide to the non-marine Gastropoda of Great
Britain and Ireland, Pliocene to Recent. Oxford, the Clarendon
Press, 275 pp., frontispiece, 14 pis., 11 text-figs.
Franzen, D. S.

1944. New state records of Mollusca from Kansas. Trans. Kansas Acad.
Sci., vol. 47, pp. 261-273, 1 pi., 1 text-fig.
Franzen, D. S., and Leonard, A. B.

1942. A preliminary survey of the Mollusca of Kingman County, Kansas.
Trans. Kansas Acad. Sci., vol. 45, pp. 334-343, 2 pis., 1 map.

1947. Fossil and living Pupillidae ( Gastropoda-Pulmonata ) in Kansas.
Univ. Kansas Sci. Bull., vol. 31 (pt. 2), pp. 311-411, 6 pis., 1 table,
15 text-figs.
Hanna, G. D.

1909. The Mollusca of Douglas County, Kansas. Nautilus, vol. 23, pp. 81-
82, 94-96.
Hickman, C. P.

1931. The spermiogenesis of Succinea ovalis Say, with special reference
to the components of the sperm. Jour. Morph., vol. 51, pp. 243-289,
8 pis., 2 text-figs.
Ingram, W. H., and Hewitt, O. H.

1943. Sporocysts of Leucochloridium in Succinea from New York State.
Nautilus, vol. 56, pp. 92-95.

Kagan, I. G.

1949. Quickella (Family Succineidae) a new host for sporocysts of Leu-
cochloridium (Trematoda:Brachylaemidae) in southeastern Mich-
igan. Jour. Parasit., vol. 35 (6) (suppl.): 38-39.
Lee, C. B.

1951a. The Molluscan family Succineidae in Michigan. Considerations of
anatomy, early embryology, and distribution. Unpublished Doctor's
dissertation, Univ. Michigan, 269 pp.

1951b. Succinea vaginacontorta (Section Calcisuccinea), a new amber
snail from Kansas. Occas. pap. Mus. Zool., Univ. Michigan, no. 533,
pp. 1-7, 2 pis., 1 text-fig.

1534 The University Science Bulletin

Leonard, A. B.

1950. A Yarmouthian Molluscan fauna in the midcontinent region of the
United States. Univ. Kansas Paleont. Contributions, art. 3, pp. 1-48,
6 pis., 4 text-figs.
Leonard, A. E.

1943. The Mollusca of Meade and Clark counties, Kansas. Trans. Kansas
Acad. Sci., vol. 46, pp. 226-240, 2 pis., 2 text-figs.
Leonard, A. B., and Leonard, A. E.

1946. Mollusca from Greenwood County, Kansas. Univ. Kansas Sci.
Bull., vol. 31, pt. 1, pp. 115-122.
Leonard, A. B., and Goble, C. B.

1952. Mollusca of the University of Kansas Natural History Beservation.
Univ. Kansas Sci. Bull., vol. 34, pt. 2, pp. 1013-1055, 5 pis.,
1 table.

Pilsbry, H. A.

1948. Land Mollusca of North America (North of Mexico). Acad. Nat.
Sci. Philadelphia, Mon. no. 3, 2 (pt. 2), xlvii + 592 pp., 585 figs.
Quick, H. E.

1933. The anatomy of British Succineae. Proc. Mai. Soc. London, vol.
20, pp. 295-318.
Shimek, B.

1935. The habitats of Iowa Succineas. Nautilus, vol. 23, pp. 6-10.
Strandine, E. J.

1941. Quantitative study of a snail population. Ecology, vol. 22 pp
86-91, 5 tables, 4 figs.
Tryon, G. W.

1866. Monograph of the terrestrial Mollusca of the United States. Amer.
Jour. Conch., vol. 2, pt. 3, pp. 218-277, 4 pis.
Webb, G. B.

1953. Anatomical studies on some midwestern Succineidae and two new
species. Jour. Tennessee Acad. Sci., vol. 28, pp. 213-220, 5 pis.

1954. Pulmonata, Succineidae: Succinea ( Desmosuccinea ) pseudavara,
new section and species. Gastropodia, vol. 1 (no. 2), pp. 10, 18-19
12 figs.

Family Succineidae (Gastropoda: Fulmonata) 1535

GUIDE TO TERMS

apl, accessory plate
ag, albumin gland
ap, appendix
ce, cutting edge
ep, epiphallus
ex, extension of appendix
fs, fertilization sac
ga, genital atrium
hd, hermaphrodite duct
hg, hermaphrodite gland
od, oviduct

USED IN FIGURES

p, penis

pg, prostate gland

pr, penial retractor muscle

ps, penial sheath

r, ribs

sr, seminal receptacle

srd, duct of seminal receptacle

sv, seminal vesicles

tva, torsion in vagina

va, vagina

vd, vas deferens

1536 The University Science Bulletin

PLATE I

Explanation of Figures

Dorsal and ventral views of shells shown in all figures

Fig. A—Quickella wandae Webb, No. 10274, April 8, 1956, 7 mi. N Ottawa,
Franklin County, Kansas.

Fig. B— Succinea ovalis Say, No. 10265, April 22, 1956, -/i„ mi. N White
Cloud, Doniphan County, Kansas.

Fig. C — Succinea vaginacontor.a Lee, No. 10266, June 3, 1956, Meade
County State Park, Meade County, Kansas.

Fig. D— Succinea pseudavara Webb, No. 10268, June 3, 1956, Cimarron,
Gray County, Kansas.

Fig. E—Quickella vagans (Pilsbry), No. 10273, July 28, 1955, Lone Star
Lake, Douglas County, Kansas.

Fig. F — Succinea concordialis Gould, No. 10270, May 16, 1956, K. U. Nat-
ural History Reservation, 4 mi. N, Vk mi. E Lawrence, Douglas County, Kansas.

Fig. G—Oxijloma retusa (Lea), No. 10277, May 11, 1956, 1% mi. S Mus-
cotah, Atchison County, Kansas.

Fig. H— Oxijloma retusa (Lea), No. 10276, September 4, 1955, %,, mi. N
Iowa Point, Doniphan County, Kansas.

Fig. I — Succinea concordialis Gould, No. 10271, June 5, 1956, 6 mi. W
Altamont, Labette County, Kansas.

Fig. J — Quickella vagans (Pilsbry), No. 10272, June 12, 1956, 2% mi. W
Burlingame, Osage County, Kansas.

Fig. K — Succinea ovalis Say, No. 10265, April 22, 1956, V-, mi. N White
Cloud, Doniphan County, Kansas.

Fig. L— Succinea ovalis Say, No. 10264, April 9, 1955, % mi. N White
Cloud, Doniphan County, Kansas.

All figures approximately X 2

Catalogue numbers are of the Mollusk Collection, University of Kansas
Museum of Natural History.

Family Succineidae (Gastropoda: Pulmonata) 1537

PLATE I

1538 The University Science Bulletin

Figures 2-4

Fig. 2. Genitalia of Succinea ovalis Say, K. U. 10264, April 9, 1955, V-, mi.
N White Cloud, Doniphan County, Kansas.

Fig. 3. Genitalia of Succinea concordialis Gould, K. U. 10327, June 2, 1956,
St. Francis, Cheyenne County, Kansas.

a, complete genitalia.

b, drawing to show penis as it appear within the sheath.

Fig. 4. Genitalia of Succinea vaginacontorta Lee, K. U. 10266, June 3,
1956, Meade County State Park, Meade County, Kansas.

Family Succineidae (Gastropoda: Pulmonata) 1539

Figures 2-4

1540 The University Science Bulletin

Figures 5-7

Fig. 5. Genitalia of Succinea psendavara Webb, K. U. 10334, June 4, 1956,
4 mi. W Medicine Lodge, Barber County, Kansas.

Fig. 6. Genitalia of Oxyloma retusa (Lea), K. U. 10285, July 14, 1955,
1/2 mi. S Muscotah, Atchison County, Kansas.

Fig. 7. Drawings of the penes and appendices of three specimens of Oxy-
loma retusa from one population, to show the variability in the appendix.
Sheath of penis not shown. Note that the degree of coiling of epiphallus
seemingly influences the position of the appendix. In a and b, the penis is
coiled in a half-turn. In c, the straightening of the epiphallus has uncoiled the
penis. The appendices shown in b and c represent about the maximum size
attained by the structure. The appendix in a is more nearly typical, although
many specimens of Oxyloma retusa possess smaller appendices than is shown
here.

Family Succineidae (Gastropoda: Pulmonata) 1541

Figures 5-7

AG-

1542 The University Science Bulletin

Figures 8-10

Fig. 8. Genitalia of Quickella vagans (Pilsbry), K. U. 10272, June 12,
1956, 2% mi. W Burlingame, Osage County, Kansas.

Fig. 9. Genitalia of Quickella wandae Webb, K. U. 10274, April 8, 1956,
7 mi. N Ottawa, Franklin County, Kansas.

Note: the penis and appendix have been twisted approximately 180 degrees
in this drawing to show structures more advantageously. The normal position
of these organs would be about the same as shown in Fig. 8.

Fig. 10. a, jaw of Succinea ovalis showing ribs on the anterior surface of
the cutting edge. This is the only species of the Succineidae in Kansas possess-
ing ribs, b, jaw of Succinea concordialis. The accessory plate (apl) shown in
Figs. 10a and 10b is present on the jaws of all species of Succineidae and is a
diagnostic feature of the family. Camera lucida drawings; figures a and b are
drawn to same scale ( X 40 ) .

Family Succineidae (Gastropoda: Pulmonata) 1543

Figures 8-10

AG — H

OD

1

i'V'l

SR

..>.Jf

TO

a p ,.., ,;;:,■.',„ , , ,r,,^

EP
PR

-Slgr

^^

8

AG —
SV —

FS —

OD— -

VD

-SRD
i-GA

A

'■?f*

-APL

10 a

plffl APL

>"T':^6r— CE
10b

/,-; ?**■

SB

,^-HG

T^4L^vOff"\H '^' HD

r eg

c

--■k.i--

PG

^i

SRi
SRD^—
GA

EP-

v-~ -^

— 1

3fi^C 55*-

-VD

.VA

AP
•PR

EX

□

26-80.50

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Kansas University Quarterly

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