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WO 



HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



BULLETIN 



OF THE 



MUSEUM OF COMPARATIVE ZOOLOGY 



AT 



HARVARD COLLEGE, IN CAMBRIDGE 



VOL. 126 



CAMBRIDGE. MASS., USA 
1961 - 1962 



Bulletin of the Museum of Comparative Zoology 
AT HARVARD COLLEGE 

Vol. 126 No. 1 



SALIENTIA OF VENEZUELA 



By Juan A. Rivero 

Institute of Marine Biology and Biology Department 

University of Puerto Rico 

Mayaguez, Puerto Rico 



With One Plate 



CAMBRIDGE, MASS., U.S.A. 
PRINTED FOR THE MUSEUM 

November, 1961 



Publications Issued by or in Connection 

WITH THE 

MUSEUM OF COMPARATIVE ZOOLOGY 
AT HARVARD COLLEGE 



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Bulletin of the Museum of Comparative Zoology 

AT HAEVAED COLLEGE 

Vol. 126. No. 1 



SALIENTIA OF VENEZUELA 



By Juan A. Rivero 

Institute of Marine Biology and Biology Department 

University of Puerto Eico 

Mayaguez, Puerto Eico 



With One Plate 



CAMBRIDGE, MASS., U.S.A. 
PRINTED FOE THE MUSEUM 

November, 1961 



No. 1 — Salientia of Venezuela 
By Juan A. Rivero 

CONTENTS 

PAGE 

Introduction 6 

Acknowledgments 6 

Historical Eesume 7 

Method of Treatment 10 

Explanation of Terms and Measurements 11 

Key to the Families of Venezuelan Frogs 16 

Pipidae 17 

Key to Venezuelan Pipa 17 

Pipa pipa (Linn6) 17 

Pipa parva Euthven and Gaige 18 

Buf onidae 19 

Key to Venezuelan Bufo 19 

Bufo guttatus guttatus Schneider 20 

Bufo ceratophrys Boulenger 22 

Bufo granulosus granulosus Spix 23 

Bufo marinus marinus (Linne) 25 

Bufo sternosignatus Giinther 28 

Bufo typhonius typhonius (Linne) 29 

Bufo typhonius alatus Thominot 31 

Leptodactylidae 32 

Key to Venezuelan leptodactylid genera 32 

Key to Venezuelan Leptodactylus 32 

Leptodactylus marmoratus hylaedactylus (Cope) 33 

Leptodactylus pentadactylus pentadactylus (Laurenti) 35 

Leptodactylus rhodomystax Boulenger 37 

Leptodactylus bolivianus Boulenger 38 

Leptodactylus mystaceus (Spix) 40 

Leptodactylus poeciloohilus dypticus Boulenger 42 

Leptodactylus sibilatrix (Wied) 44 

Leptodactylus ocellatus (Linne) 45 

Leptodactylus podicipinus petersii (Steindachner) 47 

Leptodactylus rugosus Noble 50 

Lithodytes lineatus (Schneider) 52 

Key to Venezuelan Eleutherodactylus 53 

Elcutherodactylus cornutus maussi (Boettger) 54 

Eleutherodactylus briceni (Boulenger) 56 

Eleutherodactylus turumiquirensis sp. n 57 

Eleutherodactylus terra ebolivaris sp. n 58 

Eleutherodactylus brachypodius sp. n 61 

Eleutherodactylus conspicillatus ileamazonicus ssp. n 63 



4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Eleutherodactylus stenodiscus Walker and Test 66 

Elcuthcrodactylus orocostalis sp. n 68 

Eleutherodactylus bicumulus (Peters) 70 

Eleutherodact ylus icilliamsi sp. n 72 

Eleutherodactylus rozei sp. n 73 

Eleutherodactylus urichi (Boettger) 75 

Eleutherodactylus reticulatus Walker and Test 76 

Eleutherodactylus racenisi sp. n 78 

Eleutherodactylus marmoratus (Boulenger) 80 

Eleutherodactylus anotis Walker and Test 81 

Key to Venezuelan Ceratophrys 83 

Ceratophrys calcarata Boulenger 83 

Ceratophrys cornuta (Linne) 86 

Pseudopaludicola pusilla (Ruthven) 86 

(Paludicola fischeri Boulenger) 88 

Pleurodema brachyops (Cope) 88 

Eupenrphix pustulosus ruthvcni Netting 90 

Pseudidae 91 

Pseudis paradoxus (Linne) 91 

Hylidae 92 

Key to Venezuelan hylid genera 92 

Key to Venezuelan Eyla 93 

Eyla boaus (Linne) 96 

Hyla wavrini Parker 97 

Eyla albomarginata Spix 98 

Eyla granosa Boulenger 99 

Eyla geographica geographic-a Spix 101 

Eyla crepitans Wied 103 

Eyla albopunctata midtifasciata Giinther 105 

Eyla loveridgci sp. n 108 

Eyla ranireps (Cope) 109 

Eyla taurina (Steindachner) 110 

Eyla paramica sp. n 112 

Eyla jahni sp. n 113 

Eyla platydactyla Boulenger 115 

Eyla benitezi sp. n 116 

Eyla marahuaquensis sp. n 118 

Eyla rubra Laurenti 120 

Eyla boulengeri (Cope) 121 

Eyla baumgardneri sp. n 123 

Eyla luteooellata Roux 125 

Eyla marmorata marmorata (Laurenti) 126 

Eyla tibiatrix tibiatrix Laurenti 127 

Eyla tibiatrix ingens (Duellman) 131 

Eyla vilsomana meridensis ssp. n 131 

Eyla minuta Peters 133 

Eyla misera Werner 135 



RIVERO : SALIENTIA OF VENEZUELA 5 

Eyla orophila planicola Lutz and Lutz 136 

Hyla battersbyi sp. n 138 

Corythomantis venezolana Mertens 140 

Key to Venezuelan Gastrotheca 143 

Gastrothcca ovifera (Weinland) 143 

Gastrotheca williamsoni Gaige 144 

Nototheca pygmaea (Boettger) 145 

Key to Venezuelan Phyllomedusa 147 

Phyllomedusa bicolor (Boddaert) 147 

Phyllomedusa burmeist.eri trinitatis Mertens 148 

Phyllomedusa hypocondrialis hypocondrialis (Daudin) 150 

Centrolenidae 151 

Key to Venezuelan Cockranella 151 

Cochranella buclcleyi (Boulenger) 151 

Cochranella sp 152 

Cochranella fleischmanni (Boettger) 153 

Dendrobatidae 153 

Key to Venezuelan dendrobatid genera 153 

Key to Venezuelan Prostherapis 154 

Prostherapis collaris (Boulenger) 154 

Prostherapis shrevei sp. n 155 

Prostherapis dunni sp. n 157 

Prostherapis trinitatis trinitatis (Garman) 158 

Prostherapis trinitatis mandelorum (Schmidt) 160 

Prostherapis neblina Test 161 

Prostherapis alboguttatus (Boulenger) 163 

Key to Venezuelan Phyllobates 164 

Phyllobates bromelicola Test 164 

Phyllobates brunneus (Cope) 166 

Dendrobates leucomelas Steindachner 168 

Atelopodidae 169 

Key to Venezuelan atelopodid genera 169 

Key to Venezuelan Atelopus 169 

Atelopus oxyrhynchus Boulenger 169 

Atelopus cruciger cruciger (Lichtenstein and Martens) 171 

Atelopus cruciger vogli Miiller 173 

Oreophrynella quelchii quelchii (Boulenger) 174 

Oreophrynella quelchii macconnelli Boulenger 175 

Banidae 176 

Eana palmipes Spix 176 

Microhylidae 177 

Key to Venezuelan microhylid genera 177 

Elachistocleis oralis (Schneider) 177 

Otophryne robusta Boulenger 179 

Localities 180 

References cited 195 



b BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

INTRODUCTION 

The following report contains an account of the frogs of the 
Venezuelan Republic. It is based mainly on the material collected 
by the author in Territorio Amazonas and belonging to the Uni- 
versity of Puerto Rico (U.P.R.) and on specimens deposited in 
the Museum of Comparative Zoology at Harvard (M.C.Z.), the 
American Museum of Natural History (A.M.N.H.), the Chicago 
Natural History Museum (C.N.H.M.), the Museum of Zoology 
of the University of Michigan (U.M.M.Z.), the United States 
National Museum (U.S.N.M.) and the Museo de la Universidad 
Central de Venezuela (U.C.V.), but species reported in the litera- 
ture and unavailable to the author for examination have also 
been included. The number of Venezuelan specimens studied 
comprises a total of about 1250. Twenty-four genera and ninety- 
six species are represented. 

The paper as originally written included a description of the 
physiographical provinces of Venezuela and a short discussion of 
the fauna that each of them contains. For several reasons, it has 
been found more convenient to separate the two sections and give 
priority of publication to the taxonomic portion. It is hoped that 
the section on zoogeography will follow in the near future. 

ACKNOWLEDGMENTS 

I gratefully acknowledge the courtesies extended to me by 
members of the Venezuelan government, especially Col. Dn. Mi- 
guel Nucete Paoli, then Governor of Territorio Amazonas, and the 
secretary Dr. Luis Linares. Mr. Janis Roze, of the Universidad 
Central de Venezuela, not only has loaned me the frog collection 
of his institution but has provided valuable information on the 
localities and habits of the species included. For his courtesy I 
am more than appreciative. 

I should be ungrateful if I did not express my appreciation to 
persons that helped me collect in the field : Mr. Ventura Barnes 
Jr., leader of the University of Puerto Rico Expedition to Vene- 
zuela, 1950, whose extensive knowledge of South American tropics 
made him a valuable advisor and guide ; Dr. Jenaro Maldonado, 
entomologist and companion of several night searches for frogs 
and insects ; Dr. Paul Nesbitt, anthropologist of the expedition ; 
Dr. Hans Baumgartner of Puerto Ayacucho, a great friend of 
naturalists of Territorio Amazonas and a man with a great desire 
to help others. From the latter I received a number of specimens 
as well as his experienced cooperation in the field. 



RIVERO : SALIENTIA OF VENEZUELA 7 

For the loan of specimens used in this study or for information 
regarding specimens under their custody, I am indebted to Mr. 
Charles Bogert and Mrs. Bessie Hecht of the American Museum 
of Natural History, Dr. H. Wermuth of the Berlin Museum, Dr. 
H. W. Parker, Miss A. G. Grandison and Mr. J. C. Battersby of 
the British Museum, the late Mr. K. P. Schmidt, Messrs. Clifford 
Pope and Hyman J. Marx of the Chicago Museum of Natural 
History, Drs. Norman Hartweg, Charles Walker and James A. 
Peters 1 of the Museum of Zoology of the University of Michigan, 
Dr. J. Guibe of the Paris Museum, the late Dr. E. R. Dunn of the 
Philadelphia Academy of Sciences, Dr. Robert Mertens of the 
Senckenberg Museum and Dr. Doris Cochran of the U.S. National 
Museum. 

But my chief obligations for assistance in this work are to the 
late Dr. E. R. Dunn, Mr. Benjamin Shreve, Mr. Arthur Love- 
ridge and Dr. Ernest Williams. Dr. Dunn placed at my disposal 
through letters and through personal conversation his immense 
knowledge of South American Amphibia, while Mr. Shreve, with 
his excellent taxonomic eye and accurate memory has helped me 
solve several problems that I would have been unable to solve 
myself. Mr. Loveridge and Dr. Williams not only have taken a 
personal interest in all the phases of my work, but have read the 
manuscript and suggested some changes. However, for the short- 
comings of this paper the author assumes full responsibility. 
Prostherapis dunni, Prostherapis shrevei, Hyla loveridgci and 
Eleutherodactylus williamsi have been named after these four 
persons. 

Hyla benitezi is dedicated to Dr. Jaime Benitez, Chancellor of 
the University of Puerto Rico, without whose efforts and interest 
the expedition (during the course of which many of the specimens 
here reported were collected) would never have been possible. 

Finally, I am indebted to my wife for the bibliographical work, 
for typing the manuscript and for her constant help and en- 
couragement during the course of this work. 

HISTORICAL RESUME 

One of the earliest reports of Venezuelan amphibians is that 
of Lichtenstein and Martens (1856). Recorded as coming from 
Venezuela are: Notodelphis ovifera ( = Gastrotheca ovifera), 

i Now at San Fernando Valley State College, Northridge, California. 



8 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Hyla palmata (-H. boaus), Hyla versicolor (?), Hylodes mar- 
tinicensis (= Eleutherodactylus gollmeri) ,Crossodaetylus Gaudi- 
chaudii (?), Cystignathus ocellatus (= Leptodactylus ocellatus), 
Cystignathus typhonius (= Leptodactylus sibilatrix), Bufo agua 
(= B. marinus), Bufo Leschenaultii (= B. guttatus) and Bufo 
strumosus (= B. granulosus). Phrynidium crucigerum (= Ate- 
lopus c. eruciger) is described from Veragua. In 1863, W. Peters 
described Hylodes gollmeri on tbe basis of Lichtenstein and 
Marten's Hylodes martinicensis. I have been unable to determine 
the origin of this collection, but it appears that many of the 
specimens were collected by Consul Gollmer in the vicinity of 
Caracas. Bufo guttatus in all probability came from the southern 
forests or at least from the Orinoco Delta while Gastrotheca ovi- 
fera may be the specimen or specimens collected by Appun in 
Cumbre de Valencia (Ernst, 1877: 281). It was Dr. Dunn's 
opinion (letters, 1.7.51, 7.10.51) that this collection may have 
been mixed with one from Panama probably collected by 
Warschewitz (see remarks on page 173.) 

Other important collections in the Berlin Museum were made 
by Appun, Salomon Briceno, Kumurow, Martin, Moritz, Otto, 
Iiothe, Rosenberg, Wessel and Fisher. Less rich are the collec- 
tions received from Brandt, Bancard, Eckermann, Effeldt, Ernst, 
Gundlaeh, Hagenbeck, Hiibner, Klaebisch, Kummer, Mauss, 
Thivaites, Ursulauf, Valentiner and Liming (vVermuth, letter 
25.8.51). A small collection made by Carl Sachs in Ciudad 
Bolivar and Calabozo was reported on by Peters in 1877. 

In 1892, 0. Boettger reported on the frogs of the Senckenberg 
Museum, citing eleven Venezuelan species, all coming from 
Caracas and collected (except one) by Hiibner and Schlesinger. 
In a later paper (1893), a group of seven Puerto Cabello (Coll. 
Mauss) and one "Venezuelan" (Coll. Schultzer) species were 
added to the list and in 1896, a report was made on 17 species 
collected by Hiibner in the Alto Orinoco. 

Most of the Venezuelan specimens reported by Giinther, 1858, 
and Boulenger, 1882, were acquired by the British Museum from 
Brandt and Dyson, two Natural History dealers. Dr. Parker 
informs me (letter, 25.7.51) that he has not been able to discover 
if Brandt was ever in Venezuela. D. Dyson visited that country 
in 1851-60 and collected in Colonia Tovar, Valle de Aragua, La 
Guaira and Cariaco. 

In 1877, A. Ernst published a book in which he devoted a para- 
graph to discussing the amphibians of Venezuela, mentioning 



RIVERO : SALIENTIA OP VENEZUELA 9 

four species and two additional genera without citation of species. 
Stejneger's report on the collections made by Lyons and Robin- 
son in La Guaira and San Julian came out in the Proceedings of 
the IT. S. National Museum for 1902. Apart from some separate 
descriptions and a short report by Boulenger on the frogs col- 
lected by Brieeno in Merida (1903), little was added to Vene- 
zuelan amphibiology between 1902 and 1927, when A. Lutz 
published his "Notas sobre batrachios da Venezuela e da Ilha de 
Trinidad." Until now this has been the most complete account 
of the Venezuelan frogs, and although it contains a few errors 
of interpretation, it is undoubtedly an important contribution 
and one that has been of great help in the present studies. Dr. 
Lutz's paper is based mainly on his own collections and observa- 
tions in the region of Maracay. In the same year Roux published 
the description of Ilijla luieoccllata on the basis of two male 
specimens coming from El Mene in the Falcon State (Colls. Kug- 
ler and Vonderschmitt) . 

During more recent years other authors have mentioned a few 
species of frogs. Most important among these are Schmidt 
(1932), who reported on specimens from Mt. Turumiquire and 
vicinity (Coll. Blake), Parker (1936), who had specimens from 
the Upper Orinoco (Coll. Wavrin), Shreve (1947), who reported 
on the collection made by H. G. Kugler in Falcon State, Aleman 
(1952 and 1953) who had collections from the region of Baruta, 
El Ilatillo and Kunana, respectively, and Walker and Test 
(1955) who described three species of Eleutherodactylus from 
Rancho Grande (Coll. Test). 

The most noteworthy and until now unreported collections in 
the American museums are those of Baker (U.M.M.Z. ; Carabobo, 
Yaracuy, Tachira); Barnes (C.N.H.M. ; Yaracuy, Aragua, 
D. F.) ; Beebe (U.S.N.M. ; Aragua, Monagas) ; Briceho (Me- 
rida); Carriker (U.S.N.M.; U.M.M.Z., Merida); Cherrie 
(U.S.N.M.; Bolivar) ; Holt (U.S.N.M.; Aragua, Miranda, D. F., 
Terr. Amaz.) ; Mondolfi and Vivas Berthier (U.S.N.M.; Aragua, 
Miranda, D. F., Falcon, Guarico) ; Osgood and Osgood and Con- 
over (C.N.H.M.; Merida) ; Pinkus (U.M.M.Z. ; Roraima Region) ; 
Rosenberg (Merida-); Tate and Tate and Carter (A.M.N.II. ; 
Turumiquire, Duida and Roraima regions) ; Vogl (U.S.N.M., 
C.N.H.M.; Aragua); Schultz (U.S.N.M.; Maracaibo Basin); 
Weber (M.C.Z. ; Bolivar) ; and Williamson, E. and J. (U.M.M.Z. ; 
Carabobo, Yaracuy, Tachira). 

2 Represented in most museums. 



10 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

In 1942, E. Kohl published his "Fauna Descriptiva de Vene- 
zuela," which was revised in 1949. A few species of frogs are 
included and although there are several errors, the hook fur- 
nishes common names and contains two records otherwise re- 
ported only by Ernst. 

The Universidad Central de Venezuela has a good representa- 
tion of native frogs, which has been mostly collected by the 
director of the museum, Dr. Racenis, and Mr. Janis Roze, a young 
instructor and enthusiastic curator of Herpetology. Most of the 
specimens come from Aragua, Distrito Federal and Guarico. 
Except for the types and paratypes, the tag numbers of the 
U.C.V. specimens should be considered provisional. 

METHOD OF TREATMENT 

The Venezuelan frog fauna, as known today, comprises 24 
genera and 93 species. The discussion of each of them includes 
the Venezuelan synonymy and a description that, except when 
otherwise stated, is based strictly on Venezuelan material. Where 
specimens were not available, the original descriptions have been 
used. The contents of these have been slightly modified to suit 
my purpose but in no case (and no matter how poor the origi- 
nal) has the description been altered in meaning or the original 
remarks modernized in any way. 

The group of specimens used in the writing of the description 
in each case is listed following the synonymy. Usually the speci- 
mens chosen for the description are from a single region. This 
has been done to insure the homogeneity of the sample described. 
Following the description of the species, measurements (in mm.) 
of a male and female are given, not necessarily of the largest, but 
of the best preserved specimens. Measurements of most of the 
specimens studied were recorded but it has not been considered 
desirable to publish them in the present report. 

The list of localities includes those for the material deposited 
in the museums mentioned in the introduction of this paper as 
well as those given in the literature. When a specimen was not 
examined to my complete satisfaction (because it was tempo- 
rarily missing from the shelves or for some other reason) its 
number is followed by an asterisk. The heading "Additional 
Localities" is used in each case to include records of specimens 
that have not been used in the description. 

Following the list of localities is the range of the species. This 
includes the range within Venezuela and also other countries 



RIVERO : SALIENTIA OF VENEZUELA 11 

where the forms under consideration are said to occur. This last 
is based on the literature and on the specimens deposited in the 
different museums. Although representatives of many of the 
species occurring' in the surrounding countries were carefully 
examined, in some instances it has been necessary to accept the 
records taken from the files. 

A list of most of the localities mentioned in this paper, and a 
description of those where frogs have been collected is given at 
the end of the taxonomic section. 

EXPLANATION OF TERMS AND MEASUREMENTS 

The following is a list of the terms and measurements used in 
the text. A short explanation of amphibian terminology is given 
to help students who are beginning in this field, while Spanish 
equivalents are supplied in the belief that they may be of some 
value to the trained English-speaking specialist as well as to the 
Latin American student having little knowledge of English. 

1. Head (cabeza). The head may be as long as broad (tan 
larga como ancha), longer than broad (mas larga que ancha) 
or broader than long (mas ancha que larga). The length is 
measured between the tip of the snout and the posterior mar- 
gin of the tympanum, the breadth at the greatest distance 
between the tympana. When the tympanum is hidden the 
measurements are taken from or between the angles of the 
mouth. As the ratio of length to breadth can be easily found 
from the measurements given for each species, the character 
is only mentioned in the description when the difference from 
other species in this regard should be emphasized. In some 
cases, as in the genus Leptodactylus in general, and espe- 
cially in L. bolivianus and L. occllatus, the ratio of length 
to breadth is quite variable although the proportion given is 
the usual one. 

2. Crests or ridges (crestas o aristas). Ridges that occur on 
the heads of some bufonids and hylids. The subnasal ridge 
(arista subnasal) is usually the semicircular prolongation of 
the canthal crest around the nostrils (Bufo granulosus). 
The canthal crest (cresta cantal) runs along the canthus 
rostralis; the orbitotympanic or supratympanic (orbitotim- 
panica o supratimpanica) lies above the tympanum, and the 
orbital (orbital) around the orbit. The latter may be divided 
into pre-, supra-, post- and infraorbital. 



12 



BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 










M\ 







o 

/ 





\ 



Oy ^ 



0--0 



m 



n 



°sP 



O 



\/^ 



1 



o 



o 



O Q> 



s / 



n 



^ 7\ 



W 



A 7\ 




Fig. 1. a-g, shapes of snout; h-k, shapes of tongue; 1-s, vomerine teeth; 
t-z, canthus and loreal region. See text pp. 11-16. 

3. Snout (hocieo). The snout may be semicircular (semicircu- 
lar) as in Figure la, e.g. Centrolene buckleyi; almost semi- 
circular (casi semicircular) as in Figure lb, e.g. Hyla mar- 
morata; rounded (redondeado) as in Figure lc, e.g. Hyla 



RIVERO : SALIENTIA OF VENEZUELA 13 

granosa or Leptodactylus rugosus; subovoid (subovoide) as 
in Figure Id, e.g. Leptodactylus pentadactylus ; subelliptical 
(subeliptico) as in Figure le, e.g. Leptodactylus bolivianus; 
or acuminate (acuminado, aguzado) as in Figure If, e.g. 
Leptodactylus sibilatrix. It may project over or overhang 
the mouth (saliente) in a shark-like fashion as in Figure lg, 
e.g. Leptodactylus sibilatrix; be truncate (truncado), e.g. 
Prosthcrapis trinitatis, or swollen and elevated in front 
(hinchado y elevado al frente), e.g. Hyla granosa and boans. 

4. Derm of the head involved in cranial ossification (piel de la 
cabeza comprendida en la osificacion craniana). Easily recog- 
nized by determining if the skin is free from the cranium. 
Gastrotheca ovifera is a good example. 

5. Tongue (lengua). With respect to form, the tongue may be 
circular or rounded (circular o redondeada) as in Figure 
Ih, e.g. Centrolene buckleyi; cordiform (cordiforme) as in 
Figure li, e.g. Hyla tibiatri.r; pyriform or spatulate (piri- 
forme o espatulada) as in Figure lj, e.g. Bufo guttatus; or 
oval (ovalada) as in Figure Ik, e.g. Leptodactylus bolivianus. 
With respect to its breadth, it may be broad (aneha), e.g. 
Hyla lover idgei, or narrow and with parallel edges (estrecha 
y con margenes paralelos), e.g. Bufo granulosus. It is free 
(libre) only when attached to the anterior part of the mouth. 
and adherent (adherente) when completely attached to the 
floor. When indented (indentada) behind, it is said to be 
emarginate (emarginada), nicked or notched (escotada, in- 
cisa), and when lacking the indentation, entire (entera). 

6. Vomerine teeth or odontoids (dientes u odontoides vomeri- 
anos). In a series of two on the palate, in the vicinity of the 
posterior nares. They may be distinct (distintos) as in most 
species, indistinct (indistintos), e.g. Pleurodema brachyops, 
or absent as in Bufo, Prostherapis and some hylids. They 
may form different designs as in Figures 11 to lp ; between 
(entre) the choanae (choanas) as in Figure lq ; behind 
(detras) them as in Figure lr; or behind and between 
(detras y entre) them as in Figure Is. 

7. Canthus rostralis (canto rostral). The ridge or angle formed 
between the eye and tip of the snout by the outer edges of 
the snout bones. It may be angular (angular) as in Figure 
It, e.g. Bufo guttatus (a right angle is implied) ; obtusely 
angular (angular obtuso) as in Figure lu, e.g. Hyla boans; 
rounded (redondeado) as in Figure lv, e.g. Hyla minuta, 



14 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

indistinct (indistinto), or absent (ausente) where no external 
angle is formed and the snout would present a smooth, semi- 
circular appearance if seen in cross-section. 

8. Loreal region (region loreal, region frenal, mejilla [Es- 
pada]). The area bounded anteriorly by the nostrils, pos- 
teriorly by the eyes, dorsally by the canthus and ventrally by 
the upper lip. It may be straight or vertical (recto o verti- 
cal) as in Figure lw, e.g. Bufo guttatus; almost vertical 
(casi vertical) as in Figure lx, e.g. Bufo typhonius; little 
sloping or oblique (poco inclinado u oblicuo) as in Figure ly, 
e.g. Hyla misera and crepitans; and oblique or sloping (obli- 
cuo o inclinado) as in Figure Iz, e.g. Hyla granosa. It may 
also exhibit a depression (depresion) in which case it is 
concave or excavated (concavo o excavado) or be completely 
flat or non-concave (piano o no concavo). On occasions it 
may be slightly convex (un poco convexo). 

9. Interorbital space equal, shorter or broader than an upper 
eyelid (espacio interorbital igual, mas eorto o mas ancho que 
un parpado superior). The interorbital space is measured 
in the middle, between the two upper eyelids, the eyelid by 
depressing it slightly and measuring from the origin to the 
outer margin. When the space is said to be equal to the 
eyelid it can be expected to be slightly narrower or broader 
in some specimens. As definite numbers used in the compari- 
son (e.g., interorbital space 1% times broader than an upper 
eyelid) are of no value in practice, and may even obscure 
the importance of other characters, they are used as little as 
possible in the text. 

10. Eye diameter equal, shorter or longer than distance between 
eye and nostril (diametro del ojo igual, mas corto o mas largo 
que la distancia entre el ojo y la nariz). The diameter of 
the eye is measured between the two corners, the distance 
between eye and nostril, between the anterior corner of the 
eye and the posterior of the nostril. When the diameter of 
the eye is as long as the snout (tan largo como el hocico) it 
is implied, of course, that it is also longer than the distance 
between eye and nostril. Only in genera where the character 
is variable and important, the comparative distance between 
nostril and eye and nostril and tip of the snout is given, 
e.g., nostril closer to the eye than to the tip of the snout 
(nariz mas cerca del ojo que de la punta del hocico). 



RIVERO : SALIENTIA OP VENEZUELA 15 

11. Metacarpal and metatarsal tubercles (tuberculos metacar- 
picos y metatarsicos [Espada]). Since they are present in 
most species, they are only mentioned in the description 
where they are larger or smaller than usual or when they are 
modified in such a way as to be useful in the determination 
of the species. In some cases one or both of the metacarpal 
or metatarsal tubercles are absent. 

12. Snbarticnlar tubercles (tuberculos subarticulares, nndillos o 
pelotillas infra-articulares [Espada]). Found under the 
articulations of the digits and toes. Their mention is avoided 
if they lack any distinctive feature. 

13. Web (palmeadura [Espada], membrana interdigital, mem- 
brana natatoria). Loveridge's system of measuring the 
amount of webbing by giving the number of free phalanges 
has been adopted here as it has the advantage of being 
standard, a feature that is absent in the other system (i/o 
webbed, etc.). Since different authors use different fingers 
or toes or all of them to determine the amount of webbing, 
it is difficult to make a general comparison, but in general 
it can be said that Boulenger's % webbed fingers have li/o 
free phalanges on the outer finger, his %, 2 and his %, 
23/2. When present, the disk contains the outer phalanx. 
For measuring the number of free phalanges, the fingers 
and toes are placed close together and the extension of the 
web is determined by considering the middle, not the mar- 
gins of the membrane. The web of the first finger is that 
between the first and second fingers, of the second, that be- 
tween second and third, of the third, that between third and 
fourth and of the fourth, that between third and fourth also 
but in relation to the fourth. 

14. Lateral fringes of toes and fingers (margen cutanea de los 
dedos de los pies y las manos). This character seems to be 
quite valuable in some cases. When said to be faintly indi- 
cated, the fringes may be completely absent in some speci- 
mens. On the other hand, a species that does not normally 
show lateral fringes may show some indication of them in a 
badly desiccated state. Allowance should be made for such 
cases. In a few species, e.g., Leptodactylus podicipinus, there 
are lateral flaps that may go around the toe for 14 of its cir- 
cumference. 

15. Tarsal fold (repliegue tarsico). A cutaneous fold or ridge 
that extends usually obliquely along part or all the length of 



16 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

the tarsus. In some species, e.g., Pseudopaludicola pusilla, 
the tarsal fold begins with a tubercle in the middle of the 
tarsal segment. 

16. Metatarsal fold (repliegue metatarsieo). A slight skin fold 
or close line of tubercles tbat extends along the outer margin 
of the metatarsal segment and outer toe. 

17. Hidden portions of the hind limbs (porciones ocultas de las 
extremidades posteriores) . Those parts of a frog's hind limb 
that cannot be seen from above or below when the animal is 
in the resting position. 

18. Heel of the adpressed hind limb extends to the shoulder, the 
tympanum, the eye, the nostril, the tip of the snout (la ex- 
tremidad posterior extendida a lo largo, el tobillo llega a el 
hombro, el timpano, el ojo, la nariz, la punta del hocico). 
Obtained by stretching the bind limb anteriorly and deter- 
mining the place to which the tibio-tarsal articulation ex- 
tends. Sometimes a valuable character although it should 
not be interpreted too closely. A broken limb bone alters the 
heel extension completely. 

19. Secondary sexual characters (caracteres sexuales secunda- 
rios), as the hooks or rugosities on the inner finger of some 
males, the breast spines of Leptodactylus pentadactylus, the 
chin spines of Leptodactylus rugosus, the swollen arms of 
Leptodactylus ocellatus, the external vocal pouches of many 
species, the longer arms of the male Atelopus oxyrhynchus, 
the snout ridge of Leptodactylus mystaceus, the tubercular 
dorsum of the male Hyla teturina, etc. Most of these charac- 
ters are only shown by the males during the breeding season. 

Key to flu Families of Venezuelan Frogs 

I. Vertebrae convex anteriorly, concave posteriorly (opisthocoelous) ; 
sacrum fused with the urostyle. 

Eustachian tubes opening through one orifice into the pharynx; 

tongue and eyelids absent PIPIDAE 

II. Vertebrae concave anteriorly, convex posteriorly (procoelous) ; sacrum 
articulated with the urostyle by two condyles. 

A. Right and left half of the pectoral girdle overlapping, movable 
(arciferal). 

1. No small cartilage (intercalary) between the last and penulti- 
mate phalanges; terminal phalanges simple or T-shaped; trans- 
verse processes of the sacral vertebrae (sacral diapophysis) 
cylindrical or dilated ; belly smooth or granular. 

a. Omosternum absent, parotid gland present, upper jaw 
not toothed BUFONIDAE 



RIVERO: SALIENTIA OF VENEZUELA 17 

b. Omosternum present, parotid gland absent or small and 

inconspicuous; upper jaw toothed 

LEPTODACTYLIDAE 

2. A small piece of cartilage (intercalary) between the last and 
penultimate phalanges; transverse processes (sacral diapophy- 
sis) of the sacral vertebrae generally dilated; belly granular; 
toes generally webbed. 

a. Terminal phalanges T-shaped, astragalus and calcaneum 
fused CENTEOLENIDAE 

b. Terminal phalanges claw-shaped, astragalus and calcaneum 
not fused HYLIDAE 

3. An accessory phalanx in digits; terminal phalanges simple 
(Pseudis), or claw-shaped (Lysapsus) ; transverse processes 
(sacral diapophysis) of sacral vertebrae cylindrical; toes 
webbed PSE TJDIDAE 

B. Eight and left half of the pectoral girdle fused in the middle, 
immovable (flrmisternal). 

1. Omosternum present; sternum cartilaginous; fingers and toes 
with dermal scutes on the disks DENDBOBATLDAE 

2. Omosternum absent; sternum bony; fingers and toes without 
dermal scutes on the tips ATELOPODIDAE 

III. First seven vertebrae procoelous; 8th biconcave, sacral, biconvex (di- 
plasciocoelous), articulated with the urostyle by two condyles. 

A. Teeth present, palate normal EANIDAE 

B. Teeth absent, palate with two dermal ridges in front of the 
pharynx MICEOHYLIDAE 

PIPIDAE 

Key to the Species of Pipa Recorded from Venezuela 

I. Two lateral flaps at the angle of the mouth ; one large metatarsal 

tubercle pipa 

II. Lateral flaps at the angle of the mouth absent; no metatarsal tubercles 
parva 

Pipa pipa (Linne) 

Rana pipa Linne, 1758, Syst. Nat., ed. 10: 210: Surinam. 
Pipa dorsigera Ernst, 1877, Flora y Fauna de Ven. : 281. 
Pipa pipa Dunn, 1948, Amer. Mus. Novit., no. 1384: 9. 
Pipa americana Eohl, 1949, Fauna Descr. de Ven., ed. 2: 395, fig. 177. 
No Venezuelan material examined. 

1 (M.C.Z. 6127) Trinidad. 
3 (M.C.Z. 1244) Dutch Guiana. 
Description. Head flattened, openly triangular but with a 
blunt tip ; nostrils transversely elongated, sometimes with a 



18 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

slightly elevated piece of skin at the outer edges ; a short tentacle 
under the tip of the snout; mouth very large, with two distinct 
lateral flaps at its angles ; one or more short, pointed appendages 
above the small eyes and sometimes another pair on each side of 
the snout under the upper lip ; fingers long and slender, each 
ending in four tips ; metacarpal tubercles absent ; thighs very 
short ; a very prominent inner metatarsal tubercle but no outer ; 
toes broadly webbed to the tips. Skin above, shagreened and 
with small round tubercles and lateral line organs. Below, rugose. 

Measurements. Snout-vent S 135, 9 120. 

Additional Localities. A group of petroleum geologists 
(Lopez, Davey y Rubio, 1946 : 123) have observed this species in 
the Delta Region. Ernst and Rohl do not give any special local- 
ity. 

Range. Eastern Venezuela. Trinidad and the Guianas to Matto 
Grosso; Colombia to Peru and probably Bolivia. 

Remarks. According to Rohl the common name in Venezuela 
is "sapo de celdas." 

Pipa parva Ruthven and Gaige 

Pipa parva Euthven and Gaige, 1923, Occ. Papers Mus. Zool. Univ. Mich., 
No. 136 : 1 : Sabana de Mendoza, Venezuela ; Noble, 1925, Amer. Mus. 
Novit., No. 164: 2; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 38, 41, 
pi. viii, fig. 1; Dunn, 1948, Amer. Mus. Novit., No. 1384: 8. 
Protopipa parva Carvalho, 1939, Bol. Biol., 4: 397, figs. 6b, 6c. 

1 (U.M.M.Z. 55743, type) Sabana de Mendoza. 
4 (U.M.M.Z. 57444-57447, prtps.) Sabana de Mendoza. 
1 (M.C.Z. 9012, prtp.) Sabana de Mendoza. 
47 (U.S.N.M. 115770-816) 20 km. w. Rosario, iii.42. 
Description. Head depressed ; snout rather pointed and pro- 
jecting beyond the mouth ; nostrils transversely elongated, ending 
laterally in two projecting tips; eyes small, their diameter much 
shorter than distance between eye and nostril ; interorbital space 
broad ; no dermal appendages at the tip of the snout nor in any 
other place on the head ; arms relatively short ; fingers short, with 
a small basal web, each ending in four short appendages; meta- 
tarsal and subarticular tubercles absent ; toes fully webbed, their 
tips pointed ; tarso-metatarsal articulation of the adpressed hind 
limb extends to the eye or between eye and nostril. Skin above, 
uniformly covered with keeled tubercles. Below, tubercular ex- 
cept on the lower lip. 



RIVERO: SALIENTIA OF VENEZUELA 19 

Color. Grayish brown, tinged with yellowish and with obscure 
darker markings. Below, brownish yellow. 

Measurements. Snout-vent $ 29, 9 34 ; head breadth $ 13, 
9 9.5 ; head length S 7, 9 6 ; femur 6 12.5, 9 12,3 ; tibia & 
13.5, 9 13. 

Habits. Females with swollen backs were found to be gravid. 
Apparently some hormone ( ?) stimulates swelling prior to spawn- 
ing. Some specimens only 11 mm. in length had swollen backs. 

Additional Localities. El Mene, in a roadside puddle that 
drains into Rio Cocuiza (U.S.N.M. 115818-20) ; Lagunillas 
(U.S.N.M. 11757-8) ; Rio San Juan (U.S.N.M. 115767-9) ; Zulia 
(U.S.N.M. 128845, juv.). 

Range. The arid and semiarid Maracaibo Basin and the Fal- 
con Region. Northeastern Colombia. 

Remarks. This species represents the only frog that is ap- 
parently endemic to the arid and semiarid region of northwestern 
Venezuela. It seems to be most closely related to Pipa aspera of 
the Guianas, but apparently there is complete discontinuity of 
their ranges. It is possible that in previous times an ancestral 
form occupied all the area from the Maracaibo Basin to the 
Guianas and that the range was broken by the changes that oc- 
curred during the Ice Age. Living in a medium of high specific 
heat, this mostly aquatic animal was able to persist in the Mara- 
caibo Basin while failing to survive in the Coastal Range owing 
to the peculiar conditions existing there at the time. 

I agree with Dunn (1948) in including the genera Pipa, Proto- 
pipa and Hemipipa in the single genus Pipa. 

BUFONIDAE 

Key to the Species of Bufo Reported from Venezuela 
I. Head without bony ridges. 1 

A. Snout truncate; canthus angular; a wedge-shaped, skin-covered 

ridge in front of the eye g. guttatus 

II. Head with bony ridges. 1 

A. Upper eyelid with a hornlike dermal appendage. 

1. A lateral fringe of pointed tubercles; snout with a triangular 
tip c.eratophrys 

B. Upper eyelid normal. 

1. A subnasal ridge; tympanum in contact with the descending 
ramus of the postorbital ridge ; nostrils directed upward ; heel 
never extending to parotid g. granulosus 

2. No subnasal ridge; tympanum not usually in contact with the 
descending ramus of the postorbital ridge; nostrils lateral or 
almost so; heel generally extending to parotid or beyond. 

i This character does not usually work for young specimens. Such juveniles 
should be closely examined for the presence or absence of the supraorbital ridge. 
The use of the other key characters may also be necessary. 



20 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

a. Angles of the jaw not projecting. 

1. Tarsal fold strong; snout not prominent, gradually in- 
clined downward toward the front m. marinus 

2. Tarsal fold absent; snout prominent, not gradually in- 
clined downward toward the front. 

a. Palms and soles spinulous ; metatarsal fold absent 
sternosignatus 

b. Palms and soles not spinulous ; metatarsal fold present 
typhonins aJatus 

b. Angles of the jaw strongly projecting; palms and soles not 
spinulous; metatarsal fold present t. typhonins 

Bufo guttatus guttatus Schneider 

Bufo guttatus Schneider, 1799, Hist. Amphib., 1: 218: "India Orientali"; 

Boettger, 1896, Ber. Senckenb. Naturf. Ges.: LIV. 
Bufo Leschenaultii Lichtenstein and Martens, 1856, Nomencl. Rept. Amphib. 

Mus. Berol. : 42. 

7 (U.P.R. 30-36) Pto. Ayacucho, v.50. 

4 (U.P.R. 37-40) Tapara, vi.50. 

5 (U.P.R. 42-46) Raudal de Dios, v.50. 

2 (U.P.R. 41, 47) La Culebra, 1,000 ft., v.50. 

1 (M.C.Z. 17708) Sn. Fernando, 1895. 
Description. Snout truncate, as long as, or very slightly longer 
than, the eye diameter; tongue pyriform, entire or slightly nicked 
behind ; eye diameter greater than distance between eye and 
nostril; interorbital space flat, about twice as broad as an upper 
eyelid ; canthus angular ; loreal vertical, flat ; tympanum distinct, 
y 2 the eye diameter ; a wedge-shaped, skin-covered ridge in front 
of the eye ; other head ridges absent ; sometimes the slight but 
thick supratympanic eminence appears to be an anterior exten- 
sion of the parotid ; parotids large, distinct, pitted, extending on 
the sides as far down as the lower margin of the tympanum; 
first finger much longer than second; fingers and toes swollen 
at the tips, with a thick but not very distinct lateral fringe; 
subarticular tubercles large, single, rounded, that of the first 
finger larger than the others ; tarsal fold fairly prominent ; meta- 
tarsal fold absent; inner metatarsal tubercle very prominent; 
toes taken in order from first to fifth exhibit the following 
phalanges free of web: 1*4, 1, 2, 3%, 2; heel of the adpressed 
hind limb extends to the posterior margin of the parotid. Skiu 
on the posterior part of the dorsum with numerous flat and very 



RIVERO: SALIENTIA OF VENEZUELA 21 

distinctive warts; flanks and limbs warty; a series of 3 or 4 
transverse rugae at the nape ; a loose fold of skin on each side of 
the body from parotid to groin ; ventral surfaces of the belly and 
thighs rugose and with flat warts or granules. 

Color. Above, olive or wood-brown with orange brown warts ; 
limbs, flanks and sides of the head blackish or brownish gray ; 
parotids particolored. Below, brownish gray, uniform or more 
usually with scattered, round, white spots; lower jaw generally 
margined with white dots ; limbs occasionally exhibiting dark 
blotches on the ventral surfaces. 

Measurements. Snout-vent $ 47, ? 85; head breadth $ .1.5, 
5 24.5 ; head length $ 14.1, $ 23 ; femur $ 16, $ 27 ; tibi'i 
$ 14, 9 26. 

Habits. Like B. typhonius with which it is usually found, 
this species apparently prefers the leaf -covered floors of wooded 
areas. Unlike B. t. typhonius, however, it seems to be more par- 
tial to lower grounds, the species ceasing to occur at about 400 
m. on the Marahuaca region, where B. typhonius is still 
abundant. 

The six largest adult females (U.P.R. 30-36) were collected at 
night while they were crossing a road after a heavy rain. Their 
ovaries were completely undeveloped. 

Additional Localities. Upper Orinoco (Boettger, 1896) ; Vene- 
zuela (Licht. and Mart., 1856). 

Range. The Venezuelan Guayana and probably the Delta Re- 
gion. The Guianas to northern Brasil and with all probability 
southeastern Colombia. 

Remarks. The label with the type specimen (Berlin 3517) in- 
dicates that it was collected in Surinam by Bloch. Although the 
locality records are not sufficient to draw any final conclusions, 
I consider Bufo glaberrimus and Bufo guttatus sufficiently close 
structurally to be regarded as conspecific. To avoid the use of a 
racial name until the intergradation is confirmed appears as un- 
desirable as to make premature use of the trinomial. These two 
forms are apparently not separated by any physical or faunal 
barrier, their ranges are allopatric and the differences they ex- 
hibit appear to be of racial value. 

Bufo guttatus guttatus is apparently very sensitive to changes 
of latitude and altitude (see "Habits" and "Range" above) and 
it is possible that these factors have contributed in differentiating 
glaberrimus although this animal is found as low as 500 m. in 
Colombia and Ecuador. It is distinguished from typical guttatus 



22 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

by the absence of preorbital ridges and in having a longer hind 
limb, the heel usually extending to the anterior half of the paro- 
tid. The skin is smooth in both forms up to a size of 35 or 40 mm. 
when the flat, rounded warts appear. These are generally more 
orange in guttatus than in glaberrimus. I have not discovered 
any difference in webbing between the two forms. 

The rose spot in the inguinal region described by Giinther for 
B. glaberrimus is found in a specimen from Macanal, Colombia 
(M.C.Z. 15059, 68 mm.) but not in those from Sarayacu, Ecuador 
(M.C.Z. 19603-12 plus 56 dupl., largest 53% mm.). The latter 
are generally dark in the anterior part of the belly and throat but 
the white round spots that occur in some B. guttatus are some- 
times present in B. glaberrimus. 

M.C.Z. specimens 3807-8, 2968 (largest 5iy 2 mm.) from Sali- 
dero, N. W. Ecuador undoubtedly represent a race, distinguished 
by the variegated dorsum, greater amount of web and densely 
dotted ventral surface. It is not impossible that they represent 
the immature condition of the recently described Bufo blombergi 
Myers and Funkhouser, which might also be included in the same 
Rassenkreis. Boulenger (1882) mentions a Bufo guttatus gut- 
tatus of 177 mm. (Demerara), so it appears that the species in 
general may attain considerable size. Besides its normally large 
size, however, Bufo blombergi is distinguished from glaberrimus 
by several minor differences, and geographically it is disjunct, 
being separated by the whole Andean Range. 

I believe Bufo anderssoni Melin (Taracua, Uaupes R., Brasil) 
is a synonym of Bufo guttatus guttatus. This would appear from 
the ridge in front of the eye shown in one of the figures. 

Bufo ceratophrys Boulenger 

Bufo ceratophrys Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 319, pi. 
xii, fig. 2 : Ecuador. 

1 (U.P.R. 253, juv.) Temiche, 4050 ft., v.50. 
Description. Snout with a pointed triangular tip ; nostrils 
somewhat raised laterally ; tongue narrow, entire ; eye diameter 
greater than distance between eye and nostril ; interorbital space 
broader than an upper eyelid ; canthus well defined ; loreal al- 
most straight ; tympanum hidden ; probably slight supraorbital 
and supratympanic ridges ; parotids moderate, subovate, super- 
ficial ; first finger a little shorter than second ; a moderate meta- 
tarsal fringe to the tip of the outer toe ; subarticular tubercles 
large ; toes webbed at the base only ; heel of the adpressed hind 



RIVERO: SALIENTIA OF VENEZUELA 23 

limb extends to the eye. Skin above covered with small tubercles ; 
three distinct tubercles at the angle of the mouth and several 
others at the ventral base of the arm; upper eyelid with a tri- 
angular dermal appendage ; a lateral fringe of tubercles from 
parotid to groin. 

Color. Above, brown with minute dark points and a dark spot 
on one of the sides of the dorsum ; fringes along the sides of the 
body black-edged; limbs crossbarred. Below, uniformly light 
brown. 

Measurements. Juv., snout-vent. 12 mm. 

Habits. The only specimen was collected at night on the leaf- 
covered floor of very humid forest. 

Range. Only known from Marahuaca in Venezuela. Eastern 
face of the Andes of Ecuador. 

Remarks. Although very small, the specimen shows sufficient 
characters for its proper allocation to the species. It differs from 
two Ecuadorean specimens examined (M.C.Z. 19601-2, Canelos 
to Maranon) and from the original description in having the 
first finger shorter than the second, a hidden tympanum and no 
well-developed lateral papillae. The last two characters are prob- 
ably a sign of immaturity as the smaller of the Ecuadorean 
specimens is similar in these respects. 

So far as I am aware, the species has not been reported from 
the Amazonian forests between the Guayanan Cerros and the 
Andes. It represents the first Guayanan frog showing Andean 
affinities and a confirmation of similar findings in plants, birds 
and mammals. More abundant and adequate material of Bufo 
ceratophrys may possibly show some racial differentiation from 
the Andean form. 

Bufo granulosus granulosus Spix 

Bufo granulosus Spix, 1824, Spec. Nov. Testud. Ran.: 51, pi. xxi, fig. 2: 
Bahia ; Boettger, 1892, Kat. Batr. Samni. Mus. Senckenb. : 39 ; Lutz, A., 
1927, Mem. Inst. Osw. Cruz, 20: 38, 42, pi. viii, figs. 5, 6; Parker, 1936, 
Bull. Mus. Roy. Hist. Nat, Belgique, 12: 1. 

Bufo strumosus Lichtenstein and Martens, 1856, Nomencl. Rept. Amphib. 
Mus. Berol. : 42. 

Bufo granulosus granulosus Miiller and Hellnuch, 1936, Wissen. Ergeb. 
Deutsch Gran Chaco Exped. : 13. 

13 (U.P.R. 16-28) Sn. Fernando de Atabapo, vi.50. 
1 (U.P.R. 29) Pto. Ayacucho, vi.50. 



24 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Description. Snout prominent, slightly longer than the eye 
diameter ; nostrils directed upward ; tongue narrow, entire ; eye 
diameter greater than distance between eye and nostril ; inter- 
orbital space about IV2 times broader than an upper eyelid ; 
canthus curved, denned by a horny ridge that extends anteriorly 
as a semicircle around the nostril ; loreal slightly oblique ; tym- 
panum moderate, the anterior border in contact with the post- 
orbital ridge, the posterior margin usually obscured by granules ; 
orbits encircled by a low but distinct, horny ridge ; a horny labial 
and a short supratympanic ridge ; parotids of moderate size but 
not distinctly marked ; first finger not extending beyond second ; 
a rounded palmar and a smaller and less distinct inner meta- 
carpal tubercle ; subarticular tubercles single, the second of the 
third finger double ; edges of fingers and toes serrated ; tarsal 
and metatarsal folds absent; toes taken in order from first to 
fifth exhibit the following phalanges free of web: 1%, 1%, 2,-2y 2 , 
3%, iy^-2; heel of the adpressed hind limb does not reach the 
parotid. Skin above, tubercular, spiny on the sides and limbs. 
Male with a subgular vocal sac, and dark rugosities on the inner 
side of the first digit. 

Color. Above, light brown or yellowish gray, with irregular 
dark gray markings ; head ridges black. Below, yellowish gray, 
usually marbled with darker gray on the breast and anterior part 
of the belly. 

Measurements. Snout-vent $ 46.7, 9 52 ; head breadth 8 
14.5, 9 16 ; head length $ 12.1, 9 14 ; femur $ 10, 9 13 ; tibia 
$ 13.5, 9 17. 

Habits. Most of the specimens were caught in amplexus in 
the shallow waters of a rain pool in the savannas of San Fer- 
nando. It is strange that while the B. marinus collected in the 
same locality were heavily infested with Amblyomma dissimile, 
not one of the B. granulosus was attacked by this tick. 

Of three stomachs examined, one was empty, one contained five 
small snails, mud, two small stones and a piece of wood. The 
other had one ant, one small hemipteran and three or four pellets 
of unidentifiable remains. 

Additional Localities. Arabopo (U.M.M.Z. 85131-2, 85133 [2], 
85134[2]) ; Caracas (U.C.V. 100; Boettger, 1892); Espino 
(U.C.V. 68); Lake Maracaibo (C.N.H.M. 3016[2]); Maracay 
(U.S.N.M. 97193-5; Lutz, 1927); Parmana (U.C.V. 10); Pto. 
Cabello (Licht. and Mart., 1856) ; Sanariapo (U.S.N.M. 80641- 
50) ; Upper Orinoco (Parker, 1936) ; Venezuela (Licht. and 
Mart., 1856). 



RIVERO : SALIENTIA OF VENEZUELA 25 

Range. The arid and semiarid Maracaibo Basin, the northern 
coastal belt, the arid and semiarid sections of the Coastal Range, 
the Llanos and the northern half of the Venezuelan Guayana. 
Colombia and the Guianas to northern Argentina. 

Remarks. The specimens from Arabopo (4000 ft.) have very 
prominent crests and perhaps should deserve racial recognition. 

Bufo marinus marinus (Linne) 

Rana marina Linne, 1758, Syst. Nat., ed. 10: 211: America (Bestricted to 
Surinam by Miiller and Hellmich, 1936). 

Bufo agua Lichtenstein and Martens, 1856, Nomencl. Kept. Amphib. Mus. 
Berol.: 42; Giinther, 1858, Cat. Batr. Sal. Brit. Mus.: 65. 

Bufo marinus Peters, 1877, Monatsb. Akad. Wissensch. Berlin: 460; Bou- 
lenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 315; Boettger, 1892, 
Kat. Batr. Samm. Mus. Senckenb.: 39; 1893, Ber. Senckenb. Naturf. 
Ges.: 40; 1896, Ber. Senckenb. Naturf. Ges.: LIV; Stejneger, 1902, 
Proc. U. S. Nat. Mus., 24 : 180 ; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 
20: 38, 42; Aleman, 1952, Mem. Soc. Cienc. La Salle, U\ 27, fig. 3. 

Bufo marinus marinus Schmidt, K. P., 1932, Zool. Ser. Field Mus. Nat. 
Hist., 18: 159. 

7 (U.P.R. 48-51, 76-7, 89) Sn. Fernando de Atabapo, vi.50. 

1 (U.P.R. 52) Chorro Chupadero, iii.50. 

Description. Snout slightly longer than the eye diameter; 
tongue oval or pyriform; eye diameter greater than distance 
between eye and nostril; interorbital space about iy 2 times as 
broad as an upper eyelid; canthus well defined, slightly curved 
and sloping toward the front ; loreal short, almost vertical ; tym- 
panum not very distinct on the posterior margin, y 2 to % the 
eye diameter ; head ridges well defined, the infraorbital extending 
obliquely to the angle of the jaw, the parietal variably distinct ; 
a few tubercles behind the angle of the jaw; parotids large, 
pitted, obliquely subtriangular ; a longitudinal series of tubercles 
from hand to elbow ; first finger longer than second ; a very large, 
flat palmar and a smaller inner metacarpal tubercle ; subarticular 
tubercles generally single or incompletely fused, the second of the 
third finger double ; tarsal fold strong ; metatarsal absent ; toes 
taken in order from first to fifth exhibit the following phalanges 
free of web : 1, 1 to 1%, 2, 3 to S%, V/ 2 to 1% • the web is ex- 
tended to the tips as distinct lateral fringes ; heel of the adpressed 
hind limb extends to the parotid. Skin above and on the flanks 
covered with warts of irregular size ; upper eyelids and sides of 
the head tubercular ; ventral surfaces granular, the granules be- 
ing sometimes tipped with a dark corneus matter. Male with a 
rugosity on the inner side of the first digit. 



26 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Color. Above, brown with irregular lighter and darker mark- 
ings. Below, light brownish or yellowish, generally marbled with 
gray. 

Measurements. Snout-vent $ 44, 9 79 ; head breadth $ 15, 
9 26 ; head length $ 14, 9 24 ; femur $ 16, 9 32 ; tibia $ 17, 
9 35. 

Habits. B. marinus seems to be fairly common along the Ori- 
noco River where its voice was often heard at night. Its presence 
was not detected in the Cunucunuma nor in the Upper Duida and 
Marahuaca regions. 

Several of the specimens collected in Sn. Fdo. de Atabapo were 
infested with the tick Amblyomma dissimile. One of the animals 
had been so emaciated by the infestation that for a while it was 
taken for some other form. 

Male and female specimens were collected in a shallow rain 
pool together with B. granulosus. While most of the B. granu- 
losus were in amplexus, the ovaries of the adult B. marinus were 
completely undeveloped. 

Additional Localities. Rio Albarregas (A.M.N.H. 10501) Ara- 
bopo (U.M.M.Z. 85120[3]) ; Barrancas (M.C.Z. 19916, juv.) ; Ba- 
ruta (Aleman, 1952) ; Calabozo (Peters, 1877) ; Campo del Lago, 
Lagunillas (U.S.N.M. 115692-8) ; Caracas (Boettger, 1892) ; Ciu- 
dad Bolivar (M.C.Z. 19114, 19 juv.) ; Cocollar, Cumana, Cumana- 
coa (Schmidt, 1932) ; Duida Region (A.M.N.H. 32962) ; El 
Periquito (U.C.V. 89) ; El Valle (U.S.N.M. 128850-2) ; Espino 
(U.C.V. 50) ; Kunana, Perija Mts. (Aleman, 1953) ; La Fria 
(U.M.M.Z. 55585*); La Guaira (U.S.N.M. 22537-8, 27800-1, 
27803-6, 27796); Macuto (U.C.V. 77-8*); Maracay (U.M.M.Z. 
52707-8*, Lutz, 1927) ; 30 km. of Maturin (U.C.V. 76) ; Petare 
(U.S.N.M. 121170) ; Pto. La Cruz (U.S.N.M. 121171-2) ; Pie 
del Cerro U.S.N.M. 121168) ; Puerto Cabello (Boettger, 1893) ; 
Sn. Antonio (U.S.N.M. 836119) ; Sn. Carlos (U.S.N.M. 83610) ; 
Sn. Fernando de Atabapo (U.S.N.M. 80651); Sn. Julian 
(U.S.N.M. 27807); Sta. Catalina (M.C.Z. 19915, 4 juv.); Sta. 
Elena, Bolivar (U.M.M.Z. 85121); Sta. Lucia (U.S.N.M. 
121169); Tanaguarena (U.C.V. 36); Turgua (Aleman, 1952); 
Turgua (U.S.N.M. 129265) ; Upper Orinoco (Boettger, 1896) ; 
Venezuela (C.N.II.M. 43664? [3-6 juv.]; U.M.M.Z. 55583-4*; 
Lichtenstein and Martens, 1856 ; Giinther, 1858 ; Boulenger, 
1882). 



RIVERO : SALIENTIA OF VENEZUELA 27 

Range. Probably all the physiographical provinces of Vene- 
zuela including the Andes to an elevation of 1,600 m. Texas to 
northern Brasil. 

Remarks. No constant difference in size of parotid between 
specimens from northern and southern Venezuela was observed. 
While those of the southern animals are generally of a large size, 
some of the northern specimens (U.S.N.M. 121165, 121171, 27800) 
have glands that rival in volume any of those of the Amazonian 
group, with the possible exception of the larger southern speci- 
men, an animal from San Antonio having a snout-vent length of 
142 mm. 

The coloration of U.S.N.M. 83610 (131 mm.) is unusual: the 
ventral surfaces, the under sides of the parotids, the lower jaw 
and the posterior part of the thighs are bright yellow and dark 
marbled while the throat is profusely infuscated, almost black. 
Dr. Cochran and I have agreed that this specimen represents 
B. marinus but in coloration it is not very different from a British 
Museum Pto. Cabello specimen that has been referred to B. 
crucifer (Blgr., 1882: 317, a or b). I have examined this latter 
animal but have excluded the species from the fauna of Vene- 
zuela until its presence is confirmed by further material. 

The male Bufo marinus is more tubercular above, the tubercles 
are more spiny, and the color is usually more uniform than in 
the female. 

Cochran, 1955, treats Bufo paracnemis and Bufo ictericus as 
distinct species, on the ground that the two overlap in part of 
their ranges (Rio de Janeiro and Sao Paulo). On the east, B. 
paracnemis is said to occupy the intermediate area between north- 
ern B. marinus and southern B. ictericus, occurring in Brasil in 
Minas Gerais, Bahia and Pernambuco. 

To the west the southward extension of Bufo marinus is un- 
certain, as records may refer to any of the related forms. How- 
ever, it does not appear impossible that Bufo marinus may have 
given rise to paracnemis on the east and ictericus on the west, and 
that these two forms may have come together after their inde- 
pendent origin from a common ancestor. Anyway, there does 
not appear so far to be sufficient evidence against considering 
paracnemis a race of Bufo marinus and on that basis the tri- 
nomial is used here. 

Bufo marinus andinensis Melin, 1941, was described from 
Roque, Peru. 

Bufo pythecodactylus Werner is considered a synonym of 
Bufo marinus marinus. 



28 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Bufo sternosignatus Gunther 

Bufo sternosignatus Gimther, 1858, Cat. Batr. Sal. Brit. Mus. : 68, pi. vi, 
fig. c. : Pto. Cabello, Venezuela; Cordova, Mexico (restricted to Vene- 
zuela by Boulenger, 1882); Boulenger (part?), 1882, Cat. Batr. Sal. 
Brit. Mus., ed. 2: 323; Boettger, 1892, Kat. Batr. Samm. Mus. 
Senckenb. : 39; Gunther, 1901, Biol. Centr. Amer., Bept. and Batr.: 
247; Nieden, 1923, Das Tierreieh Anura. I: 182; Lutz, A., 1927, Mem. 
Inst. Osw. Cruz, SO: 38, 42, pi. viii, figs. 3, 4. 

Bufo typhonius sternosignatus Shreve, 1947, Bull. Mus. Comp. Zool., 99 : 535. 

2 (M.C.Z. 17812-3) Distrito Acosta, 29. 
1 (M.C.Z. 26151) Cerro Cosine, x.39. 

3 (M.C.Z. 26152, 25976-7) Pauji, v. 45. 
3 (U.M.M.Z. 55690-3) Sn. Esteban. 

1 (U.M.M.Z. 55694) Kio Bejuma. 

2 (U.M.M.Z. 55695-6) Rio La Mona. 

1 (U.M.M.Z. 55697) Bet. Valencia and Caserio Silva. 
1 (U.C.V. 16) Carayaca, xii.51. 
1 (U.C.V. 28) Cerro Avila, viii.51. 

Description. This species is very similar to Bufo typhonius 
alatus, from which it can be distinguished by the following 
characters: snout not very prominent; orbital ridges slight, the 
supratympanic usually well developed, the parietal absent or 
very indistinct ; parotids subovate, not generally pointed pos- 
teriorly ; hands and feet spinulous; the subarticular tubercles not 
different from the other spinules of palms and soles ; metatarsal 
Hold absent ; heel of the adpressed hind limb extends to the 
posterior half of the parotid. Skin above and below very granu- 
lar, occasionally coarse and spinulous especially on the sides of 
the body and venter. A lateral fringe of tubercles is found in 
some but not all individuals. 

Color. This animal is usually of a reddish color when alive. 
In alcohol it turns brown and many specimens present a colora- 
tion very similar to that of B. typhonius. Some specimens are 
peculiarly marked with yellow and brown below. 

Measurements. Snout-vent $ 41, 9 49; head breadth $ 14, 
9 16; head length $ 12, 9 14; femur $ 16.5, 9 18; tibia 
$ 16, 9 18. 

Habits. Apparently a forest animal. 

Additional Localities. Caracas (Boettger, 1892) ; El Limon 
(U.S.N.M. 121173-4) ; Mamo, La Guaira (Lutz, 1927) ; Puerto 
Cabello (Gunther, 1858; Boulenger, 1882) ; Venezuela (Gunther, 
1858; Boulenger, 1882; Nieden, 1923). 



RIVERO : SALIENTIA OF VENEZUELA 29 

Range. The Coastal Range and Falcon Region. Reported from 
Colombia by Peracca, 1914 (Cafetal Camelia, nr. Angelopolis, 
1820 m.), Werner, 1916 (Canon de Tolima), and Boulenger, 
1882 (Bogota). These records probably refer to B. typhonius 
alatus. Dunn (1944) does not mention this species in his "Herpe- 
tology of the Bogota Area" but he includes Bufo typhonius in 
the actual fauna of Bogota. 

Remarks. This species is very close to Bufo typhonius alatus 
with which it has been often confused and with which it perhaps 
interbreeds (though not freely) in part or all of its range. Gen- 
erally it is well characterized (U.M.M.Z.) but specimens are 
found (M.C.Z. and U.C.V.) that are very similar to B. typhonius, 
especially in coloration and general appearance. In all these cases 
the palm and fingers are spinulous (not a sexual character) and 
the metatarsal fringe absent. The supraorbital ridge is also more 
prominent in B. typhonius and presents a pleated appearance 
in the region where a parietal ridge would join the supraorbital. 
The upper eyelid thus appears to be lower on the sides in B. 
typhonius than in B. sternosignatus. If a perpendicular line 
were to be drawn from the upper edge of the supraorbital ridge 
to the outside, the upper eyelid would be mostly above the line in 
sternosignatus and below it in typhonius. 

Bufo typhonius typhonius (Linne) 

Fana typhonia Linne, 1758, Syst. Nat., ed. 10: 211 : America. 
Bufo typhonius Boettger, 1896, Ber. Senckenb. Naturf. Ges. : LIV. 
Bufo typhonius typhonius Leavitt, 1933, Copeia : 8. 

7 (U.P.R. 4-10) Slopes Mt. Marahuaca, 4000 ft., v.50. 

7 (U.P.R. 1-3, 11-12, 14-15) Upper Cunucunuma R., v-vii.50. 

1 (U.P.R. 13) Anaben, Colombia, vi.50. 

1 (U.S.N.M. 83949) Yapacana, iv.21. 

Description. Snout prominent, with a more or less distinct 
vertical ridge at the tip, from the level of the nostrils to the edge 
of the jaw ; tongue narrow, entire ; eye diameter slightly greater 
than distance between eye and nostril; interorbital space twice 
as broad or broader than an upper eyelid ; canthus well defined, 
carved ; loreal almost vertical and a little concave ; tympanum 
moderate, oval, % the eye diameter; preorbital and descending 
ramus of the postorbital ridge slight or absent ; supraorbital ridge 
strong, anteriorly merging with the canthal, posteriorly with the 
very prominent supratympanic crest ; parietal ridge distinct, 
forming a basin with the parotid ; parotids prominent but small 



30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

and superficial; edge of the upper jaw studded with minute acu- 
minate tubercles ; two knob-like protuberances at the angle of 
the jaws; first finger not extending beyond second; tips of the 
fingers and toes swollen, their sides serrated ; tarsal fold absent ; 
subarticular tubercles large, usually single or fused ; a row of 
closely set, small spinules along the outer edge of the metatarsal 
segment and fifth toe ; toes partially webbed, the web reaching at 
least to the penultimate phalanx in all the toes except the third, 
where it does not extend beyond the proximal phalanx; heel of 
the adpressed hind limb extends to between parotid and tym- 
panum. Skin above and on the sides of the head tubercular; 
tubercles spiny on the flanks and limbs ; a fringe of pointed 
tubercles from anterior border of parotid to groin ; ventral sur- 
faces granular, some of the granules spiny on the pectoral region. 

Color. Above, brown, uniform or with irregular lighter and 
darker markings ; from tip of snout to end of urostyle a yellowish 
vertebral stripe of varying width, on either side of which, espe- 
cially in the sacral region, roundish black spots may be present 
or absent ; flanks below the lateral fringe, occasionally lighter or 
darker than the dorsum. Below, yellowish brown, plain, infuscate 
or spotted; limbs with broad, more or less distinct crossbars. 

Measurements. Snout-vent $ 42, 9 61 ; head breadth $ 14, 
9 21 ; head length S 14, 9 18; femur <J 17, 9 25 ; tibia S 17, 
9 24. 

Habits. In life this amphibian closely resembles a dead leaf, 
its serrated sides and coloration contributing to make the simi- 
larity more striking. Belying on its ability to escape detection 
the toad usually remains motionless on the approach of danger, 
though sometimes taking several hops before merging again with 
the leaves carpeting the forest floor. 

The call of this species is probably a low-toned guttural sound, 
similar in some respects to that of Bufo marinus but much shorter 
and lower. This noise was traced to specimens of Bufo typhonius 
on several occasions but in no instance was the animal actually 
seen calling. 

Of 6 stomachs examined all contained ants, each of 3 held a 
curculionid beetle, one an isopteran, two a piece of leaf, and one 
a seed (probably ingested with the ants). An engorged specimen 
contained 30 ants, 1 mm. in length, and 2 wingless wasps, each 2 
mm. long, besides other unidentifiable insect remains. 

Additional Localities. Upper Orinoco (Boettger, 1896). 

Range. The southern part of the Venezuelan (hiayana. Ac- 
cording to Leavitt the subspecies "extends from Venezuela, south 
and east through the Guianas and well south through Brasil." 



BIVERO : SALIENTIA OP VENEZUELA 31 

Bufo typhonius alatus Thominot 

Bufo alatus Thominot, 1884, Bull. Soe. Philom. Paris, 8: 151: Obispo, Isth- 
mus of Panama. 
Bufo typhonius alatus Leavitt, 1933, Copeia : 8. 
2 (U.M.M.Z. 55571-2) Bejuma. 

1 (U.M.M.Z. 55573) Estacion Ferrocarril, Tachira. 

2 (U.M.M.Z. 55574-5, juv.) La Fria. 

1 (U.M.M.Z. 85124) W. of Wiaka Piapu Mt. 
1 (U.M.M.Z. 85125) Wiaka Piapu Mt. 

Description. Bufo typhonius alatus differs from the typical 
form in lacking the bony protuberance at the angle of the jaws 
and in having a longer hind limb, the heel of the adpressed hind 
limb extending to the tympanum or slightly beyond. The paro- 
tid is usually narrower and pointed posteriorly. 

Measurements. 6, snout-vent 50.5; head length 15; head 
breadth 18; femur 23; tibia 21. 

Habits. Apparently a forest animal. 

Range. The Coastal Range and the northwestern slopes of the 
Occidental Andes, southeastern Venezuela. According to Leavitt : 
"as much of Central America as it inhabits and extends south 
and east into Venezuela and south and west into Colombia." 
British Guiana. 

Remarks. I have been unable to confirm all the characters men- 
tioned by Leavitt, but the absence of jaw knob and the shorter 
hind limbs are perhaps sufficient for racial recognition of this 
form. On the other hand, the distribution of the two races is 
quite problematic and occasional specimens of Bufo typhonius 
from the Coastal Range have well developed bony protuberances 
at the jaws. The Colombian frogs examined lack the jaw knobs 
but a trace of these is found in Panamanian examples. 

Bufo typhonius alatus occurs in southeastern Venezuela and 
in the western part of British Guiana (U.M.M.Z. 63045, Tukeit 
Hill, Potaro R.) and with little doubt it will be found in the 
Delta Region. Specimens with all the characters of Bufo typho- 
nius typhonius have been collected as far north as Dunoon 
(M.C.Z. 6073, about 45 km. south of Georgetown, on the margin 
of the Demerara R.). It is thus possible that this form passes 
north through the valley of the Demerara or that it moves west- 
ward along the forested north coast of South America from the 
eastern Guiana. If the two animals are really races of each 
other, it appears that the zone of intergradation is longitudinal, 
more or less along the Essequibo River. 



32 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

LEPTODACTYLIDAE 

Key to the Genera of Venezuelan Leptodactylidae 

I. .Sternum cartilaginous. 

A. Tympanum absent. 

1. No vomerine teeth; a tubercle on the forearm and another on 
the tarsus Pseudopahidicola 

B. Tympanum present. 

1. Toes with disks, without web ; vomerine teeth behind the 
choanae 1 Eleutherodacti/his 

2. Toes without disks, with web Ceratophrys 

II. Sternum with a bony style. 

A. Upper jaw without teeth; a small parotid gland Eupcmphix 

B. Upper jaw with teeth ; no parotid gland. 

1. A large inguinal gland Pleurodema 

2. No inguinal gland. 

a. Terminal phalanges T-shaped Lithodytes 

b. Terminal phalanges simple Leptodactylus 

Key to the Species of Leptodactylus Reported from Venezuela 

I. Tips of the toes dilated into small, but distinct disks ; first finger not 
extending beyond second; 22 mm. marmoratiis hylaedactylus 

II. Tips of the toes not dilated into disks; first finger longer than second. 
A. Two or more dorsal folds. 

1. Dorsal folds not more than two. 

a. Dorsal folds not extending posteriorly beyond sacrum ; lat- 
eral fringes of toes slight; thighs marbled with reddish; 
breeding male with two breast and one thumb spine but no 
apparent external vocal sac; 150 mm. p. pentadacti/his 

b. Dorsal folds extending to inguinal region. 

1. Lateral fringes of toes very distinct (lateral flaps) ; no 
white longitudinal line on the posterior part of the 
thighs. 

a. Heels do not overlap when brought together in the 
anal region (tibia and femur more or less equal) ; 
anterior convexities of vomerine teeth not extending 
to choanae; 70 mm rhodomystax 

b. Heels overlap considerably when brought together in 
the anal region (tibia longer than femur) ; anterior 
convexities of vomerine teeth reaching the choanae 
and generally extending beyond their posterior mar- 
gin; 73 mm bolivianiis 

2. Lateral fringes of toes absent or faintly indicated ; a 
white, longitudinal line along the posterior part of the 
thighs. 

i Eleutherodactylus conspicillatus occasionally shows a trace of web in Ecua- 
dor. Some Antillean Eleutherodactylus also possess webs. 



RIVERO : SALIENTIA OP VENEZUELA 33 

a. An uninterrupted, white streak along the margin of 
the upper jaw to the shoulder ; dorsal folds dark on 
the anterior end; 47 mm mystaceus 

b. Two or more bars from lower eyelid to upper lip ; 
dorsal folds usually light colored throughout, 45 
mm poecilochilus dypticus 

2. Dorsal folds more than two; 1 spotted above. 

a. Snout acuminate, well projecting beyond the mouth ; lateral 
fringes of toes absent or slight; a longitudinal white line or 
close series of dots along the posterior part of the thighs; 
male with no breast or thumb spines but two black ex- 
ternal vocal sacs ; 40 mm sibilatrix 

b. Snout subelliptical, not extending much beyond the mouth; 
lateral fringes of toes distinct; no white longitudinal line 
along the posterior part of the thighs ; a series of white 
dots along the margin of the lower jaw; male with two 
large thumb spines ; 70 mm ocellatus 

B. No dorsal folds. 2 

1. Toes with very distinct lateral fringes; male with two thumb 

spines but no breast spines; color not black; 37 mm 

podicipinus petersii 

2. Toes without lateral fringes ; male with breast spines and one 
thumb spine; dorsum very rugose; color usually black; 60 mm. 
rugosvs 

Leptodactylus marmoratus hylaedactylus (Cope) 

Cystignaihus hylaedactylus Cope, 1868, Proc. Acad. Nat. Sci. Phila.: 115: 

Napo and Marafion rivers. 
Leptodactylus discodactylus Boettger, 1896, Ber. Senckenb. Naturf. Ges. : 
LIT. 

1 (U.P.R. 102) Anaben, Colombia, vi. 50. 

1 (U.P.R. 103) Marahuaca, 4,050 ft., v.50. 

1 (U.P.R. 228) La Culebra, 1,000 ft., vi.50. 
Description. Snout subelliptical, projecting slightly beyond the 
mouth ; tongue narrow, oval, indistinctly nicked behind ; vomerine 
odontoids in two short oblique series behind the choanae, 
their posterior extremities directed inwards; eye diameter 
slightly greater than distance between eye and nostril; inter- 
orbital space broader than an upper eyelid; canthus absent; 
loreal moderately inclined, concave ; tympanum about % the eye 
diameter ; a slight supratympanic fold to the shoulder and a 

i Sp.eci.meus in which some of the folds are not easily seen are found. See 
"Remarks" under L. holirianits. 

2 L. podicipinus petersii may occasionally show very slight folds on the anterior 
part of the hack. 



34 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

small, glandular ridge from shoulder to the angle of the mouth ; 
first finger not extending beyond second, which is more or less 
equal to fourth ; subartieular tubercles of fingers and toes very 
large and prominent ; fingers distally swollen ; toes dilated into 
small disks; tarsal and metatarsal folds present but indistinct; 
toes almost free, without lateral fringes; heel of the adpressed 
hind limb extends to the tympanum. Skin above, with numerous 
tubercles and short longitudinal folds, especially on the posterior 
half ; a loose lateral fold from supratympanic fold to the inguinal 
region. Below, smooth, with a discoidal, ventral fold. Breeding 
male with a slight anterior ridge on the snout and a pair of 
longitudinal external vocal sacs that extend to the base of the 
humerus. 

Color. Above, grayish or light brown, spotted with darker 
brown or black ; usually a triangular spot between the eyes ; limbs 
crossbarred. Below, uniformly white or infuscated on the throat 
and breast. 

Measurements. $ snout-vent 22.5 ; head breadth 8 ; head length 
8 ; femur 9 ; tibia 11. 

Habits. All specimens were collected on the forest floor. 

Additional Localities. Caripito (U.S.N.M. ? 2) ; Minapana 
(U.M.M.Z. 55550) ; Boettger's, 1896, L. discodactylus from the 
Upper Orinoco (Tiriquin and Guavamaco) was found to repre- 
sent this species. 

Range. The Coastal Range, the Delta Region and the Vene- 
zuelan Guayana. Southeastern Colombia, British Guiana, Suri- 
nam and south to the Napo and Maranon rivers. 

Remarks. A specimen from Surinam in the M.C.Z. is of a 
lighter ground color and the dorsolateral spots are better defined 
than in the Venezuelan specimens. The frog from Marahuaca is 
darker than the other two animals and in this respect it is closer 
to a specimen from Oyapock River, Brasil. In the two frogs 
from Caripito in the U.S.N.M., the vomerine teeth are almost 
transverse, the tongue not nicked and there is no metatarsal fold. 
U.M.M.Z. 55550 also has transverse vomerine teeth. This speci- 
men is a little desiccated. 

This form is very similar to L. marmoratus (L. dyptix, auet.) 
from which it is distinguished by the presence of small disks. T 
have considered the two forms conspeeific as one seems to repre- 
sent the other in the northern part of the continent (as already 
noticed by Cochran, 1955 : 309) and some specimens from Bolivia 
show an intermediate condition in disk development (M.C.Z. 



RIVERO : SALIENTIA OF VENEZUELA 35 

15583 and others). The only obstacle to this conclusion is Melin's 
L. rugosus from Manaos, Brasil, which is considered similar to 
these forms and according to the author, does not have disks. 
He had only one specimen and it is possible that occasional speci- 
mens without disks may be found in the north. I rather suspect 
that this animal had small disks. 1 

The situation suggested would result in L. marmoratus Parker, 
1835, L. melini, Lutz and Kloss, 1952, and probably L. andreae 
Miller, 1923, being referred to the synonymy of L. marmoratus 
hylaedactylus. 

It is possible that Steindachner's (1867) figure of Adenomera 
marmorata refers to the northern form since slight swellings that 
could be disks are shown on the toes, and the type locality (of the 
figured animal) is unknown. 2 If this is the case, then the north- 
ern form should be called L. marmoratus marmoratus and the 
southern L. m. dyptyx. For the present it seems best to follow 
Parker (1932) and apply the name L. m. marmoratus to the 
southern animal. 

Leptodactylus pentad act ylus pentadactylus (Laurenti) 

Eana pentadactyla Laurenti, 1768, Syn. Eept. : 32: Indus, i. e. Surinam (re- 
stricted by L. Miiller, 1927). 
Leptodactylus pentadactylus Lutz, A., 1927, Mern. Inst. Osw. Cruz, 20: 40. 
5 (U.P.R. 78-82) Casurua, iv.50. 
1 (U.P.R. 83) Anaben, Colombia, vi.50. 
Description. Head considerably broader than long; snout sub- 
ovoid; tongue widely oval, emarginate behind; vomerine odon- 
toids in two arched series, with their anterior convexities between 
the choanae ; eye diameter more or less equal to distance between 
eye and nostril ; interorbital space narrower than an upper eye- 
lid ; canthus obtusely angular ; loreal sloping, concave ; tympanum 
large, obliquely oval, % the eye diameter; a supratympanic fold 
from posterior corner of eye to shoulder ; first finger longer than 
second ; tarsal fold strong ; soles smooth ; subarticular tubercles 
of toes conical; Aveb absent or slight; lateral fringes of toes 
faintly indicated ; heel of the adpressed hind limb extends to the 
tympanum. Skin above, finely granular; a pair of dorsolateral 
folds from upper eyelids to sacral region and sometimes two 

i After this statement was written Lutz and Kloss (1952) collected in the same 
general region and found a small Leptodacti/lus with disks in the toes which they 
regard as Melin's form. 

2 There is, apparently, no statement of the type locality in the jar of the speci- 
men figured, for Parker, 19o2, does not mention it. 



36 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

other loose lateral folds from supratympanic fold to groin ; sides 
of the head granular. Belly and throat smooth ; an indistinct 
ventral discoidal fold. Breeding male with a bicuspid or tricuspid 
tubercle on each side of the breast ; arms swollen ; a strong spine- 
bearing tubercle on the inner side of the first digit and numerous 
small spinules on each side of the throat, breast and lower parts 
of the humerus ; no apparent external vocal sac. 

Color. Above, yellowish brown or chestnut, with darker, black 
margined, transverse bars : a black streak along the canthus 
rostralis and supratympanic fold ; upper lip with dark triangular 
spots; limbs crossbarred; a usually interrupted black line along 
the hind part of the f orelimb ; posterior part of the thighs black 
and salmon or pinkish yellow marbled ; groin and flanks generally 
tinged with salmon. Belly and throat white, immaculate or 
marbled with brown ; lower jaw dark and white barred ; throat 
infuscate; a black ventral spot at the base of the arm. 

Measurements. Snout-vent $ 146, 5 160 ; head breadth $ 57, 
9 60 ; head length S 49, 9 51 ; femur $ 54, 9 51 ; tibia $ 64, 
9 65. 

Habits. This species occurs commonly along the Cunucunuma 
River where the Maquiritare Indians use it as food. Its voice was 
often heard at night but all our specimens were brought by In- 
dians, who apparently objected to one of their greatest delicacies 
being thrown into a jar of formalin, preferring to bring a few 
themselves rather than permitting the collection of too many. 
A large foam nest attributed to this species by the Indians was 
found in a small depression between the roots of a tree. There 
were neither eggs nor tadpoles present and the nearest water was 
that of the river about 200 }-ards away. 

Of several stomachs examined, two were empty, one had the 
remains of a small frog, many large pieces of leaves and a few 
legs and other insect remains ; another contained a huge scorpion, 
by whose sting the predator was apparently unaffected (it was 
puncturing the stomach wall) and several sticks, leaves and other 
unidentifiable remains. 

Additional Localities. iy 2 hr. above Paso del Diablo (A.M.N. II. 
23164). 

Range. The Venezuelan Guayana. The species extends from 
Central America and Trinidad to Peru and southern Brasil. The 
race goes as far south as Ceara. 

Remarks. It would be interesting to discover how this species 
reached Central America as it has not yet been discovered in 



RIVERO: SALIENTIA OF VENEZUELA 37 

northern Venezuela nor in the Llanos. There is a specimen from 
the Andes of Bogota (not examined) in the collection of the 
American Museum. 

Leptodactylus rhodomystax Boulenger 

Leptodactylus rhodomystax Boulenger, 1883, Proc. Zool. Soc. London: 637, 
pi. lviii, fig. 2 : Yurimagas, Huallaga Eiver, Northern Peru. 

1 (A.M.N.H. 23175) Esmeralda, 28. 

Description. Head a little broader than long; snout subovoid 
to rounded ; tongue with a slight notch behind ; vomerine odon- 
toids in two very slightly arched series behind the choanae ; eye 
diameter equal to distance between eye and nostril ; interorbital 
space more or less of the same breadth as an upper eyelid ; tym- 
panum distinct, % the eye diameter; canthus flat, indistinct; 
loreal sloping, slightly concave ; a thick supratympanic fold from 
lower eyelid to shoulder ; an elongated glandule at the angle of 
the mouth ; first finger longer than second but much shorter than 
third ; subarticular tubercles moderate ; a strong tarsal fold and 
a very pronounced flap from outer metatarsal tubercle to tip of 
the last toe ; other toes similarly fringed and with a short web 
(2, 2, 3, 4, 4) ; inner metatarsal tubercle more protuberant and 
elongated than the outer ; heel of the adpressed hind limb extends 
to middle of the eye. Skin above somewhat rugose, with two 
glandular folds from posterior corner of the eye to inguinal re- 
gion ; flanks with several rounded glandules. Below, smooth. 

Color. Above, vinaceous-brown, the dorsolateral folds and some 
of the lateral glandules of a blackish or dark gray color ; upper 
lip light, vermiculated or spotted with darker ; apparently a 
canthal and a supratympanic streak ; anterior and posterior as- 
pect of the thighs marbled with brown and black, the rest of the 
limbs with obscure dark spots. Below, yellowish-white ; lower lip 
grayish, dotted with white ; probably there were white spots on 
the throat and pectoral region ; distal portion of the hind limbs 
speckled with dark. 

Measurements. 2 snout-vent 70; head breadth 24; head length 
25.5; femur 33; tibia 34.5. 

Range. The Venezuelan Guayana. The species is said to occur 
in British Guiana, Colombia, Brasil and Peru. 

Remarks. I have decided to follow rather uncritically Dr. 
Dunn's determination of this specimen until topotypical material 
is available for comparative purposes. From a glance I took at 



38 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

specimens of L. stictigularis Noble (= L. rhodomystax, fide Par- 
ker, 1935) from British Guiana and at a specimen from Rio 
Uaupes determined as L. rhodomystax in A.M.N.H. (39791), I 
got the impression that this animal is none of them and that if 
the latter is the real rhodomystax then stictigularis is not an 
absolute synonym. The Rio Uaupes form seems to me identical 
with the animal pictured and described by Lutz (1926: 149, pi. 
xxxii) and I do not think it represents L. mystaceus as thought 
by Parker (op. cit.). At least it is much larger than any mys- 
taceus I have seen and the color has a reddish tinge not found in 
that species. The animal here described has very strongly de- 
veloped lateral fringes on the toes, the second finger is only a 
little shorter than, and the third much longer than, the first. I 
noted that the first finger of L. stictigularis is about as long as 
the third and the second is only % the first. The lateral fringes 
are present but not even half as prominent as in the Venezuelan 
animal. 

Leptodactylus bolivianus Boulenger 

L,eptodactylus bolivianus Boulenger, 1898, Ann. Mus. Civ. Stor. Nat. Genova, 
19: 131: Barranca and Misiones Mosetenes, Bolivia; Lutz, A., 1927, 
Mem. Inst. Osw. Cruz, 20: 39, 45, pi. x, figs. 10, 11; Schmidt, K. P., 
1932, Zool. Ser. Field Mus. Nat. Hist., 18: 160; Shreve, 1947, Bull. Mus. 
Comp. Zool., 99: 535; Aleman, 1952, Mem. Soc. Cienc. Nat. LaSalle, 
12: 26. 

Leptodactylus ncellatus Stejneger (not Linne), 1902, Proc. U. S. Nat. Mus., 
24: 180. 

2 (U.P.R. 84, 86) Anaben, Colombia, vi.50. 

1 (U.P.R. 85) Sn. Fdo. de Atabapo, vi.50. 

2 (U.P.R. 87-8) La Culebra, 1000 ft. iv.50. 
4 (M.C.Z. 25985-8) Pauji, iv-xi.45. 

1 (M.C.Z. 26143) Riecito, 29. 
Description. Head usually longer than broad ; snout subellip- 
tical; tongue broad, oval, emarginate behind; vomerine odontoids 
in two arched series behind the choanae, their anterior convexities 
extending sometimes to the middle of the inner margin of the 
latter; eye diameter shorter than, or equal to, distance between 
eye and nostril ; interorbital space equal to an upper eyelid ; 
canthus rounded but distinct; loreal somewhat sloping, concave; 
tympanum large, circular, distinct, % the eye diameter; first 
finger longer than second ; subarticular tubercles of fingers large, 
rounded, of toes, oval or conical ; soles mostly smooth ; toes very 



RIVERO : SALIENTIA OF VENEZUELA 39 

slightly webbed, the web extending to the tips as conspicuous 
lateral fringes; heel of the adpressed hind limb extends to the 
eye. Skin above, smooth, with two dorsolateral folds from upper 
eyelid to groin; a supratympanic fold from posterior corner of 
the eye to shoulder ; also occasionally present is a slight pair of 
lateral folds; an elongated glandular fold at the angle of the 
mouth. Below, smooth, with a discoidal fold on the belly. Arms 
in the breeding male swollen and bearing one or two tubercles 
on the inner side of the first digit. 

Color. Above, dark or light gray, generally marked with dark 
blotches on the posterior half; a black canthal streak which con- 
tinues behind the eye as a temporal spot to the shoulder ; a few 
black blotches sometimes follow this spot on the anterior part of 
the flanks; dorsal folds usually black-edged; between them a 
dark, central marking broadens in front to form an an- 
teriorly truncate spot between the eyes ; tympanum chestnut ; 
upper lip light, sometimes with dark, vertical spots; posterior 
part of the flanks near the inguinal region with black blotches 
and smaller white spots ; posterior aspect of the thighs marbled ; 
rest of the hind limb crossbarred, the bars being black and very 
distinct on the hidden portions of the tibiae. Below, uniformly 
grayish white or slightly marbled with gray; lower lip with 
round, white dots. 

Measurements. Snout-vent $ 72, 9 73 ; head breadth $ 24.5, 
9 24 ; head length $ 25, 9 25.3 ; femur $ 31, 9 31; tibia S 39, 
9 41. 

Habits. Some of the specimens were collected in a partially 
inundated, largely treeless area near the river ; others were found 
in a trail passing through very tall grass about 50 feet from the 
river edge and 25 from the forest. 

Of three stomachs examined, one contained a pentatomid bug, 
a spider and a small beetle. The second had one complete beetle 
and other beetle remains, while the third was empty. 

Robinson (in Stejneger, 1902) describes the breeding habits 
of this frog : ' ' This frog makes a noise like the sound of water 
gurgling from a bottle, only it is a single note and louder. They 
make in the weeds in the water's edge a bird's nest of bubbles or 
more like the whipped-up white of eggs and even more gelatinous. 
The depression in the center goes entirely through and the frog 
sits in the water below, with its nose and eyes appearing in the 
bottom of the nest." 



40 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Additional Localities. Baruta (Aleman, 1952) ; Caracas (Lutz, 
1927) ; nr. Carapa (U.S.N.M. 80613-4) ; Casiquiare, 8 mi. below 
Orinoco (A.M.N.H. 23167) ; Cocollar (Schmidt, 1932) ; Encon- 
trados (C.N.H.M. 2603); Higuerote (U.C.V. 51); La Fria 
(U.M.M.Z. 55582) ; La Guaira (U.S.N.M. 22539, 27793 [= Stej- 
neger, 1902] ) ; La Trinidad (C.N.H.M. 28114, 35988-9) ; Maracay 
(Lutz, 1927) ; Palma Sola (U.M.M.Z. 55581) ; Parmana (U.C.V. 
66) ; Rio Sn. Esteban (U.M.M.Z. 55580) ; Sosa (U.C.V. 80) ; 
Sucre (C.N.H.M. 17769-71 [= Schmidt, 1932?]) ; Turgua (Ale- 
man, 1952) ; Turmero (C.N.H.M. 2734). 

Bange. Probably found in all the physiographical provinces of 
Venezuela. Central America to Peru, Bolivia and Brasil. 

Remarks. Specimens from north of the Andean System seem 
to have a lighter ground color, more blotches on the posterior 
part of the body and flanks and more numerous and distinct 
crossbars on the limbs. In the two Culebra specimens the dorsum 
is somewhat rugose and with small scattered tubercles and the 
ventral surfaces are lightly marbled. No appreciable difference 
has been found between animals from southern Venezuela and a 
topotypical specimen. 

Boulenger describes the male of the species as having a single 
tubercle on the inner finger, apparently a sign of immaturity, 
for sexually mature specimens have two tubercles. 

This species is allied to Lcptodactylus ocellatus from which it 
can be distinguished by its coloration, by having not more than 
two dorsal folds, and by the lesser development of the forelimbs 
in the breeding male. An occasional specimen is found in which 
these characters do not work. The greater breadth of the inter- 
orbital space in L. bolivianus may be useful in these doubtful 
cases. 

A specimen from Colombia deposited at the Museum of Ver- 
tebrate Zooloffv does not seem to have anv dorsal folds. 



'&. 



Leptodactylus mystaceus (Spix) 

Rama mystacea Spix, ]824, Spec. Nov. Test. Ran.: 27, pi. iii, fig. 1, 3: Bahia. 
?Leptodactylus poecilochilus Boettger, 189G, Ber. Senekenb. Naturf. Ges. : 
LIV. 

1 (II.P.R. 91) Anaben, Colombia, vi.50. 

2 (U.P.R. 37-38) Marahuaca, v.50. 

Description. Head broad at the base; snout subovoid, extend- 
ing well beyond the mouth; tongue oval, emarginate behind; 
vomerine odontoids in two slightly arched series behind the 



RIVERO : SALIENTIA OP VENEZUELA 41 

choanae, fairly close together and extending outwards to about 
the middle of the posterior margin of the latter; eyes dorsally 
depressed, their diameter equal to distance between eye and 
nostril ; interorbital space somewhat broader than an upper eye- 
lid ; canthus not very distinct ; loreal oblique, concave ; tympanum 
distinct, % the eye diameter; a supratympanic fold from pos- 
terior corner of eye to shoulder ; an elongated gland at the angle 
of the mouth ; first finger longer than second ; tarsal fold slight, 
metatarsal absent or faintly indicated ; soles slightly tubercular ; 
subarticular tubercles of toes and fingers large, prominent ; toes 
free or with a rudimentary web ; lateral fringes of toes faintly 
indicated ; heel of the adpressed hind limb extends to the eye. 
Skin above and on the flanks and limbs shagreened ; a pair of 
dorsolateral folds from upper eyelid to groin. Belly and throat 
smooth; thighs granular; a ventral discoidal fold. Breeding 
male with a sharp anterior ridge in front of the snout and two 
external vocal sacs indicated by a pair of shallow pigmented 
grooves at the sides of the throat. 

Color. Above, brown, with the dorsal folds of a black or dark 
color on the cephalic and light on the caudal extremities ; a well 
defined black canthal, and supratympanic streaks ; upper lips 
margined with dark; a white "mustache" above the dark line of 
the upper lip extends posteriorly to the shoulder; tympanum 
chestnut; a dark saddle-shaped marking between the eyes; a 
broad, dark brown line along the posterior surface of the fore- 
limbs and a white, slanting one on the anterior side of the 
humerus ; thighs with a dark streak or series of spots along their 
anterior border and a white, dark-edged longitudinal line along 
their hinder aspect; hind limbs with broad crossbars; hidden 
portions of the tibiae with well-defined black spots. Below, white ; 
the throat, chest and limbs sometimes speckled with brown. 

Measurements. Snout-vent S 47, 9 48.5 ; head breadth <$ 17, 
5 17.3; head length $ 17, 9 17 ; femur <J 21, 9 22 ; tibia $ 24, 
9 25.5. 

Habits. Of two stomachs examined, one was empty and the 
other contained a cricket and a seed. 

The female specimen in the collection is full of very large, 
yellow eggs. 

Additional Localities. Boettger's, 1896, L. poecilochilus from 
Upper Orinoco may refer to this species. 

Range. The Venezuelan Guayana. Southeastern Colombia and 
British Guiana to southern Brasil. 



42 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Remarks. Specimens from Rio Humboldt, Brasil and Buena- 
vista, Bolivia, do not show any appreciable difference from the 
Venezuelan animals. 

The species is related to L. poecilochilus from which it can be 
distinguished by the different coloration of the snout and its 
tubercular soles. The white coloration of the loreal area is more 
extensive dorsally in L. poecilochilus and not narrow and well 
defined as in L. mystaceus. This "mustache" is not interrupted 
by spots or lines in L. mystaceus. 

Leptodactylus poecilochilus dypticus Boulenger 

Lepto dactyl us dypticus Boulenger, 1918, Ann. Mag. Nat. Hist., (9) 2: 431: 
Andes of Venezuela; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 39, 45, 
pi. x, figs. 12, 13; Shreve, 1947, Bull. Mus. Comp. Zool., 99: 535. 

2 (M.C.Z. 25989-90) Pauji, iv, xi.44. 

1 (U.M.M.Z. 55554) Palma Sola. 

1 (U.M.M.Z. 57483) Sabana de Mendoza. 

1 (U.M.M.Z. 57484) Guachi. 
Description. Head broad at the base; snout subovoid, longer 
than the eye diameter ; tongue oval, emarginate behind ; vomerine 
odontoids in two long, slightly arched and close-together series 
behind the choanae, their anterior convexities not extending be- 
yond the horizontal of the posterior margin of the latter ; eye 
diameter equal or slightly greater than distance between eye and 
nostril ; interorbital space equal to or narrower than an upper 
eyelid ; canthus indistinct ; loreal sloping, slightly concave ; tym- 
panum distinct, % the eye diameter ; a supratympanic fold from 
lower eyelid to shoulder ; an elongated glandule at the angle of 
the mouth ; first finger much longer than second ; fingers and 
toes distally swollen ; tarsal fold not distinct, metatarsal slightly 
indicated ; inner metatarsal tubercle small, subarticular tubercles 
large and very prominent ; soles usually smooth or nearly so ; toes 
free, without lateral fringes; heel of the adpressed hind limb 
extends to between eye and nostril. Skin above, smooth, with 
two dorsolateral folds that extend from upper eyelid to groin ; 
another pair of loose folds along each side of the body; flanks 
with small, scattered warts. Below, smooth, with a discoidal fold 
on the belly. 

Color. Above, gray or light brown, with a pair of dark, elon- 
gated spots behind the scapulae, at the edge of the dorsolateral 
folds ; a distinct, black canthal streak that does not extend to 
the eye ; a black supratympanic streak ; a black, broad, anterior 
and a narrow posterior line from eye to dark-margined upper 



RIVERO : SALIENTIA OF VENEZUELA 43 

lip; tympanum chestnut; interorbital space with a narrow trans- 
verse line ; dorsolateral folds light colored ; forelimbs crossbarred ; 
thighs with a broad dark streak or series of large spots along 
the anterior border, their posterior aspect with a white, dark- 
edged, longitudinal line ; sometimes the thighs are closely cross- 
barred above ; hidden portions of the tibiae with very distinct 
black bars. Below, milky white ; lower lip edged with a dark 
grayish line which disappears anteriorly. 

Measurements. 9 Snout-vent 45 ; head breadth 16 ; head 
length 16; femur 12; tibia 14. 

Additional Localities. Maracay (Lutz, 1927). 

Range. The Coastal Range, the Falcon Region, the Maracaibo 
Basin and the lower slopes of the Western Andes. Northeastern 
Colombia. 

Remarks. The number of specimens available have been insuf- 
ficient to determine whether the Venezuelan form is really 
identical with the Colombian-Central American poecilochilus. 
They clearly show, however, that the two are very closely related 
and probably conspecific. 

Four of the five Costa Rican frogs examined are of the "quat- 
rilineatus" type which is said to be the least common within the 
range of L. p. poecilochilus (Dunn, 1940: 106). M.C.Z. 7997 from 
the same locality and collection as the "quatrilineatus" group is 
mostly plain above and similar in many respects to M.C.Z. 16069 
from Rio Frio, Colombia and to the Venezuelan animals, for 
which the striped coloration has never been reported. Apart 
from this, little is left to separate the two forms. It appears that 
L. p. poecilochilus is more spotted on the dorsum, has better de- 
fined spots between upper lip and eye and a broader white line 
along the posterior aspect of the thighs. The canthal streak seems 
to extend to the eye in most poecilochilus while in dypticus it 
stops before reaching the orbit. It would be necessary to have 
more non-" quatrilineatus" specimens to determine if these dif- 
ferences really exist and if there are any others. 

M.C.Z. 25989 has a rosy tinge on the flanks and thighs and 
both 25989 and 25990 are of a light gray color, differing in this 
respect from the U.M.M.Z. specimens, in which the color is brown. 
This may be due to differences in the preservatives used. In 
25990 the transverse bars of the thighs are narrow and numerous 
while in the other specimens they are indistinct or absent. 

The male L. p. dypticus from Rio Frio, Colombia (37.5 mm.) 
has a very pronounced ridge in front of the snout. U.M.M.Z. 



44 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

57484 looks somewhat different on account of its unusually nar- 
row head but it agrees in other characters. 

In the files of the American Museum of Natural History I 
noticed that there are one or more specimens from Medellin, 
( !olombia, determined as L. dypticus, but probably representing 
the typical form. I believe the Argentinian specimens mentioned 
by Dunn (op. cit.) refer to some other species, perhaps L. mys- 
taceus. 

Leptodactylus sibilatrix (Wied) 

Sana sibilatrix Wied, 1824, Abbild. Naturg. Bras., No. 4: 5, pi. 5, fig. 2: 

East Coast of Brazil. 
Cystignathus typhonius Lichtenstein and Martens, 1856, Nomenel. Rept. 

Amphib. Mus. Berol.: 39. 
Leptodactylus mystacinus Boettger (part: not Burmeister), 1892, Kat. 

Batr. Samm. Mus. Senckenb.: 30; Nieden (part?), 1923, Das Tierreich. 

Anura I: 485; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 39, 45. 
Leptodactylus typhonius Boettger, 1892, Kat. Batr. Samm. Mus. Senckenb. : 

31; 1896, Ber. Senckenb. Naturf. Ges.: LIV; Lutz, A., Mem. Inst. Osw. 

Cruz, 20 : 39, 46. 

9 (U.P.R. 92-100) Pto. Ayacucho, iii, vi.50. 

Description. Head longer than broad ; snout acuminate, pro- 
jecting well beyond the mouth; tongue oval, emarginate behind; 
vomerine odontoids in two slightly arched series close to- 
gether behind the choanae ; eye diameter equal or slightly greater 
than distance between eye and nostril ; interorbital space more 
or less equal to an upper eyelid ; canthus absent ; loreal sloping, 
slightly concave; tympanum distinct, % the eye diameter; a 
slight supratympanic fold from lower eyelid to shoulder ; an 
elongated gland at the angle of the mouth ; first finger longer than 
second ; tarsal fold usually slight ; metatarsal fold may be faintly 
indicated ; soles with minute, flat granules ; toes free, their lateral 
fringes absent or faintly indicated in very dry specimens; heel 
of the adpressed hind limb extends to the eye. Skin above, 
smooth, with several (usually more than 5) dorsal, longitudinal 
folds. Below, smooth, with a distinct discoidal fold on the belly. 
Male with two lateral external vocal sacs; arms not enlarged; 
inner side of the first digit without tubercles. 

Color. Above, brown or tan, spotted with darker brown mark- 
ings ; one or two spots generally present between the eyes ; tym- 
panum chestnut, with a white rim; limbs crossbarred; posterior 
surface of the thighs marbled, but with a well defined, longitudi- 
nal, white line or continuous series of dots along its lower part. 
Belly milky white ; vocal sacs pigmented with dark gray or black. 



RIVERO : SALIENTIA OF VENEZUELA 45 

Measurements. Snout -vent <$ 42.5, 9 39.5; head breadth 
6 14, 9 13.1 ; head length $ 15, 9 15; femur $ 18.2, 9 18.5; 
tibia $ 24, 9 22. 

Habits. Several specimens were collected at night, after heavy 
rain, in the shallow pools of a grassy (2 or 3 inches grass) area. 
Others were collected in tall grass while calling at night. The 
voice is probably a short "huit" sound, as described by Lutz 
for L. mystaceus. 

Additional Localities. Arabopo (U.M.M.Z. 39757, 85201 [5], 
85205) ; Caracas (Senckenb. Mus. 1244b [= Boettger, 1892: 39, 
part]); Chicare (Caicara?) (U.S.N.M. 36369-70); Espino 
(U.C.V. 69) ; 10 km. from La Pascua (U.S.N.M. 128839) ; Mara- 
cay (Lutz, 1927) ; Pie del Cerro (U.S.N.M. 121148) ; Roraima 
(A.M.N.H. 39752) ; Sabana de Mendoza (U.M.M.Z. 75478) ; Sta. 
Elena, Bolivar (U.M.N.Z. 85204) ; Sosa (U.C.V. 86) ; Upper Ori- 
noco (Boettger, 1896) ; Wiaka Piapu, 5000 ft. (U.M.M.Z. 
85200[2]). 

Range. The Maracaibo Basin, the Coastal Range, the Llanos 
and the Venezuelan Guayana. Northern Colombia to northern 
Argentina. 

Remarks. Specimens from Sabana de Mendoza are of a much 
darker ground color ; the spots are rather obscure and the dorsal 
folds are absent or reduced to two slight ones. Animals from 
Sta. Marta, Colombia, are not very different from those of the 
rest of Venezuela, but a lot from Magdalena Valley, Colombia, 
have a somewhat rugose skin and as dark a color as the specimens 
from Sabana de Mendoza. These last animals probably represent 
a subspecies. 

L. sibilatrix can be confused with young L. act 11 at us. from 
which it can be distinguished by its more projecting and acumi- 
nate snout, by the absence of white dots on the lower lip and of 
lateral fringes on the toes and by the presence of a longitudinal 
white line on the posterior part of the thighs. The breeding male 
of L. occllatus has enlarged forearms and two tubercles on the 
inner side of the first digit but no apparent external vocal sacs. 

Leptodactylus ocellatus (Linne) 

liana ocellata Linne, 1758, Syst. Nat., ed. 10: 211: America. 

Cystignathus occllatus Lichtenstein and Martens, 1856, Nomencl. Sept. Am- 

phib. Mus. Berol. : 39. 
Leptodactylus occllatus Boettger, 1893, Ber. Senckenb. Naturf. Ges.: 40; 

Lutz, A., 1927, Mem. Inst, Osw. Cruz, 20: 45. 



46 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

2 (U.M.M.Z. 83718) Sn. Fdo. de Apure. 

1 (U.M.M.Z. 89583) Rio Araana, nr. Maturin. 

1 (U.C.V. 11) Parmana, vi.51. 

4 (U.C.V. 31-33, 35) El Sombrero, viii.51. 

Description. Head longer than broad; snout subelliptical ; 
tongue broad, oval, emarginate behind ; vomerine odontoids in two 
arched series behind the choanae, their anterior convexities ex- 
tending sometimes to the middle of the inner margin of the latter ; 
eye diameter equal to distance between eye and nostril ; inter- 
orbital space narrower than an upper eyelid ; canthus absent ; 
loreal somewhat sloping, concave ; tympanum distinct, % the eye 
diameter ; a supratympanic fold from posterior corner of eye to 
shoulder and a thick glandular ridge from this to the angle of 
the mouth ; first finger longer than second ; tarsal fold strong, 
metatarsal indistinct or absent ; soles mostly smooth ; subarticular 
tubercles of fingers large and rounded, of toes, conical ; toes 
slightly webbed, the web extending to the tips as conspicuous 
lateral fringes ; heel of the adpressed hind limb extends to the 
eye. Skin above, smooth, with several (usually more than 5) 
longitudinal folds; flanks sometimes with small, rounded glan- 
dules. Below, smooth, with a discoidal fold on the belly. Arms 
in the breeding male much swollen and bearing two blunt tu- 
bercles on the inner side of the first digit. 

Color. Above, light brown spotted with a darker shade of 
brown ; occasionally some of the spots are elongate and follow the 
dorsal plicae for part of their length ; a usually symmetrical spot 
between the eyes ; a canthal and a temporal streak ; thighs marbled 
behind : hind limbs crossbarred. Belly and throat white, uni- 
form, mottled or dusted with brown ; a series of white dots along 
the margin of the brown pigmented lower jaw. Throat may have 
white spots on a brownish background. 

Measurements. Snout-vent $ 71.5, 9 60 ; head breadth $ 25.5, 
9 21 ; head length $ 24, 9 22 ; femur $ 31, 9 25; tibia S 34, 
9 28.5. 

Additional Localities. Puerto Cabello (Boettger, 1893) ; Ven- 
ezuela (Licht. and Mart., 1856). 

Range. The coastal belt and the Llanos. So far as 1 am aware, 
the records given here represent the most northerly for the 
species. To the south it extends as far down as Argentina, where 
a local race (L. occllatus bonariensis) is recognized by Cei (1956) . 

Remarks. See remarks under L. sibilatrix and L. bolivianus. 



RIVERO : SALIENTIA OF VENEZUELA 47 

Leptodactylus podicipinus petersii (Steindachner) 

Platymantis petersii Steindachner, 1864, Verhandl. Zool. Ges. Wien.: 254, 

pi. xvi, figs. 2, 2a, 2c: Marabitanas, Brasil. 
Leptodactylus caliginosus Boulenger (not Girard), 1903, Ann. Mag. Nat. 
Hist., (7) 11: 481; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 39, 45, 
pi. xi, figs. 14, 15 ; Shreve, 1947, Bull. Mus. Comp. Zool., 99 : 536. 
1 (U.P.R. 101) Sn. Fdo. de Atabapo, vi.50. 
6 (M.C.Z. 26144-6 --3 dupl.) Cerro Cosme, x.30. 
1 (M.C.Z. 2970) Merida. 

1 (U.M.M.Z. 55551) Lagoon Palma Sola. 

2 (U.M.M.Z. 55552-3) La Fria, rt. fork R. Oropito. 
Descriptio?i. Head broader than long ; snout subovoid ; tongue 

oval, slightly nicked behind; vomerine odontoids in two little 
arched series behind the choanae, their anterior convexities not 
extending beyond the horizontal of the posterior margin of the 
latter ; eyes small, their diameter equal to distance between eye 
and nostril ; interorbital space more or less equal to an upper 
eyelid ; canthus indistinct ; loreal sloping, not concave ; tympanum 
circular, % the eye diameter ; first finger longer than the second ; 
subarticular tubercles moderate; a strong tarsal and a distinct 
metatarsal fold ; metatarsal tubercles small but distinct ; soles 
usually tubercular; toes slightly webbed, the web extending to 
the tips as very conspicuous lateral fringes ; heel of the adpressed 
hind limb extends to the posterior corner of the eye or a little 
beyond. Skin above, with small tubercles on the posterior % ; 
upper eyelids usually granular ; flanks sometimes with round and 
elongated glandules. Ventral surfaces of the belly and throat 
smooth ; an indistinct discoidal fold on the belly. Breeding male 
with two pointed tubercles on the inner side of the first digit; 
no distinct external vocal sac. 

Color. Above, grayish brown with occasional darker spots ; a 
dark, anteriorly truncate spot between the eyes is generally lim- 
ited in front by a light transverse line ; snout lighter than the 
body color ; a light, not very distinct, slanting line from lower 
eyelid to angle of jaw and usually some others in front of these ; 
hind part of the thighs obscurely marbled with light ; a not very 
well defined light line along the lower posterior aspect of thighs ; 
limbs crossbarred, sometimes indistinctly. Ventral surfaces dirty 
white, sometimes marbled or freckled with brown on the throat, 
breast and distal portions of the hind limbs; lower lip usually 
margined with white dots. 



48 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

Measurements. Snout-vent $ 37, 9 41 ; head breadth $ 13, 
9 14 ; head length $ 12.5, 9 14 ; femur $ 18, 9 17 ; tibia 6 20, 
9 18.5. 

Additional Localities. Albarregas River (A.M.N.H. 10517-9) ; 
Arabopo (A.M.N.H. 39758-9; U.M.M.Z. 85197[4]) ; Bonicito 
River (U.M.M.Z. 100056) ; Carayaca (U.C.V. 17) ; Caripito 
(U.S.N.M. 17088-9) ; Casiquiare, 8 mi. above Capibara (A.M.N.H. 
23166) ; El Limon (U.S.N.M. 121146) ; Los Canales, Planta 
Electrica de Naiguata (U.S.N.M. 128838); ?Maracay (Lutz, 
1927); Merida (A.M.N.H. 3136; U.S.N.M. 118176; Boulenger, 
1903) : Petare (U.S.N.M. 121147) ; Pie de Avila (U.C.V. 110) ; 
Pto. Ayacucho, nr. Venado (U.S.N.M. 80666, 80671-5) ; Rio 
Cotiza, Camino de Galipan (U.S.N.M. 117526, 121137) ; Rio 
Pescado (A.M.N.H. 23182); Roraima (A.M.N.H. 39753). 

Range. All the physiographical provinces of Venezuela, in- 
cluding the Andes to an elevation of at least 1600 m. The 
species extends from Colombia and Trinidad to Buenos Aires. 
The race is found in the northern part of the continent and as 
far south as Bolivia and Minas Gerais. 

Remarks. Dr. Cochran has found (1955:328) that Leptodac- 
tylus caliginosus Girard is a strict synonym of Leptodactylns 
ocellatus (Linne). Specimens from southern Brasil referred to 
caliginosus by various authors are considered by her to be identi- 
cal with L. podicipinus Cope. Such being the case, the name 
available for the northern form which has been called L. calig- 
inosus seems to be Leptodactylns petersii (Steindachner). 

For the study of podicipinus podicipinus and podicipinus 
petersii, I have examined a total of 155 specimens from Colombia, 
Venezuela, the Guianas, Brasil, Bolivia and Ecuador, and for 
comparison with these I had a few examples of melanonotus from 
Central America and validus from Trinidad. 

The change from petersii to podicipinus seems to occur in Bo- 
livia on the west and in Minas Gerais on the east, but the nature 
of the change and the direction it follows is not easy to see without 
resort to statistical measures. There are sufficient specimens in 
the different museums and it is hoped that this type of analysis 
can be done in the near future. Meanwhile, the following observa- 
tions may be helpful. L. p. podicipinus differs from L. p. petersii 
in its smaller size (average for 15 adult specimens, 10 9 9,5 
i £ , 32.2 for L. p. petersii and 29.5 for L. p. podicipinus) , in 
having a shorter femur (the length of the femur equals half the 
distance from anus to interorbital line in podicipinus and from 



RIVERO: SALIENTIA OF VENEZUELA 49 

anus to the middle of the snout [Bolivia] or beyond the tip in 
petersii), in having an interorbital spot that extends much farther 
back on the dorsum, in the characteristic white spotting on the 
venter and in lacking a longitudinal, white line along the pos- 
terior part of the thighs and a well defined supratympanic fold. 
L. p. petersii has a broader head and usually transverse bars on 
the upper surface of the hind limbs and brown freckles on the 
distal lower surface. 

In Bolivia, typical L. p. podicipinus occurs in Cachuela Esper- 
anza and Rurrembaque but in the Upper Beni (farther south!) 
it occurs as a larger form, many specimens lose the ventral spots, 
and the longitudinal, white line in back of the thighs comes out. 
There is a form in Buena Vista which I think should be referred 
to the northern race though somewhat divergent from it and 
possessing certain characters of its own (long, narrow head; gray 
color; 2 black spots on the sides of the anus). It seems to me 
that the change from one race to the other is not along a well- 
defined latitudinal boundary ; instead there are many finger-like 
projections of one race into the "supposed" range of the other. 
Along the eastern base of the Andes and along the lower course 
of Rio Beni, typical podicipinus occurs. However, the more sou- 
thern Upper Beni specimens show some transitional characters 
toward the eastern variant occurring in Buena Vista and the 
latter toward the northern form, which probably meets podicip- 
inus on the east and north of this region. In Maracaju, Salobra 
and Corumbo in Matto Grosso only podicipinus seems to occur, 
while in Minas Gerais both forms exist. L. natalensis should per- 
haps by synonymized with L. p. petersii. 

In the north, L. p. petersii has undergone considerable modifi- 
cation in some places. In Colombia, for example, the species 
seems to attain enormous (relatively speaking) proportions, while 
in the Upper Rupununi of British Guiana, specimens with very 
dark dorsal coloration and densely reticulated ventral surfaces 
occur. This ventral pattern is found in specimens from other 
places but in this region it is very distinct and apparently gen- 
eral (A.M.N.H. 46437, 27 untagged specimens). 

Leptodactylus validus seems to me referable to this same species 
and perhaps even Leptodactylus melanonotus may be included in 
the Rassenkreis but these problems are left until a complete 
study can be made. 

Aleman's, 1952, Leptodactylus species off. petersii from Turgua 
may be referable to this form. 



50 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Leptodactylus bugosus Noble 

Leptodaotylus rugosus Noble, 1923, Zoologica (N.Y.), 3: 297: British 
Guiana; Parker, 1936, Bull. Mus. Boy. Hist. Nat. Belgique, 12: 2. 
7 (U.P.R. 190-1, 62-6) Cerro Turuma Chica, vi.50. 

6 (U.P.R. 53-4, 67-70) Pto. Ayacucho, iii.50. 

7 (U.P.R. 55-61) Sanariapo, iii.50. 

5 (U.P.R. 71-5) Casa de Julian, iv.50. 

Description. Snout subovoid, longer than the eye diameter; 
tongue oval, entire or very slightly nicked behind ; vomerine 
odontoids close together and forming two arched groups behind 
the choanae, their anterior convexities usually extending to the 
middle of the inner margin of the latter; eye diameter greater 
than distance between eye and nostril ; interorbital space nar- 
rower than an upper eyelid; canthus depressed; loreal sloping, 
concave; tympanum distinct, circular, % the eye diameter; a 
supratympanic fold from posterior corner of eye to shoulder ; one 
or two glandular tubercles between angle of jaw and arm ; first 
finger longer than second ; fingers and toes distally swollen ; 
tarsal fold slight to moderate, metatarsal absent ; an elongate 
oval inner and a small, rounded, outer metatarsal tubercle ; toes 
practically free, without lateral fringes; heel of the adpressed 
hind limb extends to the posterior corner of the eye. Skin above 
with irregularly scattered warts of unequal size ; flanks usually 
with numerous round or elongated glandules. Below, smooth, 
except for the hind part of the thighs and sides of the belly 
which are grandular ; a ventral discoidal fold. Breeding male 
with a pair of spines on the breast, two pigmented external vocal 
sacs, numerous spinules on the chin and a pointed tubercle on 
the inner side of the first digit. 

Color. Above, black or brownish gray, sometimes obscurely 
spotted with a darker shade ; several light colored lines radiate 
from eye to upper lip ; a distinctive, white interorbital line and 
occasionally a pair of vermicular, whitish ones originate in the 
posterior corner of the eye and converge posteriorly to separate 
again and form an hour-glass figure on the anterior part of the 
dorsum; hinder part of the thighs obscurely marbled; limbs 
crossbarred, the bars not clearly delimited in some specimens. 
Belly and lower surfaces of limbs white or brownish, uniform or 
somewhat marbled ; throat generally infuscate in the male. 

Measurements. Snout-vent S 67, 9 67 ; head breadth $ 25, 
? 25 ; head length S 24, 9 22.5 ; femur $ 31, 9 29 ; tibia S 30, 
9 31.5. 



RIVERO : SALIENTIA OP VENEZUELA 51 

Habits. The voice of this species is very similar to the noise 
produced when a stone is thrown into a quiet pool of water. It 
is usually heard at dusk in the stony places which the animal 
generally inhabits. Many of the specimens were collected under 
a large rock on top of a hill of bare granite (Turuma Chica) 
where the dampness and the small, stingless bees that nested 
under the stone provided a clear motive for their site preference. 
All the stomachs of these specimens and of some of those col- 
lected elsewhere were full of these bees. Evidently this is not 
their only food, one of the stomachs also having revealed a pair 
of beetle elytra. 

In Puerto Ayacucho most of the specimens were also found 
hiding under granitic rocks or in the proximity of them and the 
same is true for the small specimens collected in Casa de Julian. 
Here, a small, swift stream runs for about 100 yards over a bed 
of solid granite. The specimens, together with a few long-tailed 
tadpoles that very probably belong to this species, were found 
under a loose stone at the edge of the stream. Very little water 
reached the stone and the tadpoles seemed to have some difficulty 
in w r riggling about, but stones in deeper water or in muddy 
places nearby revealed no amphibian life. 

The material from Sanariapo comes mostly from the edge of 
the river (collected at night) with granite rock at not more than 
a few hundred feet away. A few small and probably recently 
metamorphosed specimens were seen at a short distance from this 
place on an extensive "laja" of black bare granite that contained 
water in many of its holes and depressions. Crustaceans and 
mosquito larvae were also collected in these pools. 

I have the impression that Leptodactylus rugosus is very much 
restricted to black, crystallized rock and that it will not be found 
outside of the Guayana Highlands. 

Additional Localities. Auyantepui (A.M.N.H. 46038-9) ; Bor- 
der between British Guiana and Venezuela (U.M.M.Z. 85211) ; 
Canal Casiquiare, between Colombia and Venezuelan border 
(A.M.N.H. 23160-2) ; Puerto Ayacucho, near Venado (U.S.N.M. 
80664-73) ; Puerto Ayacucho (A.M.N.H. 23209-19) ; Sanariapo 
(U.S.N.M. 80035-8) ; Upper Orinoco (Parker, 1936) ; 20 mi. west 
of Wiaka Piapu Mts. (U.M.M.Z. 85158 [2]) ; Wiaka Piapu Mts. 
(U.M.M.Z. 85209 [2]). 

Range. The Venezuelan Guayana. British Guiana, southwest- 
ern Colombia and probably northern Brasil. 



52 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Remarks Examination of the paratype (A.M.N.H. 3790), now 
in M.C.Z., and later of the type, has revealed close morphological 
agreement with the animals reported here, although probably 
due to the preservative used by the collector, the color is, as 
described by Noble, reddish brown. His specimens were 16.5 to 
41 mm. in snout-vent length. 

In reporting on an Upper Orinoco collection in the Brussels 
Museum, Parker (1936:2) identified three specimens as L. rugo- 
sus but pointed out that both males in the collection had single 
nuptial spines while the female was distinctly larger than Noble 's 
(49 mm.). Noble's specimens did not have nuptial spines. 

Leptodactyhis rugosus Melin, 1941, is preoccupied by L. rngo- 
sus Noble. For Melin 's form, Lutz and Kloss have created the 
name L. melini. I regard this form as a synonym of L. marmora- 
tus hylaedactylns (see page 35). 

Lithodytes lineatus (Schneider) 

Bana Lineata Schneider, 1799, Hist. Amphib., 1: 138. 

1 (U.P.R. 104) La Culebra, 1000 ft,, vi.50. 

Description. Physiognomy of Dendrobates; snout somewhat 
truncate ; tongue oval, entire and free behind ; vomerine odon- 
toids in two short, slightly arched series behind the choanae. 
their anterior convexities extending to the middle of the inner 
margin of the latter ; interorbital space slightly broader than 
an upper eyelid ; tympanum almost as large as the eye ; canthus 
not distinct, rounded ; loreal nearly vertical, not distinctly con- 
cave ; first finger longer than second ; tips of fingers and toes 
distally swollen; a moderate tarsal and a metatarsal fold; toes 
free, with an indication of a lateral fringe ; heel of the ad pressed 
hind limb extends to the posterior corner of the eye. Skin above 
and on sides of the head, lower lip and limbs studded with regu- 
larly distributed tubercles ; on the thighs the tubercles are limited 
to a central longitudinal band. Ventral surfaces, smooth. 

Color. Above, dark grayish-brown with two longitudinal 
stripes that begin at the tip of the snout and end in the groins ; 
several red spots on the thighs, hidden portions of the tibiae, 
tarsii and feet, Below, grayish, with round white spots on the 
belly and throat. 

Measurements. $ snout-vent 45.5 ; head breadth 8.5 ; head 
length 8.5 ; femur 13.5 ; tibia 9.5. 

Habits. The only specimen was collected at night at the edge 
of a savanna, more toward the forest side than to the grassland. 



RIVERO : SALIENTIA OF VENEZUELA 53 

Range. The Venezuelan Guayana. Colombia (Rio Garagoa), 
Trinidad, British Guiana, Ecuador and probably northern Brasil. 

Remarks. I have examined very large specimens of this 
species (50 mm.) and all of them have T-shaped terminal phal- 
anges. 

There is practically no difference between animals from Trini- 
dad, British Guiana, Venezuela and Ecuador. 

Key to the Species of Eleutherodactylus Reported from 

Venezuela 

I. First finger longer than second. 

A. Head and mouth very large; dorsum tubercular and ridged 

cornutus maussi 

II. First ringer equal to, or shorter than second. 

A. Tympanum % to % the eye diameter. 

1. First finger shorter than second. 

a. Margins of the frontoparietals elevated ; skin above smooth ; 
vomerine teeth oblique briccni 

b. Margins of the frontoparietals not elevated; skin above 

finely granular; vomerine teeth transverse 

turumiquirensis 

2. First finger equal to second. 

a. Heel of the adpressed hind limb extends to the nostril or 
beyond terraebolivaris 

b. Heel of the adpressed hind limb does not reach the nostril. 

1. Thigh y 2 the length between anus and anterior corner 
of the upper eyelid; eye diameter shorter than distance 
between eye and nostril; belly granular. . . . brachypodius 

2. Thigh Y2 the length between anus and middle or tip of 
the snout; eye diameter equal to distance between eye 

and nostril; belly smooth or granular 

conspioillatus ileamazonicus 

B. Tympanum not more than % the eye diameter. 

1. Tympanum small and indistinct; disks small, narrow, more or 
less pointed stenodiscus 

2. Tympanum distinct, even if small; disks not pointed. 

a. A brown or black lateral band. 

1. Eye diameter greater than distance between eye and 
nostril orocostalis 

2. Eye diameter not greater than distance between eye 
and nostril bicumuhis 

b. No lateral band. 

1. A white (red?) area at the groin and another under the 
arm williamsi 



54 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

2. Not as above. 

a. Eye diameter greater than distance between eye and 
nostril. 

(1) Anterior and posterodistal portions of thighs 
stained with orange brown; a W-shaped scapu- 
lar marking rozci 

(2) Not as above orocostalis 

b. Eye diameter not greater than distance between eye 
and nostril. 

(1) Disks not broader than the tympanum; thighs 
usually tinged with red urichi 

(2) Disks broader than the tympanum; thighs not 
tinged with red. 

a. Heel extending to the nostril or beyond .... 
reticulatus 

b. Heel not extending to the nostril. 

(i) Thighs with white spots and/or bars; 36 

mm.; Coastal Eange racenisi 

(ii) Thighs without white spots or bars; 15-22 

nun.; Venezuelan Guayana. .marmoratus 

C. Tympanum absent 1 anotis 

Eleutherodactylus cornutus maussi (Boettger) 

Hylodes maussi Boettger, 1893, Ber. Senckenb. Naturf. Ges. : 39: Puerto 
Cabello, "Venezuela; Nieden, 1923, Das Tierreich. Anura I: 466; Lutz, 
A., 1927, Mem. Inst. Osw. Cruz, 20: 40. 

1 (M.C.Z. 10178, juv.) Pto. Cabello, 1895. 

1 (U.M.M.Z. 56774, juv.) Sn. Esteban. 

1 (U.C.V. 38) Guamitas, iii.51. 

7 (U.C.V. 19, 106, 113, 116-8, 1159) Rancho Grande, 49-51. 
Description. Head large ; snout broad, subovoid, sloping to- 
ward the front; vomerine odontoids forming two slightly arched 
groups behind the choanae, their anterior convexities touching 
the posterior margin of the latter; tongue rounded, entire and 
Y 3 free ; interorbital space more or less as broad as an upper 
eyelid ; margins of the frontoparietals prominent ; eye diameter 
equal or slightly shorter than distance between eye and nostril ; 
canthus obtusel.v angular ; loreal strongly sloping, concave ; tym- 
panum distinct, % the eye diameter ; a row of four or more 
tubercles along the forearm ; metacarpal tubercles large, sub- 
articulars very large, rounded and prominent, especially on the 
first finger; fingers free but with thick lateral fringes, including 

i Present, l>tit small and indistinct in young specimens, which can be dis- 
tinguished by having red on the venter and/or limits. 



RIVERO: SALIENTIA OF VENEZUELA 55 

the outside of the first; first finger longer than second, which is 
more or Jess equal to fourth ; digits distally dilated ; a tarsal fold ; 
an indistinct, outer and a small, elongated, inner metatarsal 
tubercle ; subarticular tubercles of toes large, conical and very 
prominent ; disks of toes small but distinct ; heel of the adpressed 
hind limb extends to between eye and nostril. Skin above and on 
the sides of the head and flanks tubercular ; upper eyelid tubercu- 
lar, one of the posterior tubercles being longer than the others; 
a pair of longitudinal, curved ridges from behind the upper 
eyelid to the post-scapular region ; usually a cross ridge or row 
of tubercles unite the two longitudinal ridges in the middle, 
forming with them a ) — ( figure; behind this figure, two other 
ridges, usually less prominent and closer together occur ; another 
very short, oblique ridge or series of tubercles at the angle of the 
mouth ; on the limbs some rows of tubercles fuse to form oblique 
ridges. Throat, belly and thighs thickly granular. 

Color. Dark gray (pinkish in life), the ridges and some of the 
tubercles being sometimes of a darker color; a light line from 
lower eyelid to near the angle of the mouth is generally followed 
anteriorly by a triangular darker spot, another light line and 
another dark spot ; limbs sometimes crossbarred ; posterior part 
of the thighs of the same color as the body. Below, dark brownish 
gray, the granules being lighter than the ground color ; throat and 
limbs usually darker than the belly. 

Measurements. 9 snout-vent 59.5; head breadth 24; head 
length 19; femur 26; tibia 26.5. 

Habits. In Venezuela this appears to be mainly a subtropical 
forest form. 

Additional Localities. Puerto Cabello (Nieden, 1923; Lutz, 
1927 [cited from Boettger, 1893]). 

Range. The Coastal Range and probably the Andes. The 
species extends from Colombia to Ecuador. 

Remarks. U.C.V. 38 has a much lighter color than the Rancho 
Grande specimens, the limbs are lighter than the body color, and 
the venter is plain white. 

Topotypical specimens of E. cornutus cornutus (M.C.Z. 19640- 
2; Canelos, Ecuador) have a more depressed snout than maussi; 
the dorsum occasionally has extensive dark spots (M.C.Z. 19642), 
the posterior part of the thighs is black with light pinkish (red?) 
spots, and the groins, anteroventral and ventrodistal portions of 
the thighs are equally spotted or more broadly marbled ; there 
are large pinkish spots on the hidden portions of the tibia and 



56 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

tarsus and in specimen 19642, one of the lines that radiates from 
eye to lip is broad and very distinct. 

It is very possible that Hylodes sulcatus Cope, 1874, from 
Nanta, Peru and Ctenocranius koki Melin, 1941, from Taracua, 
Brasil, are conspecific with Eleiitherodactylus cornutus (Es- 
pada), 1870, from San Jose de Moti, Ecuador. The first, however, 
is described as having an areolate belly. 

If the large-headed Eleiitherodactylus of the comutus-bipor- 
catus group is found to constitute a genus, the name Straboman- 
tis Peters, 1863, will probably be the available name while 
Limnophys Espada, 1870, and Ctenocranius Melin, 1941, will be- 
come synonyms. So far, I do not think that the characters men- 
tioned by Melin are sufficient to separate the two groups. 

Eleutherodactylus briceni (Boulenger) 

Hylodes briceni Boulenger, 1903, Ann. Mag. Nat. Hist., (7) 11: 481: Merida, 
Venezuela; Nieden, 1923, Das Tierreich. Anura I: 466; Lutz, A., 1927, 
Mem. Inst. Osw. Cruz, 20 : 39. 

2 (M.C.Z. 3888, 7601, cotypes) Merida. 

Description. Snout subovoid ; tongue broadly oval, entire or in- 
distinctly nicked and free behind ; vomerine odontoids forming 
two short, oblique and close together series behind the choanae, 
converging posteriorly and not extending outwards beyond the 
vertical of the inner margin of the latter; interorbital space 
slightly concave, broader than the upper eyelid; margin of the 
frontoparietals slightly elevated ; eye diameter equal to distance 
between eye and nostril ; canthus distinct, somewhat curved ; 
loreal sloping, concave; tympanum small, not very distinct, 
about !/2 the eye diameter; a pair of indistinct metacarpal tu- 
bercles ; subarticular tubercles moderate, round ; first finger 
shorter than second ; disks small, truncate, equal or slightly 
smaller than the tympanum ; disks of the toes smaller than those 
of the fingers ; an indistinct outer, and an oval, inner metatarsal 
tubercle; toes free; heel of the adpressed hind limb extends to 
the anterior corner of the eye. Skin above, smooth. Below, smooth 
in the throat, breast and central part of the belly; sides of the 
belly sometimes with flat, irregular sized granules. 

Color. Above, brown with darker, sometimes cbevron-shaped 
markings ; an interorbital band ; dark, narrow canthal and supra- 
tympanic streaks; flanks brown with darker reticulations or light 
centered, dark brown spots; limbs diffusely crossbarred. Below 
whitish, reticulated or marbled with brown. 



RIVERO : SALIENTIA OF VENEZUELA 57 

Measurements. Snout-vent <5 ? 28, 5 36 ; head breadth $ t 10, 
$ 16 ; head length $ ? 9, 9 12.5 ; femur $ ? 11.5, 9 16 ; tibia 
S ? 13, 9 17.5. 

Additional Localities. Merida (U.M.M.Z. 46461, 51262, co- 
types; Nieden, 1923) ; nr. Merida (A.M.N.H. 10507). 

Range. The subtropical zone of the Merida Andes. 

Remarks. The species has, even when preserved, a very unique 
look about the eyes that makes it easily distinguished anywhere. 

The two folds that were described on the anterior part of the 
back were only detected in U.M.M.Z. 51262, probably a male. 

Eleutherodactylus turumiquirensis sp. n. 

Type. American Museum of Natural History No. 22557, a $ 
from La Trinidad, Mt. Turumiquire, in cave at 5,500 ft. Coll., 
Tate and Clements. 

Diagnosis. A large Eleutherodactylus with broad head, trans- 
verse vomerine teeth, curved canthus rostralis, tympanum y 2 the 
eye diameter, no outer metatarsal tubercle, heel extending to be- 
tween eye and nostril, and light colored (red?) spots on the 
anterior part of the thighs. 

Description. Head as broad as long; snout broad, subovoid ; 
tongue wide, emarginate and free behind ; vomerine odontoids 
small, in two transverse groups behind the small, round choanae ; 
eye diameter shorter than distance between eye and nostril ; inter- 
orbital space about 1% times broader than an upper eyelid ; 
canthus curved, well defined ; loreal strongly sloping, concave ; 
tympanum small, V2 the eye diameter ; two moderate metacarpal 
tubercles ; subarticular tubercles moderate ; disks large, about % 
the size of the tympanum ; first finger reaching the base of the 
disk of the second ; femur half the length of the body ; no outer 
metatarsal tubercle ; toes free, their disks smaller than those of 
the fingers ; heel of the adpressed hind limb extends to between 
eye and nostril. Skin above, finely granular, the granules being 
smaller in the head region. Below, smooth on the throat, chest 
and antero-central portion of the belly; rest of the belly and 
posterior part of the thighs granular. 

Color. Above, mostly plain brown with an obscure interorbital 
bar and two other diffuse dark spots on the scapular region ; 
upper lip with dark, vertical bars; thighs, hidden portions of 
the tibiae and posterior parts of the flanks with several large, 
yellowish (red?) spots. Below, white, marbled on the throat and 
more loosely marbled (some brown reticulations) on the belly. 



58 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Measurements. 9 snout-vent 40 ; head breadth 14 ; head length 
14; femur 20; tibia 21.5. 

Remarks. The paratypes are both females with the same data 
as the type and agreeing in most characters : 

A.M.N.H. 22558, snout-vent 46 ; head breadth 17 ; head length 
16.5; femur 21.5; tibia 24. The spots on the limbs and flanks are 
larger and much better defined ; there is a very small and indis- 
tinct outer metatarsal tubercle. 

A.M.N.H. 22559, snout-vent 39.5; head breadth 14; head length 
14 ; femur 20 ; tibia 23. This specimen is rather desiccated ; there 
are no distinct light spots on the thighs and tibiae, but, on the 
other hand, there are some dark, lightly margined ones. The 
heel reaches the tip of the snout. 

The species is quite distinct from everything known from Vene- 
zuela. 

Eleutherodactylus terraebolivaris sp. n. 

Hylodes gollmeri Peters, 1863, Monats. Akad. Wissensch. Berlin: 409; Lutz, 

A., 1927, Mem. Inst, Osw. Cruz, 20: 40. 
Hylodes gollmeri Niecien, 1923, Das Tierreich. Anura I: 464 (part). 

Type. Museum of Comparative Zoology, no. 31062, a 9 from 
Rancho Grande, Edo. Aragua. Coll. J. A. Rivero, 20 Dec. 1958. 

Diagnosis. An Eleutherodactylus of moderate size, with long 
snout ; sharp canthus rostralis ; eye diameter shorter than dis- 
tance between eye and nostril ; broad disks ; heel of the anteriorly 
adpressed hind limb extending to the tip of the snout or beyond ; 
granular venter ; no femoral spots ; two small, black spots on the 
scapular region, and usually, two other black spots on the snout, 
in front of the eyes. 

Description. Head longer than broad ; snout long, subelliptical ; 
tongue oval, emarginate or free behind ; vomerine odontoids in 
two oblique series behind and between the choanae, their posterior 
extremities directed inwards ; eye diameter shorter than distance 
between eye and nostril ; interorbital space as broad as an upper 
eyelid ; canthus angular ; loreal almost vertical, scarcely concave ; 
tympanum distinct, % the eye diameter ; a supratympanic fold 
from posterior corner of the eye to shoulder ; an indented outer, 
and an oval, prominent, inner metacarpal tubercle ; subarticular 
tubercles conical, distinct; first finger equal to second; disks 
truncate, about % the size of the tympanum ; an indistinct, short 
tarsal fold originating below the middle of the tarsus and ex- 
tending to the inner metatarsal tubercle ; toes free ; heel of the 



RIVERO: SALIENTIA OF VENEZUELA 59 

adpressed hind limb extending to beyond the tip of the snout. 
Skin above and on the flanks finely shagreened. Throat and 
anterior part of the belly, smooth ; posterior part of the belly 
with small granules. 

Color. Above, brown, with irregular, obscure spots ; some of 
these chevron shaped ; two small, elongated spots on the snout ; 
two rounded, small scapular spots and two smaller ones in back 
of these ; loreal region, especially the upper part, darker than 
the rest of the head ; tympanum traversed above by a black 
supratympanic streak ; limbs crossed by dark, light margined 
bars. Ventral surfaces white, uniform except on the limbs, where 
some infuscation occurs. 

Measurements. 9 Snout -vent 31; head breadth 10.9; head 
length 12.7; femur 11.3; tibia 18.7. 

Habits. The species is quite abundant in the subtropical forests 
of the Coastal Range, occurring on the margins of streams to- 
gether with Atelopus c. cruciejer and Prostherapis t. trinitatis. 

Additional Localities. Avila, Cerro (U.C.V. 120-3, II.P.R. 
653) ; Bejuma (U.M.M.Z. 55545) ; Rancho Grande (U.C.V. 18, 
72, 105). 

Range. The Coastal Range of Venezuela. (See below.) 

Remarks. The following specimens are designated as para- 
types: U.S.N.M. 117527-36, 128807-34, Los Canales; U.S.N.M. 
128835, Quebrada Chacaito; U.S.N.M. 128836, Camino de Gali- 
pan. Rio Cotiza; U.S.N.M. 128884, Curupao ; U.M.M.Z 55545, 
Bejuma; U.C.V. 120-3, Cerro Avila; U.C.V. 18, 72, 105, Rancho 
Grande ; U.P.R. 653, Pie del Avila. In general physiognomy all 
paratypes agree with the type, but the following variations may 
occur : 

1. The vomerine teeth may be very strong and joining in the 
middle. 

2. The heel of the adpresesd hind limb may only extend to the 
tip of the snout. 

3. The dorsal surfaces are occasionally granular and/or with 
scattered tubercles. 

4. Two small tubercles are generally present on the snout, in 
the middle of the two black spots. 

5. Two scapular spots are present in greater or lesser degree 
of distinctness in all individuals. 

6. The type is an exception in having an almost smooth belly ; 
in all paratj'pes the posterior part of the belly is distinctly 
granular, the granules being larger than the dorsal granules. 



60 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

7. A narrow vertebral line and sometimes two broad paraver- 
tebral bands may be present in some individuals. 

8. Adult size seems to be variable. Specimens of 50 mm. have 
been measured. 

9. Some of the specimens from Cerro Avila at U.C.V. were 
greenish gray when fresh. 

Examination of a specimen labeled as "Typus" of Eleuthero- 
dactylus gollmeri (Berlin, Mus. 3168, Caracas) reveals this ani- 
mal to represent the Central American form currently known 
as Elciitherodactylus lanciformis, an animal well characterized 
by its narrow disks and smooth belly. As Peters' description of 
E. gollmeri also corresponds to the Central American form, it 
appears that the type was somehow mislabeled or that a collection 
of Central American frogs got mixed with one from Venezuela, 
with the result that the name E. gollmeri has persistently, though 
incorrectly, been applied to the Venezuelan species. The name 
Eleutherodactylus terraebolivaris is here suggested for the Cara- 
cas and Coastal Kange form previously known as E. gollmeri. 

Under the name of E. gollmeri, a frog has been reported from 
Ecuador (Parker, 1936; Andersson, 1945; Boulenger, 1882; 
Peracca, 1904), Bolivia (Andersson, 1906), Matto Grosso (Bou- 
lenger, 1903), and Orgel Gebirge, Brasil (Bauman, 1912). The 
record from Orgel Gebirge probably refers to E. gilntheri while 
that from Matto Grosso may represent the E. conspicillatus 
ileamazonicus here described. It seems to me that two specimens 
from Bolivia at M.C.Z. (10094-5) represent E. longirostris and 
that may also be the case with other records from Bolivia and 
Ecuador. Andersson (1945:28) seems to include a composite of 
E. c. conspicillatus and perhaps E. longirostris, under his E. 
gollmeri. 

Some specimens of E. terraebolivaris may be almost impossible 
to separate morphologically from E. c. conspicillatus, especially 
if in the latter, thigh spots are indistinct and webs are lacking 
(as they sometimes are). E. terraebolivaris is always granular 
on the belly, even although in most cases, the granules may be 
restricted to the posterior half. E. c. conspicillatus is usually 
smooth below, and when granular, the granules are usually finer 
than the dorsal granules. One can say that if it is absolutely 
smooth it is not E. terraebolivaris, but the opposite cannot be 
said of E. c. conspicillatus. 

E. conspicillatus ileamazonicus is also similar to E. terraeboli- 
varis, but it is apparently separated from the range of that 
species (here presumed to be limited to the subtropical regions of 



RIVERO : SALIENTIA OP VENEZUELA 61 

the Coastal Range of Venezuela) by the whole Llanos. Neither of 
the two forms has been recorded from the Guianas nor the 
Delta Region, and tcrracbolivaris is not known from the 
easternmost section of the Coastal Range. So far as known, the 
two forms are thus completely discontinuous in their ranges. 
On the other hand, E. c. conspicillatus has been collected by 
Stebbins at Meta, Colombia (vide), and if Boulenger's "E. goll- 
meri" from Matto Grosso represents E. conspicillatus ileamazoni- 
cus, it would follow that the race would occupy most of the 
Amazonian region, west to the Andes, where the typical form 
occurs. 

The proposed relationship of E. c. conspicillatus and E. c. 
ileamazonicus is admittedly based on little evidence, but the 
morphological similarity of frogs belonging to this general group 
of Eleutherodactylus is so great, and the relationships between 
each other are so obscure at present that it has not been con- 
sidered desirable to propose a new species name for the Ama- 
zonian form. 

Related to E. terracbolivaris are E. longirostris, from Bolivia 
and Ecuador, and Central American E. gollmeri and E. fitzingeri, 
this last a species which may be found to be conspecific with 
E. c. conspicillatus. Conspicillatus, gollmeri and longirostris may 
sometimes have the scapular and snout spots that are found in 
E. terraebolivaris. 

Aleman's, 1952, Eleutherodactylus cf. gollmeri from Turgua 
most probably refers to this species. 

Eleutherodactylus brachypodius sp. n. 

Type. Museum of Comparative Zoology No. 28568, a 9 from 
the Upper Cunucunuma region, Territorio Amazonas. Coll. J. A. 
Rivero, May to June, 1950. 

Diagnosis. A large Eleutherodactylus with long snout, tym- 
panum % the eye diameter ; first finger as long as the second 
and very short hind limbs, the heel of which does not reach the 
nostril. 

Description. Head longer than broad; snout long, subelliptical 
but with a rather blunt tip ; tongue broad, granular, emarginate 
and free behind ; vomerine odontoids in two short, oblique groups 
behind and between the choanae, their posterior extremities con- 
vergent ; eye diameter slightly shorter than distance between eye 
and nostril ; interorbital space a little narrower than an upper 
eyelid ; canthus well defined ; loreal scarcely oblique, slightly con- 
cave ; tympanum % the eye diameter ; a supratympanic fold 



62 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

from posterior corner of the eye to shoulder ; a terminally divided 
palmar and an oval, more prominent, inner metacarpal tubercle ; 
subarticular tubercles moderate ; first finger as long: as second ; 
disks truncate, about % the diameter of the tympanum ; ap- 
parently an indication of a tarsal fold on the distal half of the 
tarsus ; metatarsal tubercles moderate ; toes free, disks of the toes 
smaller than those of the fingers ; thigh half the length between 
anus and anterior margin of upper eyelid ; heel of the adpressed 
hind limb extends to the posterior corner of the eye. Skin above, 
shagreened ; flanks granular. Below, smooth on the throat, breast, 
and central part of the belly ; sides of the belly and hind part of 
the thighs granular. 

Color. Above, very dark slate gray, almost black ; snout rather 
browner; a black canthal and fine supratympanic streak; tym- 
panum chestnut ; lower part of the loreal region and lower part 
of the flanks grayish white ; limbs grayish with darker diffuse 
spots. Below, dirty white. 

Measurements. 9 snout-vent 38.5; head breadth 14; head 
length 15 ; femur 16 ; tibia 18. 

Remarks. U.P.R. 213, a 9 from the forest floor of Mt. Mara- 
huaca, 5,000 ft. and U.P.R. 210, a 9 from a "conuco" or Indian 
plantation at La Culebra, 1000 ft. are very similar in structure 
to E. braehypodius but have not been designated as paratypes 
on account of some differences that may or may not be of signifi- 
cance. In both, the hind limbs look longer, the heel reaches 
between eye and nostril, the color above is much lighter (yellow- 
ish gray) and with irregular darker markings, there is no distinct 
separation between dorsal and ventral color, and the canthus in 
no. 213 is more vertical. Both specimens contain eggs and they 
agree in structural characters. 

Eleutherodactylus braehypodius differs from E. ejollmeri in its 
less tapering snout, finely granular posterior part of the belly, 
absence of scapular spots and much shorter hind limbs ; from the 
Venezuelan E. conspiciUatus in its less tapering snout, light 
loreal region, absence of chevron markings on the dorsum and 
much shorter hind limbs. 

Unfortunately, the exact locality where the type of E. braehy- 
podius was collected was not recorded. From my field notes I 
deduct that it is either Casa de Julian or Marahuaca, 4,050 ft. 



RIVERO: SALIENTIA OF VENEZUELA 



63 



Eleutherodactylus conspicillatus ileamazonicus ssp. n. 

Figure 2 

Hylodes frenatus Boettger, 1896, Ber. Senckenb. Naturf. Ges., LIV (nom. 
nud. ) . 
Type. Museum of Comparative Zoology No. 30397 a 6 from 
Temiche, Mt. Marahuaca, 4050 ft,, S. Venezuela. Coll. J. A. 
Eivero, May 1950. 




■• *** .-.. r_. . 





Fig. 2. Eleutherodactylus conspicillatus ileamazonicus ssp. 
M.C.Z. 30397. 



n. 



Type 



Diagnosis. A medium-sized Eleutherodactylus differing from 
the typical form in its smaller size, shorter legs and in lacking 
toe webs and white spotting or marbling on the posterior part of 
the thighs. 

Description. Head longer than broad, snout subelliptieal ; 
tongue oval, free and nicked behind ; vomerine odontoids in two 
strongly oblique and separated groups behind and between the 
small, round choanae, their posterior extremities convergent; 
eye diameter equal to distance between eye and nostril ; inter- 
orbital space more or less as broad as an upper eyelid ; canthus 
well denned ; loreal almost vertical, slightly concave ; tympanum 
distinct, % the eye diameter ; a supratympanic fold from pos- 
terior corner of the eye to shoulder ; a divided outer and an oval, 



64 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

inner metacarpal tubercle ; subarticular tubercles distinct, 
rounded ; fingers free, the first more or less equal to second ; 
largest disk about % the size of the tympanum ; a round outer 
and an oval inner metacarpal tubercle ; toes free ; heel of the 
adpressed bind limb extends to between eye and nostril. Skin 
above, finely granular, with larger granules or tubercles scattered 
among the finer ones especially behind the eyelids and toward 
the flanks. Below, granular on the sides of the throat, belly and 
binder aspect of thighs. A ventral discoidal fold and a moderate, 
subgular vocal sac. 

Color. Above, brown with irregular, confluent markings on the 
back ; one of the dorsal markings forms a chevron in the middle 
of the dorsum ; another, forming an interorbital spot, sends two 
extensions backward to the sides of the chevron ; two dark dots 
on the snout, in front of the eyes, and in the center of each of 
these a small tubercle ; a dark supratympanic and a canthal 
streak ; loreal region lighter than the body color and with a few 
short dark bars on the lip ; a few dark spots on the anterior distal 
border of the thighs; limbs crossbarred; posterior part of the 
femur uniform brown, of the tarsus and sole dark brown. Below, 
brownish w T hite. 

Measurements. $ snout-vent 31.6; head length 12.8; head 
breadth 11; femur 16; tibia 17.6. 

Habits. All specimens were collected on the forest floor or on 
the steins of plants at not more than a foot from the ground. 
These frogs were heard at dusk and dawn but not during the 
intervening hours of the night. The species apparently prefers 
"conucos'' and cleared areas where the conditions are not very 
humid or the vegetation dense. The voice is something like a 
"toot, toot, toot." 

Range. The Venezuelan Guayana. The species extends from 
the Colombian Andes to Peru. Tt has been reported from east 
and west of the Andes and also from Matto Grosso (Cope, 
1887a). A number of records of E. gollmeri from Ecuador (Bou- 
lenger, 1882 ; Peracca, 1904 ; Parker, 1936 ; Andersson, 1945) may 
refer to this species or to E. longirostris. 

Remarks. The following specimens are designated as para- 
types : 

U.P.R. 105, a 9 from Alto Cunucunuma. Coll. J. A. Rivero, 
April 1950. Snout-vent 39 ; head length 15.6 ; head breadth 14.1 ; 
femur 19 ; tibia 22. 



RIVKRO : SALIENTIA OF VENEZUELA 65 

U.P.R. 107, a i from Mt. Marahtiaca, 4050 ft. Coll. J. A. 
Rivero, May 1050. Snout-vent 29.7; head length 11.7; head 
breadth 11; femur 15.7; tibia 17. 

U.P.R. 38, an immature specimen from Mt. Marahuaca, 4050 
ft. Coll. J. A. Rivero, May 1950. Snout-vent 18.3. 

U.P.R. 108, a S from Tapara, Alto Cunucunuma. Coll. J. A. 
Rivero, April 1950. Snout-vent 24; head length 10.4; head 
breadth 9.5 ; femur 13 ; tibia 14.4. 

U.P.R. 212, a S from Tapara, Alto Cunucunuma. Coll. J. A. 
Rivero, April 1950. Snout-vent 26.7; head length 11; head 
breadth 9.5; femur 12.5; tibia 15.5. 

Paratypes agree morphologically with the type except for the 
fact that the tympanum can only be V2 the eye diameter and the 
heel of the adpressed hind limb may reach only to the anterior 
corner of the eye. All specimens from Mt. Marahuaca or the 
Cunucunuma River have irregular or chevron-shaped markings 
on the dorsum, but in the specimen 211 from Mt. Duida, they 
are not apparent. The basic body color may be of a variable 
shade of brown and the loreal region may be lighter or darker 
than the body color. The two snout dots are present in all speci- 
mens from Mt. Marahuaca, although not too well defined in 
some. There is no marbling or spotting on the hind part of the 
thighs of any of the examples. 

Other material examined includes specimens from the foot- 
hills of Mt. Duida, 750-800 ft. (A.M.N.H. 23205, 23176) ; Caiio 
Pescado, 325 ft. (A.M.N.H. 23183-4) ; Cerro Yapacana (U.S.N.M. 
83950) ; Iguapo, S. Venezuela (Senekenb. Mus. 51844-5) and Mt, 
Duida, 2000-4000 ft., (U.P.R. 211). The latter is more or less 
of a solid color (some darker scribblings may be detected) and 
does not show the two snout dots of the type and paratypes. 

The Venezuelan form differs from Ecuadorian material in its 
smaller size (males especially are smaller), total absence of web 
in the toes, lack of any marbling or spotting on the posterior 
aspect of the thighs, and shorter hind limbs. Out of 43 specimens 
of Ecuadorian and Peruvian E. conspicillatus examined, only 4 
did not have any web (the largest four — one of them 65.1 mm. 
in snout-vent length) while all of them had some marbling or 
spotting (even if not too clear in a few) on the posterior aspect 
of the thighs. 

Morphologically, this species is almost identical with E. terrae- 
bolivaris but the Venezuelan ra( ,r> can be distinguished by its 
darker color, smooth or finelv granular belly and shorter hind 



66 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

limbs. The length of the femur from the anus is included twice 
in the distance between anus and tip of the snout in conspicillatus 
ileamazonicus but in gollmeri the double measurement usually 
extends beyond the tip of the snout. In E. c. ileamazonicus the 
extended heel does not pass beyond the nostril while in E. terrae- 
bolivaris it usually extends considerably beyond the tip of the 
snout. A good distinguishing character is the appearance of the 
belly. In E. c. ileamazonicus the belly is smooth on the center or 
finely granular (the granules not larger than the dorsal granules) 
while in E. terreabolivaris the ventral granules are large and 
distinct. 

A 31 mm. 9 specimen of E. conspicillatus, from nr. Sn. Mar- 
tin, Meta, Colombia (Berkley) has the thigh marbling of the 
topotypical form. The species has also been reported from Matto 
Grosso (Cope, 1887a), but these examples are said to have lacked 
the thigh marblings of the Andean animal. 

The type and one paratype (U.P.R. 105) are very similar in 
dorsal pattern and body form to Boulenger's (1882, pi. xiv) 
figure of E. conspicillatus. 

Hylodes peruvianas Melin, 1941, is probably a synonym of 
E. c. conspicillatus. 



Eleutherodacttlus stenodiscus Walker and Test 

Figure 3 

Eleutherodactylus stenodiscus Walker and Test, 1955, Occ. Pap. Mus. Zool. 
Univ. Mich. No. 561 : 2. Pico Periquito, Rancho Grande. 
2 (U.M.M.Z. 109870, paratypes) Pico Periquito, Rancho 

Grande, 1375 m. 
Description. Head slightly longer than broad; snout long, 
subovoid ; tongue rounded, % free, slightly nicked behind ; vo- 
merine odontoids small, oblicpie, converging posteriorly, between 
and immediately behind the round choanae, their anterior extremi- 
ties at level with the posterior margin of the latter ; eye large, its 
diameter as long as the snout ; interorbital space slightly broader 
than an upper eyelid; canthus well denned but rounded; loreal 
moderately sloping, concave ; tympanum very small and indis- 
tinct, less than % the eye diameter ; two tubercles at the angle 
of the jaw; two or three small, indistinct tubercles on the arm 
close to the wrist ; first finger shorter than second, which is more 
or less as long as fourth ; an elongated inner and a divided outer 



RIVERO : SALIENTIA OF VENEZUELA 



67 



metacarpal tubercle ; disks subtriangular, pointed, that of the 
first finger much smaller than others; a few tubercles at the 
heel, one or two of which are very prominent and distinct ; a 
slight tarsal fold ; metatarsal tubercles small; subarticulars small, 




Fig. 3. Eleutherodactylus stenodiseiis Walker and Test. Paratype 
U.M.M.Z. 109870. 



not prominent; toes free, their disks pointed, as large as those 
of the fingers ; heel of the adpressed hind limb extends to between 
eye and nostril. Skin above shagreened or granular, with scat- 
tered, small tubercles ; upper eyelids with several small tubercles 
and a distinct prominent one near the top. Below, granular on 
the belly and posteroventral aspect of the thighs ; throat smooth. 
Color. Above, grayish brown, with diffuse darker blotches ; 
eyelids dark brown; a light interorbital line diffuses anteriorly 
and blends with the brown color of the snout ; a dark brown 
canthal spot and a supratympanic streak ; two brown bars radiat- 
ing from eye to upper lip ; between these, and anterior to the 
foremost, two other smaller dark spots; white labial dots occur 
between the brown bars and spots; two distinct, rounded brown 
spots above the groins ; seat with a brown triangular spot that 



68 BULLETIN : .MUSEUM OF COMPARATIVE ZOOLOGY 

extends distally halfway on the posterior part of the thighs ; 
thighs with or without crossbars above ; humerus, tibia and tarsus 
with dusky crossbars or diffuse blotches. Ventral surfaces light 
brownish, with greater inf uscation on the throat and limbs ; lower 
lip margined with white dots; tarsus and foot dark brown. 

Measurements. Snout-vent 18.5 ; head length 6.9 ; head breadth 
6.8 ; femur 8.5 ; tibia 9.5. 

Habits. Collected in the forest floor at 1375 m., "where there 
were considerable accumulations of leaf litter" (Walker and 
Test, op. tit.). 

Range. Known only from the type locality. 

Remarks. The main variation is said to be in dorsal pattern. 
The lumbar spots were absent in 5 of the 24 individuals available 
to Walker and Test, but even those 5 had a dorsolateral stripe 
terminating above the groin. In paratype 109870 (field no. AB 
4392), there are 2 small spots in front of the lumbar ones. A 
dusky diagonal bar from shoulder to sides is present in this 
specimen, and according to the description, in the type and all 
paratypes. In 109870 (field no. AB 4380) it is, however, limited 
to a small diffuse spot above the shoulder. Chevron-shaped 
markings or longitudinal stripes are described for some of the 
specimens. 

E. stenodiscus is similar to the Jamaican E. andrewsi and to 
a lesser extent, to E. emiliae from Cuba. Both of these species 
have pointed disks, two lumbar spots and a black seat, but the 
tympanum is larger and more distinct in the two Antillean forms, 
and in E. emiliae the eyes are small and the head much broader 
at the base. 

Among the Venezuelan Eleutherodactylus, stenodiscus can bo 
easily distinguished by its characteristic coloration, indistinct, 
almost hidden tympanum, and pointed disks. 

Eleutherodactylus orocostalis sp. n. 
Figure 4 

Type. Universidad Central de Venezuela No. 2003, a 9 from 
El Junquito, D.F. Coll. Roze, 12 April 1951. 

Diagnosis. A small Eleutherodactylus with subtriangular 
snout; head slightly broader than long; eye diameter greater 
than distance between eye and nostril ; heel extending to the eye ; 
dorsum tubercular, with a large tubercle on the upper eyelid 
and another at the heel; color variable, usually black or very 
dark brown above; marbled or reticulated below. 



RIVERO : SALIENTIA OF VENEZUELA 



69 



Description. Head slightly broader than long; snout subtri- 
angular ; tongue oval, free and nicked behind ; vomerine odon- 
toids in two small, rounded groups behind and between the 
ehoanae, their exterior extremities not extending to the vertical 
of the inner margin of the latter ; eyes protuberant, their diameter 








Fig. 4. Eleutherodactylus orocostalis sp. n. Type TJ.C.V. 2003. 



longer than distance between eye and nostril but slightly shorter 
than the snout ; interorbital space as broad as an upper eyelid ; 
canthus well defined ; loreal little inclined, concave ; tympanum 
distinct, % the eye diameter; two small tubercles between tym- 
panum and arm ; others between elbow and wrist ; metacarpal 
tubercles small ; first finger shorter than second ; first disk small 
and narrow, only slightly broader than phalanges; others rela- 
tively large, almost as large as the tympanum ; a large tubercle 
at the heel and other smaller ones at the tarsus ; a rounded outer 
and an elongated inner metatarsal tubercle; subarticular tu- 
bercles moderate ; heel of the adpressed hind limb extends to the 
middle of the eye. Skin above, tubercular; upper eyelid with a 
large tubercle and other smaller and less distinct ones; upper 
flanks tubercular. Bellv granular. 

Color. Above, very dark brown, almost black ; limbs with nar- 
row oblique listing over a lighter background. Below, dusted and 
marbled with brown. 



70 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Measurements. 2 snout-vent 20.5; head breadth 8.2; head 
length 8 ; femur 10.1 ; tibia 10.2. 

Habits. During the daytime, E. orocostalis is usually found 
under the leaf litter in heavily wooded areas. 

Remarks. Paratypes (U.C.V. 2001-7, El Junquito) agree in 
morphological characters with the type except for the fact that 
the heel may extend to the anterior corner of the eye or slightly 
beyond, and the dorsal skin may be less or more tubercular than 
in the type specimen. In size, none of the examples available 
surpasses the type. 

Coloration is quite variable, the following patterns being the 
most common : 

a. Yellowish brown, tan or chestnut with a dark brown spot 
on the middle of the body between occiput and sacrum and two 
broad, dark brown, lateral bands that cover the loreal region 
and extend posteriorly to the groins. 

b. Very dark brown with darker, almost black markings. 

c. Variegated with greenish gray and dark gray. 

The loreal region may be of a solid color or with distinct cross- 
bars that radiate from eye to upper lip ; flanks solid brown or 
with spots or oblique light and dark bands; limbs distinctly 
crossbarred, the bars on the thighs being sometimes separated 
by white spaces ; posterior part of the thighs solid brown or with 
obscure, small, lighter spots. All specimens have a large tubercle 
on the upper eyelid, another at the heel, two or three at the arm 
and two or three at the tarsus. The heel tubercle is mammillary 
and very distinct in some specimens. In most, a W is formed 
by four or five tubercles on the scapular region. 

E. orocostalis is similar to E. rozei (see "Remarks" under that 
species) and, to some extent, to the young of E. bicumulus. Tn 
this latter species the snout is longer, the orbital diameter not 
being greater than the distance between eye and nostril, the dor- 
sum is not tubercular, and the heel extends anteriorly to between 
eye and nostril. 

Bleuthebodactylus bicumulus (Peters) 

Eylodes bicumulus Peters, 1863, Monatsb. Akad. Wissensch. Berlin: 410: 
Caracas; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus. ed. 2: 215; Nieden, 
1923, Das Tierreich. Anura I: 465; Lutz, A., 1927, Mem. Inst. Osw. 
Cruz, 20: 39. 

2 (Berlin Mus. 4899, cotypes) Caracas 



RIVERO : SALIENTIA OP VENEZUELA 71 

Description. Snout subovoid; tip of snout rounded; tongue 
large, slightly nicked behind ; vomerine odontoids in two small, 
rounded and close together groups between and behind the choa- 
nae ; eye diameter as long as distance between eye and nostril ; 
interorbital space as broad as an upper eyelid ; canthus distinct ; 
loreal somewhat sloping, concave; tympanum small, % the eye 
diameter, with a distinct supratympanic fold ; first finger shorter 
than second ; disks well developed ; a small tubercle at the heel ; 
outer metatarsal tubercle indistinct, inner more prominent, elon- 
gated ; heel of the adpressed hind limb extends to the middle of 
the eye. Above, smooth ; flanks and posterior part of the eyelids 
with small, scattered granules. Below, smooth on the anterior 
part of the chest and throat ; belly and thighs granular. 

Color. Above, tan ; a black canthal streak continues posteriorly 
over the tympanum and forms a broad lateral band that extends 
to the groins ; limbs crossbarred ; posterior aspect of the thighs 
dark brown. Chest and belly white, immaculate or slightly 
reticulated. 

Measurements. Snout-vent 34.9 ; head length 11.9 ; head 
breadth 12.5 ; femur 16.5 ; tibia 18.9. 

Remarks. The name E. bicumulus has been applied to a num- 
ber of forms but cotypes reveal a well characterized animal not 
too easily confused with any of the other Coastal Range forms. 
In general physiognomy, E. bicumulus looks somewhat like E. 
urichi but it is apparently a larger species and has the lateral 
band that immediately separates it from urichi. Some specimens 
of E. guntlieri and E. orocostalis also show lateral bands. The 
first is a southern species with small disks not much broader than 
the phalanges, while E. orocostalis is a very small form with 
short snout and usually tubercular dorsum. 

Through the courtesy of Dr. Charles Walker and Dr. Frederick 
Test, I have been able to examine three specimens of E. bicumu- 
lus collected by the latter in Venezuela. As these may be used 
for a separate report by Dr. Test they are not included here, but 
it may be well to point out that they agree in most essentials 
with the cotypes, the lateral band occurring in all specimens in 
greater or lesser degree of intensity. In the younger examples, 
the snout seems to be longer and the loreal region more vertical, 
resembling the condition in E. terraebolivaris. They can be dis- 
tinguished, however, by the shorter hind limbs, presence of the 
lateral band and absence of scapular spots. 



72 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



Eleutherodactylus williamsi sp. n. 
Figure 5 

Type. Universidad Central de Venezuela No. 2012, a ? from 
El Junquito, D. P. Coll. Roze, 16 October 1949. 

Diagnosis. A small Eleutherodactylus with short, transverse 
vomerine teeth slightly behind the choanae ; tympanum % the 
eye diameter ; a row of tubercles between elbow and wrist ; first 
finger shorter than second ; heel of the adpressed hind limb ex- 
tending to nostril; tubercular dorsum, granular belly and a 
white (pink or red?) spot in the axilla. 

Description. Snout subovoid; tongue oval, free and slightly 
nicked behind; vomerine odontoids in two short, transverse 
groups slightly behind the choanae ; eye diameter little shorter 




Fig. 5. Eleutherodactylus williamsi sp. n. Type TT.C.V. 2012. 



than distance between eye and nostril; interorbital space about 
iy 2 times broader than an upper eyelid ; canthus distinct, curved ; 
loreal slightly oblique, concave; tympanum small, % the eye 
diameter; a slight supratympanic fold; two small tubercles be- 
tween tympanum and arm; a row of small tubercles between el- 
bow and wrist; metacarpal tubercles indistinct; subarticular 



RIVERO : SALIENTIA OF VENEZUELA 73 

tubercles rounded, not very prominent ; first finger shorter than 
second ; disks fan-shaped, the largest more or less as large as 
the tympanum; a small tubercle at the heel; metatarsal tubercles 
very small ; heel of the adpressed hind limb extends to nostril. 
Skin above, mostly smooth on the head, densely tubercular on 
the body and sparsely tubercular on the limbs ; some tubercles 
fuse and form long ridges on the sides of the dorsum ; a row of 
three small tubercles on the head between occiput and tip of the 
snout ; upper eyelid slightly tubercular ; flanks tubercular above, 
granular below. Belly granular. 

Color. Snout dark chestnut; body behind a narrow, light, in- 
terorbital line very dark brown; a white (pink or red?) spot in 
the axilla and in the inner angle of the elbow ; rest of the humeral 
segment light brown ; a broad, dark band around the radio-ulnar 
segment ; an extensive, but not well delimited, white, inguinal 
spot ; anterior part of the thighs with a few light spots, posterior 
part brown with obscure light spots above and a few dark in- 
distinct ones below; rest of the hind limbs uniform dark brown. 
Below, dusted or speckled with brown. 

Measurements. $ Snout -vent 23; head breadth 9.5; head 
length 9 ; femur 12 ; tibia 13. 

Remarks. Eleutherodactylus williamsi is quite distinct from 
everything known from Venezuela and can be distinguished from 
other species by its tubercular dorsal surface and very typical 
coloration. 

Eleutherodactylus rozei sp. n. 
Figure 6 

Type. ! niversidad Central de Venezuela No. 2018, a <3 ? from 
Curucuruma, Edo. Aragua. Coll. Boze, 22 March 1951. 

Diagnosis. A small Eleutherodactylus with rounded, trans- 
verse vomerine teeth slightly behind the choanae; eye diameter 
greater than distance between eye and nostril; tympanum % 
the eye diameter; a row of three indistinct tubercles between 
elbow and wrist ; first finger shorter than second ; granular belly ; 
distinctly crossbarred limbs and brownish orange regions on the 
anterior and posterior aspect of the thighs. 

Description. Snout short, subovoid ; tongue oval, free and 
slightly emarginate behind ; vomerine odontoids in two round, 
transverse groups behind the choanae; eyes large, their diameter 
slightly greater than distance between eye and nostril ; inter- 
orbital space as broad as an upper eyelid; canthus very distinct, 



74 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



curved; loreal scarcely oblique, slightly concave; tympanum 
small, ]/ 3 the eye diameter; a slight supratympanic fold from 
posterior corner of the eye to shoulder; two small tubercles be- 
tween tympanum and arm ; a row of three small tubercles between 
elbow and wrist ; metacarpal tubercles very small and indistinct ; 




Fig. 6. Eleutherodactylus rosei sp. n. Type U.C. V. 2018. 

subarticular tubercles small and indistinct; first finger shorter 
than second; disks large, fan-shaped, the largest more or less 
equal to the tympanum ; a small tubercle at the heel ; metatarsal 
tubercles minute ; heel of the adpressed hind limb extends to be- 
tween eye and nostril. Skin above, with scattered small tubercles ; 
a small, flat tubercle on each upper eyelid; flanks tubercular. 
Belly finely granular. 

Color. Above, gray with an indistinct bar between the eyes 
and a large W-shaped marking on the occipital region ; a diffuse, 
dark spot in front of the sacrum; a black canthal and a supra- 
tympanic streak; several distinct, black bars radiating from 
lower eyelid to upper lip ; posterior part of the flank and groin 
with an orange brown spot that extends along the anterior part 
of the thighs to the knees; buttocks with several wavy lines and 
round spots; rest of the thighs orange brown on the posterior 
part ; fore and hind limbs distinctly crossbarred with black ; 
ventral surfaces speckled with brown ; outer edge of tarsus and 
fifth toe with a light line. 



RIVERO : SALIENTIA OF VENEZUELA 75 

Measurements. $ snout-vent 20 ; head breadth 7.5 ; head length 
7.5; femur 10; tibia 10.5. 

Remarks. This species agrees in some respects with Eleuthero- 
dactylus inguinalis Parker, 1940, from New River, British Gui- 
ana, but, as well as I can make out from the description of that 
species, it differs in several important characters. The head 
is definitely not depressed as described for E. inguinalis, but 
rather high and Avith very large, protruding eyes; the largest 
disks are not more than % the width of the eye (y 2 in inguina- 
lis) ; the subarticular tubercles cannot be called prominent and 
there is no "black bordered inguinal ocellus." 

Eleutherodactylus rozei is quite similar to E. orocostalis, from 
which it can be best distinguished by its coloration. In E. rozei 
the markings on the body, especially the thighs, are very distinct 
and contrasting, while in E. orocostalis the dorsal color is usually 
black or with obscure markings. Besides, the golden brown stains 
at the proximal anterior and distal posterior portions of the 
thighs of E. rozei are lacking in E. orocostalis. The snout of the 
latter is also more pointed and triangular. 

Eleutherodactylus urichi (Boettger) 

Hylodes urichi Boettger, 1894, Journ. Trin. Field Nat. Club, 2\ 88: Trini- 
dad. 
Eleutherodactylus gollmeri Schmidt, K. P. (not Peters), 1932, Zool. Ser. 

Field Mus. Nat. Hist., 18: 159. 
Eleutherodactylus bicumulus Schmidt, K. P. (not Peters), 1932, Zool. Ser. 
Field Mus. Nat. Hist., 18: 159. 

11 (C.N.H.M. 17777-87) Mt. Turumiquire, 7000-8000 ft., 32. 

Description. Snout long, subovoid ; tongue usually small, oval 
or spatulate, free and entire ; vomerine odontoids in two very 
short and slightly oblique groups behind and between the 
choanae, their posterior extremities converging; interorbital 
space iy 2 to 2 times broader than tbe narrow upper eyelid; eye 
diameter equal or slightly shorter than distance between eye 
and nostril; canthus distinct, usually curved; loreal moderately 
inclined, almost flat; tympanum small, % the eye diameter; 
metacarpal tubercles not very distinct; subarticular tubercles 
moderate, round ; first finger shorter than second and with a much 
smaller disk; disks moderate, the largest almost as large as the 
tympanum ; metatarsal tubercles small, the other very indistinct ; 
toes free ; heel of the adpressed hind limb extends to the middle 
of the eye. Skin above, smooth or with a slight tuberculation on 



76 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

the posterior half ( S ?) ; usually a small tubercle on the upper 
eyelid ; flanks smooth or granular. Below, smooth on the throat 
and chest; very distinctly granular on the belly (with occasional 
exceptions) and finely or more coarsely granular on the thighs. 

Color. Above, uniformly golden or purplish brown, very rarefy 
with blotches or large spots; upper eyelids occasionally edged 
with white ; thighs, groin and hidden portions of tibiae reddish. 
Color of the belly generally purplish between the lighter colored 
granules; throat and chest dirty white or brownish (reticulated 
in 17784, which is spotted above). 

Measurements. Snout-vent $ 29, 9 31 ; head breadth <j 10, 
9 11.5; head length $ 9.8, $ 11; femur 6 13.5, 9 13; tibia 
$ 14, 9 14. 

Range. The eastern section of the Coastal Range. Trinidad 
and the Guianas. 

Remarks. The Venezuelan E. urichi differs from topotypical 
material in its larger size (largest of 10 Trinidad specimens, 
snout-vent 24 mm.), longer and narrower snout, and in the ab- 
sence of body markings in most of the specimens. Of 10 Trinidad 
specimens examined, all have a short, black supratympanic streak, 
9 have distinct or indistinct crossbars on the hind limbs, 7 have 
vertical bars on the upper lip and 8 have some kind of markings 
above. 

Of the 11 Venezuelan examples, only two have a slight 
supratympanic streak and distinct spotting of the dorsal sur- 
faces. C.N.H.M. 17784 is strikingly different from the other 
specimens in coloration, but it agrees in most morphological 
characters. 

Although the Venezuelan E. urichi probably represents a race, 
I have preferred to use the binomial until specimens from other 
northern South American countries have been examined. 



Ele[ t tiierodactylus reticulatus Walker and Test 

El.eutlierodaet ylus reticulatus Walker and Test, 1955, Occ. Pap. Mus. Zool. 
Univ. Mich., No. 561: 4: Pico Periquito, Raneho Grande, 1275 in. 

1 (U.M.M.Z. 109873, paratype) Raneho Grande, 1090 m., 51. 

Description. Head broader than long; snout subovoid ; tongue 
rounded, % free, slightly nicked behind ; vomerine odontoids in 
two small, round and close together series between and just be- 
hind the small choanae ; eye diameter as long as distance between 
eye and nostril ; interorbital space as broad as an upper eyelid ; 



RIVERO : SALIENTIA OF VENEZUELA 77 

eaiithus distinct ; loreal moderately oblique, concave ; tympanum 
small, indistinct, % the eye diameter; a very slight supratym- 
panic fold ; two large tubercles at the angle of the jaw ; a few 
very small tubercles at the elbow; one distinct humeral tubercle 
near the wrist; first finger shorter than second, which is shorter 
than fourth ; an elongated large and a flattened divided outer 
metacarpal tubercle ; disks somewhat indented at their free end, 
the largest larger than the tympanum; some very small tubercles 
at the heel ; no tarsal fold or tubercle ; an oval, rather prominent 
inner and a smaller, rounded outer metatarsal tubercle ; sub- 
articular tubercles rounded, not too prominent; disks of toes 
smaller than those of the fingers ; heel of the adpressed hind 
limb extends to the tip of the snout. Skin above, smooth, except 
for an occasional tubercle; one distinct tubercle at each eyelid. 
Ventral surfaces granular on the throat, belly and posteroventral 
aspect of thighs. 

Color. Above, grayish brown, with two darker blotches at the 1 
scapular region, one in the center, anterior to the sacrum, sev- 
eral small spots or dots at the coccygeal region, and other not so 
well defined mottles or freckles all over the dorsum; no ap- 
parent canthal streak; a narrow, not too distinct supratympanic 
streak; two dark lines radiating from eye to upper lip; anterior 
to this, another triangular labial spot; a few white, irregular 
spots at the groin ; anterior and posterior aspects of the thighs 
equally dotted with white, on a chestnut background ; rest of the 
hind limbs with diffuse, dusky crossbars. Below, brownish, reticu- 
lated with lighter, the hind limbs being darker and with better 
defined white dots ; a series of white dots along the margin of 
the lower lip. 

Measurements. 9 , snout-vent 35 ; head length 12.5 ; head 
breadth 13.3; femur 18; tibia 19. 

Habits. Walker and Test state that some of the specimens were 
collected during the daytime under leaf litter, while another 
example was caught at night while sitting on the leaf of a plant 
at about four feet from the ground. 

Range. Only known from the type locality. 

Remarks. The tympanum of this species, although clearly evi- 
dent, is not too distinct, its margins being more or less level 
with the skin surface. 

The type is described as having a tubercular upper surface and 
a pair of short, crescentic glandular folds in the scapular region. 
The living coloration is said to be "clay brown, with blackish 



78 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



markings on face, back, and legs; venter purplish black and 
white." 

In general form of head and snout this species agrees with E. 
guntheri and E. sancta-martae, which, however, are very dif- 
ferent in other respects. E. fitzingeri and E. conspicillatus which 
are also spotted on the posterior part of the thighs, have a good- 
sized, distinct tympanum and a long, subelliptical snout with a 
strong canthus rostralis. 

Eleutherodactylus racenisi sp. n. 

Figure 7 

Type. Universidad Central de Venezuela No. 2014, a 9 from 
El Junquito, D. F. Coll. Roze, 14 April 1951. 




Fig. 7. Eleutherodactylus racenisi sp. n. Type U.C.V. 2014. 

Diagnosis. A moderate sized Eleutherodactylus with oblique, 
posteriorly converging vomerine teeth situated behind the choa- 
nae; tympanum % the eye diameter; a small tubercle on the 






RIVERO : SALIENTIA OF VENEZUELA 79 

distal portion of the radio-ulnar segment ; first finger shorter than 
second ; large, fan-shaped disks slightly larger than the tym- 
panum and heel of the adpressed hind limb extending to the eye. 

Description. Snout subovoid; tongue oval, slightly nicked and 
free behind ; vomerine odontoids in two oblique, posteriorly con- 
verging groups behind the choanae ; eye diameter equal to 
distance between eye and nostril ; interorbital space slightly 
broader than an upper eyelid ; canthus moderate, curved ; loreal 
oblique, concave; tympanum % the eye diameter; two small tu- 
bercles at the angle of the mouth ; a flat, indistinct supratympanic 
fold ; a small tubercle on the distal portion of the radio-ulnar 
segment and one or two less distinct ones between this and the 
elbow ; metacarpal tubercles indistinct ; subarticular tubercles 
moderate, not very prominent ; first finger shorter than second ; 
disks large, fan-shaped ; the largest slightly larger than the tym- 
panum ; a small tubercle at the heel ; an oval, not very distinct, 
inner and a small, rounded, outer metatarsal tubercle ; toes free ; 
heel of the adpressed hind limb extends to the middle of the eye. 
Skin above, smooth, slightly tubercular in front and behind the 
tympanum ; a transverse fold on each upper eyelid. Below, 
smooth. 

Color. Above, brown with chocolate brown markings of irregu- 
lar form ; one marking forms a canthal streak, one crosses the 
intercanthal space in front of the eyes while others form a 
posteriorly diffuse spot between the eyes ; a pagoda-shaped mark- 
ing in front of the sacrum ; a diffuse but extensive temporal spot 
that extends back on the flanks and a few radiating bars between 
lower eyelid and upper lip ; anterodorsal part of the thighs with 
a row of rounded, white ( pink or red ? ) spots from base to knee 
(in left thigh the spots are confluent and not well defined) ; upper 
part of the thighs with indistinct dark and light bars; posterior 
aspect uniform brown ; forelimbs and rest of hindlimbs cross- 
barred. Ventral surfaces dusted with brown, the throat more 
profusely so than the rest of the body ; tibial segment with a 
large, white (pink or red?) spot; tarsus dark brown; meta- 
tarsus wtih a light outer line that extends to the disk of the 
last toe. 

Measurements. 9 Snout-vent 36 ; head breadth 13 ; head length 
12; femur 14.5; tibia 16. 

Remarks. Elcutherodactylus raccnisi is somewhat similar in 
coloration to E. oriceni but it differs from this species in having 
a narrower head, no elevation of the frontoparietals, slightly 



80 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

smaller tympanum, and in other characters. It also hears some 
superficial resemblance to E. megalops of the Sta. Marta region 
of Colombia, and E. reticulatus of the Coastal Range of Vene- 
zuela. It can be distinguished from both by the smooth dorsum 
and venter, shorter hind limbs, different color of the thighs, and 
different position of the vomerine odontoids. 

Eleutherodactylus marmoratus (Boulenger) 

E\jlodcs marmoratus Boulenger, 1900, Trans. Linn. Soc. London, S: 50, pi. 
V, fig. 0: Mr. Eoraima, 3500 ft. 

5 (U.P.R. 214-18) Mt, Marahuaca, 4,050 ft., v.50. 

Description. Snout short, subovoid to subelliptical ; tongue 
oval, free and slightly nicked behind ; vomerine odontoids form- 
ing two short, slightly oblique groups behind and between the 
round choanae, their posterior extremities directed inwards; in- 
terorbital space broader than an upper eyelid; eye diameter as 
long as distance between eye and nostril ; canthus distinct ; loreal 
almost straight, concave; tympanum small, not more than Y 3 
the eye diameter; metacarpal tubercles slight, subarticular tu- 
bercles rounded, moderate; fingers free, the first shorter than 
second ; disks truncate, of about the same size as the tympanum ; 
metatarsal tubercles small, the inner a little more oval than the 
outer; toes free; heel of the adpressed hind limb extends to 
between eye and nostril. Skin above generally sparsely tubercu- 
lar ; loreal region granular ; flanks, belly and ventral surface of 
the thighs granular; a transverse pectoral fold. Male with a 
large, subgular vocal sac. 

Color. Above, brownish gray with darker, sometimes obscure 
spots; upper eyelids dark; a black supratympanic streak; two 
black, curved, longitudinal lines from near the upper eyelid 
toward the scapular region; a few, sometimes indistinct bars 
radiate from eye to upper lip ; hind limbs crossbarred ; ventral 
surfaces white or greenish white, immaculate. 

Measurements. Snout-vent $ 17.5, 9 22 ; head breadth $ 6, 
9 8 ; head length $ 6.2, 9 7.9 ; femur $ 1.1, 9 10.1 ; tibia 

$ 9, 9 11. 

Habits. Collected at night in shrubs and small trees where 
their voices were heard but could not be recorded. 

Remarks. Two of the three male specimens are longitudinally 
striped with brown, but they were taken in the same place, and 
morphologically they agree with the other three examples. 



RIVERO: SALIENTIA OF VENEZUELA 81 

The non-striped male was sent to Dr. Parker for comparison 
with the type. He pronounced it conspecific with the Roraima 
form. 

See remarks under Otophryne robust a. 

Eleutherodactylus anotis Walker and Test 

Eleutherodactylus anotis Walker and Test, 19.1.1, Occ. Pap. Mus. Zool. Univ. 
Mich. No. 561: 7: Eancho Grande, 1090 m. 
1 (U.M.M.Z. 109877, paratype) Rancho Grande, 1090 m., 51. 

Description. Head hroader than long • snout subovoid ; tongue 
rounded, % free, indistinctly nicked behind ; vomerine odontoids 
round, transverse, close together and immediately behind the 
small choanae ; eye diameter as long as distance between eye and 
nostril ; interorbital space as broad as an upper eyelid ; canthus 
rostralis distinct; loreal region little oblique, concave above; 
tympanum completely hidden ; a slight fold above its site ; a few 
tubercles at the elbow and three or four very indistinct ones 
along the posterior side of the arm ; a palmar and an inner 
metacarpal tubercle; subarticular tubercles rounded, not very 
protuberant; first finger shorter than second, which is shorter 
than fourth ; disks moderate ; a few slight tubercles at the heel ; 
no tarsal fold : inner metatarsal tubercle elongated, outer 
rounded, conical ; toes free, their disks smaller than those of the 
fingers; heel of the adpressed hind limb extends to between eye 
and nostril. Skin above, rugose and tubercular, especially be- 
hind the angle of the jaw, in the temporal region and anterior, 
upper flanks ; a few tubercles at the upper eyelid ; some tubercles 
elongate and form a rough W-shaped figure behind the nape. 
Below, with slight granulation on the venter and posteroventral 
aspect of the thighs. 

Color. Above, grayish brown, with irregular mottling and 
dusting, lighter on the snout, in front of a narrow, posteriorly 
diffused interorbital line ; loreal region with dark brown bars 
that radiate from eye to upper lip ; anterior and posterior aspect 
of the thighs uniform, with some infuscation ; limbs crossbarred. 
Below, brownish white, with greater infuscation on the limbs, 
throat and anterior belly, where light points and reticulations 
can be seen ; a series of white dots margin the lower jaw. 

Measurements. 9 Snout -vent 41 ; head length 15 ; head breadth 
]6; femur 20.5; tibia 23.5. 

Habits. During the day E. anotis hides in rock crevices and 
at night comes out and is found on rock outcroppings in or near 
streams (Walker and Test, op. cit.). 



82 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Range. Known only from within 1 km. of the type locality 
and from a stream beside the Maracay-Charoni road, at 1300 m. 

Remarks. According to Walker and Test, there seems to be 
little structural variation in the species. The head in the larger 
individuals is broader than in the smaller ones and the heel may 
reach to the tip of the snout. A narrow dorsolateral fold that 
extends from above the shoulder halfway to the groin is described 
for the type but is not evident in paratype U.M.M.Z. 109877. 
Neither is an ill-defined femoral ocellus or a distinct dorsal tri- 
angular blotch. One of the immature specimens available to 
Walker and Test had a broad, light, vertebral stripe. 

The coloration of the living animal is described as follows : 
dorsal surfaces greenish to olive, sometimes overlaid with reddish 
cinnamon, or with scattered, pale gray flecks; posterior part of 
the belly and underside of the thighs and shanks reddish, coral 
red or brownish salmon. 

Other large Eleutherodactylus without tympanum are E. ven- 
trivittatus Andersson, 1 E. surdus (Boulenger) and E. roseus 
(Boulenger). 

E. ventrivittatus is distinctly marbled below and has white and 
black thigh bars; E. surdus (from W. Ecuador) has a smooth 
skin, chevron-shaped markings on the back and white spots on 
the posterior aspect of the thighs, while E. roseus has a very 
oblique loreal region, smooth dorsal skin, a white streak on the 
canthus and edge of the upper eyelid and pink spots on upper 
eyelids and flanks. E. whymperi (Boulenger) does not have a 
tympanum but it is a relatively small, compact and rotund form 
with no similarity to E. anotis. Lutz and Kloss described E. car- 
valhoi from the Alto Solimoes, Amazonas, apparently from im- 
mature specimens without exposed tympanum. This species has 
a vertical loreal region and a light spot on each flank, at the 
lumbar area. 

Included under this species are U.C.V. 44, from El Junquito, 
and U.C.V. 2008-9 from La Culebra, Edo. Miranda. These speci- 
mens have a small, indistinct tympanum, but this may be a sign of 
immaturity in this species, as all three examples agree with adult 
E. anotis in other important morphological characters, and in 
some details of their distinct coloration. 

i Perhaps a synonym of E. ventrimarmoratus (Boulenger), although this spe- 
cies is described as having a small tympanum. M.C.Z. 19643-7 and 24437 are 
probably referable to this species, although determined as E. latidiscus. The 
smaller specimens of these do not have any apparent tympanum, are very tuber- 
cular above, have comparatively smaller disks and show a uniform ventral 
surface. 



RIVERO : SALIENTIA OF VENEZUELA 83 

U.C.V. 44 is only 9 mm. long. The color above is greenish 
gray, with a dark green spot on the middle of the body, in front 
of the sacrum ; the loreal region and tip of the snont are coral 
red; the limbs, coral red, crossbarred with dark grayish green; 
there is a whitish area at the knee, and the ventral surfaces are 
much infuscated on the throat and anterior part of the belly. 
In coloration, this little frog looks more like a gaudy, coral reef 
crab than an Eleutherodactylus. 

U.C.V. 2009 is reddish or brownish gray with darker mottlings 
on the upper flanks and dorsolateral region, and it has a greenish, 
well defined interscapular spot and another spot of the same 
color on the middle of the sacral region ; the venter is salmon red. 
In No. 2008 the general dorsal color is gray with irregular spots 
of different shades of gray and a light green interorbital spot ; 
the hind limbs are crossbarred except for the distal portions of 
the tibiae, which are of a contrasting milky white color ; the 
distal portion of the femur and proximal part of the tibia is 
blackish ; the distal portion of the tibia and most of the tarsus, 
white. The peculiar coloration of these small frogs make them 
look as if they were covered with lichens. 

Key to the Species of Ceratophrys Recorded from Venezuela 

I. Toes Vs webbed; heel of the adpressed hind limb extends to the tym- 
panum ; inner metatarsal tubercle shovel-shaped, with a sharp edge ; 

no orange or red on the back calcarata 

II. Toes % webbed; heel of the adpressed hind limb extends to between 
eye and nostril; inner metatarsal tubercle oval, blunt; orange and red 
on the back cornuta 

Ceratophrys calcarata Boulenger 

Ceratophrys calcarata Boulenger, 1890, Proc. Zool. Soc. London: 327, pi. 
xxvi : Colombia. 

1 (U.P.R. 196) Territorio Amazonas, i.48. 

1 (U.P.R, 197) Pto. Ayacucho, vi.50. 
Description. Head much broader than long, bony and very 
high ; snout short and rounded ; tongue rounded, slightly emargi- 
nate and free behind; "vomerine odontoids" present as small 
patches in front of the large, transverse choanae of the male, 
absent in the female ; eye diameter a little shorter than distance 
between eye and nostril ; nostril closer to the eye than to the tip 
of the snout; a longitudinal frontal furrow; interorbital space 



84 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

concave, as broad as an upper eyelid; a triangular dermal ap- 
pendage and several other smaller papillae on the upper eyelid ; 
canthus descending almost vertically to the tip of the snout; 
loreal slightly sloping ; tympanum not very distinct, rather more 
than y 2 the eye diameter ; a flat and broad, bony supratympanic 
ridge; a slight glandular ridge along the forearm; a large but 
somewhat diffuse outer and a smaller oval inner metacarpal tu- 
bercle ; fingers free, the first stronger and longer than second ; 
last finger scarcely extending to the penultimate articulation of 
the longest third ; a distinct tarsal fold ; a horny spade-like, inner 
metatarsal tubercle, the outer absent or indistinct; subarticular 
tubercles of toes moderate, smaller than those of the fingers ; toes 
about y 3 webbed ; heel of the adpressed hind limb extends to the 
tympanum. Skin above, rugose and warty, the larger warts with 
longitudinal ridges ; flanks and ventral surface of the body and 
thighs granular. 

Color. Above greenish gray or whitish with olive green mark- 
ings ; a dark green, usually T-shaped spot in the frontal furrow ; 
several radiating bars from eye to upper lip ; two light-colored 
broad lines originate on the eyelid appendages, meet in front of 
the occipital region and continue posteriorly to the anus, sending 
ramifications between the darker spots of the body ; in the sacral 
region this stripe is usually divided by an oval, central spot ; a 
pair of externally concave, reniform spots between the supra- 
orbital ridges and the central longitudinal stripe ; limbs cross- 
barred. Ventral surfaces yellow white, with brown variegations ; 
lower lip with a series of dark, elongated spots that usually ex- 
tend to the region of the anterior girdle. 

Measurements. 9 Snout-vent 79 ; head breadth 45 ; head 
length 31 ; femur 27 ; tibia 22. 

Habits. "While in San Fernando de Atabapo I heard of a 
woman in Puerto Ayacucho who had been bitten by a poisonous 
toad when she went out at night to get a chicken in her yard. 
Upon returning to Puerto Ayacucho a reward was offered for 
the "poisonous toad" and when a man appeared with a "sapo 
de cachos" tied to a string, the animal was shown to Justina 
Manrique for identification. That was, in effect, the supposed 
poisonous animal, but since the small teeth failed to penetrate 
the skin of my finger, she insisted that the toad loses its poisonous 
properties when kept in captivity. The animal was always ag- 
gressive and held on to a stick with the tenacity of a bulldog. 
Justina still showed a pair of punctures on the ankle and it might 



RIVERO : SALIENTIA OP VENEZUELA 85 

be supposed that she was the victim of a poisonous snake which 
was approaching the toad. On being questioned if she knew 
the toad to be poisonous she answered that she had heard of a 
poisonous "sapo" with "horns" but had never seen one before. 
The doctor who was said to have attended her and who ' ' gave her 
some shots" could not be found. 

The other 9 specimen (U.P.R. 196) was obtained through the 
courtesy of Dr. Hans Baumgardner of Puerto Ayacucho, who 
said that the toad was common in ditches and on a nearby granite 
hill, where, after rain, water accumulated in depressions. Al- 
though a whole night was spent searching for them, no Cerato- 
phrys could be found on this hill nor in any other place in Puerto 
Ayacucho. 

Range. The Venezuelan Guayana and probably the Llanos, 
the coastal belt, the Falcon Region and the lower Maracaibo 
Basin, northeastern Colombia and perhaps the southern Guianas 
and the savannas of N. Brasil. The presence of this species in 
Santa Marta region of Colombia and in Puerto Ayacucho may 
indicate that it is also found in the Llanos and in the arid and 
semiarid regions of N. Venezuela. It is not known by the people 
of San Fernando de Atabapo, an indication that its distribution 
is affected by the higher humidity and other ecological factors 
of this region. 

Remarks. Adolpho Lutz, (1927: 40) records seeing a Cerato- 
phrys from Venezuela but failed to obtain specimens and was 
uncertain whether the frog was a calcarata or cornuta. It might 
well have been an Eleutherodactylus cornutus maussi. 

The Venezuelan animals seem to have a higher head and a 
shorter and more rounded snout than the specimens from north- 
eastern Colombia. A male of this latter group has the following 
measurements : snout-vent 72 ; head breadth 45 ; head length 33 ; 
femur 24.3 ; tibia 23. A vocal sac is present and is generally 
dark in color. 

No vomerine teeth have been detected between the choanae 
but the male toads from Colombia have two whitish patches far 
in front of the openings. These are well raised and represent 
protuberances of some bone, but as the head of this form is so 
much compressed from back to front, it is possible that the 
cranial bones are equally modified or moved and these oclontoids 
may actually come from the vomer, although their position is 
closer to that of the premaxillary of normal forms. The matter 
is under investigation. 



86 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Ceratophrys cornuta (Linne) 

Sana cornuta Linng, 1758, Syst. Nat,, ed. 10: 212: Virginia (in error). 
Ceratophrys cornuta Ernst, A., 1877, Flora y Fauna de Yen.: 281; Rohl, E., 
1949, Fauna Desei'. de Yen., ed. 2: 406, fig. 185. 
No material examined. 

Original Description. (Based on Seba 1, pi. LXXII, figs. 1, 2.) 
Eyelid conical; aspect horrible. 

Color. According to Rohl : above green, brown and black, with 
an orange stripe that extends from the head to the end of the 
dorsum; shoulders reddish green; hind limbs green, streaked 
with lighter green. Jimenez de la Espada (1875:32) speaks of 
an irregular mixture of orange, red, purple and dark chestnut 
spots on the dorsum, with emerald green and purple bars alter- 
nating on the sides of the head and limbs (C. megastoma) . 

Range. ?Venezuela. Surinam, Brasil, Bolivia, Ecuador. 

Remarks. There is no doubt that the color description given 
by Rohl is not that of Ceratophrys calcarata nor of Eleuthero- 
dactylus cornutus maussi, the other species with which it could 
be confused. I feel, however, very uncertain about the identity 
of this animal and about its presence in Venezuela. The A.M.N.H. 
has British Guiana specimens under this name (not examined) 
and its occurrence in this country has been confirmed bv Craw- 
ford (1931:11). 

Like Boulenger (1882), Crawford speaks only of a grayish 
color, the head and back with dark markings and the flanks 
marbled with brown. I believe C. calcarata may extend to Brit- 
ish Guiana, and it is not improbable that these records may refer 
to that species and not to C. cornuta. 

According to Rohl the common name for this species in Vene- 
zuela is "rana cornuda." 

PSEUDOPALUDICOLA PUSILLA (Rllthven) 

Paludirola pusilla Buthven, 1916, Occ. Papers Mus. Zool. Univ. Mich., No. 

30: 1-3: Fundaeion, Colombia. 
PahtdicoJa exigua Boettger, 1896, Ber. Senekenb. Naturf. Ges. : LIV. 
12 (IT.P.R. 23745, 247-9) Pto. Ayacucho, vi.50. 
2 (U.P.R. 252, 254) Pto. Ayacucho, vi.50. 
Description. Snout subovoid or subelliptieal ; tongue narrow, 
somewhat pyriform, entire and free behind ; eye diameter greater 
than distance between eye and nostril but shorter than the snout ; 
interorbital space equal to, or a little broader than an upper eye- 
lid ; canthus rounded ; loreal oblique, concave ; tympanum hidden ; 



RIVERO : SALIENTIA OP VENEZUELA 87 

one distinct and one or two other less prominent tubercles be- 
tween elbow and wrist; subarticular tubercles large, rounded, 
that of the inner finger much larger than the others ; fingers free, 
first shorter than second; tips of fingers swollen; usually a 
tubercle at the heel; an oblique tarsal fold that commences with 
a tubercle at the middle of the tarsus ; metatarsal tubercles small 
but very prominent, the outer commonly pointed ; subarticular 
tubercles of toes prominent; toes with a short web that extends 
to the small disks as distinct lateral fringes ; heel of the adpressed 
hind limb extends to the middle or anterior corner of the eye. 
Skin above, generally rugose and warty (apparently smooth in 
females) ; a flat, oblique and indistinct fold from lower eyelid 
to base of the humerus. Below, smooth. Male with two slight 
subgular vocal sacs. 

Color. Above, brown or dark grayish-brown, generally with 
darker spots or large yellowish blotches that may cover most of 
the dorsum ; an occasional specimen with a greenish yellow 
vertebral stripe ; upper lip vertically spotted with white ; a dark 
longitudinal line along the posteroventral part of the thighs. 
Below, white, immaculate or speckled with brown, especially on 
the throat and limbs. 

Measurements. Snout-vent S 12, $ 17 ; head breadth S 4.6, 
2 5 ; head length <£ 4.9, $ 6 ; femur $ 5.5, $ 7 ; tibia $ 5.7, 
9 7. 

Habits. All the 12 males were caught on the sandy shores of 
a stream. There was small and sparse forest growth on the 
margins, deep forest upstream, and savannas nearby. The two 
gravid females were found in a grassy savanna near a large 
granite outcropping. For a good illustration of the habitat of this 
animal and probably the exact location where the two females 
were collected, the reader is referred to Hitchcock (1947, fig. 13). 

The tremendous jumps that this little animal is capable of 
performing make it extremely difficult to catch. Sometimes on 
opening one's hand expecting to see the frog, one finds only dust 
and leaves, for the little animal is jumping several feet away. 

Additional Localities. Pto. Ayacucho nr. Venado (U.S.N.M. 
80674) ; nr. Rosario (U.S.N.M. 115760-7) ; Upper Orinoco (Boett- 
ger, 1896). 

Range. The Venezuelan Guayana (lower extension of the 
Llanos into Pto. Ayacucho), the arid and semiarid Maracaibo 
Basin and probably the Llanos, the Falcon Region and the 
coastal belt. Northeastern Colombia across central and southern 



88 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Venezuela to British Guiana. It probably occurs also in the 
Llanos of Colombia, the savanna country of the eastern Guiana 
and in N. Brasil. 

Remarks. The two females from Pto. Ayacucho have a more 
pointed and longer snout than the males, and since both of them 
are full of eggs, the heel scarcely reaches the shoulder. 

Very little difference has been found between this species from 
southern Venezuela and a paratype from Colombia. There is 
only one tubercle on the forearm of the latter. 

Six specimens from Inirida in the Senckenberg Museum 
(Paludicola exigua Boettger, 1896) were found to represent this 
species. 

Paludicola pischeri Boulenger 

PaludicoJa fischeri Boulenger, 1890, Proc. Zool. Soc. London: 327, pi. xxv, 

fig. 2: Venezuela; Nieden, 1923, Das Tierreich. Anura I: 512. 
Paludicola Fisdheri Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 39, 46. 

No material examined. 

Description. Parker (1927:473) has placed Paludicola 
fischeri under the synonymy of Physalaemus gracilis Blgr., stat- 
ing that he has been unable to find any character by which to 
distinguish the two species or any data to confirm the locality. 
Since Physalaemus gracilis is a southern form not likely to occur 
in Venezuela, it should not be included for the present in the 
fauna of this country. A Physalaemus 1 has been taken by 
Dunn at Villavicencio and Llanos de Boyaca of Colombia (1944a : 
512). Probably this form will also be found in Venezuela. 

Pleurodema brachyops (Cope) 

Pleurodema bibroni var. B. Giinther, 1858, Cat. Batr. Sal. Brit. Mus. : 32. 
Lystris brachyops Cope, 1868, Proc. Acad. Nat. Sci. Pliila. : 312: Magdalena 

B., Colombia. 
Pleurodema Sachsi Peters, 1877, Monatsb. Berl. Akad. Wissensch. : 460. 
Paludicola brachyops Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 232; 

Boettger, 1893, Ber. Senckenb. Naturf. Ges.: 40. 
Pleurodema brachyops Nieden, 1923, Das Tierreich, Anura, I: 499; Lutz, 

A., 1927, Mem. Inst, Osw. Cruz, 20: 39, 46, pi. 11, fig. 17; Parker, 

1927, Ann. Mag. Nat. Hist., (9) 20: 473; Schmidt, K. P., 1932, Zool. 

Ser. Field Mus. Nat. Hist,, 18: 160. 
2 (U.S.N.M. 115757-8) Campo del Lago, Lagunillas, iv.42. 
1 (U.S.N.M. 36371) Chicara (Caicara?), 05. 

1 I have been unable to find to what species Dunn referred. 



RIVERO: SALIENTIA OP VENEZUELA 89 

5 (U.S.N.M. 117525, 128841-4) Sierra de Sta. Ana, Falcon, 39. 

1 (U.S.N.M. 128840) Palenque, iii.39. 

Description. Snout short and rounded; nostrils closer to the 
tip of the snout than to the eye ; tongue oval, entire or indis- 
tinctly nicked behind ; vomerine odontoids in two short, oblique 
groups between the choanae, their posterior extremities directed 
inwards; eye diameter as long as the snout; interorbital space 
equal or narrower than an upper eyelid ; canthus rounded ; loreal 
almost vertical, not concave ; tympanum not very distinct, about 
y 2 the eye diameter; metacarpal tubercles oval, distinct; fingers 
free, the first a little longer than second which may be slightly 
longer than fourth ; subarticular tubercles of fingers and toes 
very prominent; tips of fingers swollen; two sharp, inwardly 
directed, shovel-shaped, metatarsal tubercles; toes with a slight 
web that extends to the tips as thick lateral fringes; heel of the 
adpressed hind limb extends to the shoulder. Skin above, with 
scattered small warts and short glandular ridges ; two prominent 
and relatively large lumbar glands. Below, smooth. Male with 
a subgular vocal sac. 

Color. Above, light yellowish brown or gray, marbled and 
spotted with darker brown ; a dark spot under the eye ; some- 
times a vertebral stripe ; lumbar glands blackish brown, spotted 
with pink (red?) ; inguinal region, underside of the tibiae and 
a transverse spot at the knee, pink. 

Measurements. Snout-vent $ 31.5, $ 34.5 ; head breadth 
<$ 13, $ 13.5; head length $ 10.5, 9 10.5 ; femur $ 12, $ 11; 
tibia $ 12.5, $ 13. 

Additional Localities. Caracas (Boettger, 1892) ; Cocollar 
(C.N.H.M. 17783 [^Schmidt, 1932]); Distrito Acosta (C.N.H.M. 
25898) ; Maracay (Lutz, 1927) ; Sn. Fdo. de Apure (Peters, 
1877) ; Venezuela (U.S.N.M. 36372-5; Giinther, 1858; Boulenger, 
1882; Boettger, 1893; Nieden, 1923). 

Range. The arid and semiarid Maracaibo Basin, the coastal 
belt, the Coastal Range and the Llanos. Northeastern Colombia 
and the islands north of Venezuela to British Guiana and north- 
ern Brasil. Dunn (1944) says it also occurs in Panama. 

Remarks. There seems to be considerable variation in the 
coloration of this form. Some have a canthal streak, others verti- 
cal spots on the upper lip, a vertebral stripe or black color ex- 
tending outside of the lumbar gland. In the specimen from 
Cocollar there are a very few light spots in these glands. I failed 
to detect vomerine teeth in U.S.N.M. 115758. 



90 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

Eupemphix pustulosus BUTHVENi Netting 

Engystomops pustulosus Boettger, 1892, Kat. Batr. Samm. Mus. Senckeub. : 

33. 
Eupemphix pustulosus Boettger, 1893, Ber. Senckenb. Naturf. Ges.: 40; 

Aleman, 1952, Mem. Soc. Ciene. Nat. La Salle, 12: 26, fig. 6. 
Eupemphix pustulosa Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 38, 42, pi. 

xii, figs. 25, 26. 
Eupemphix ruthveni Netting, 1930, Ann. Carneg. Mus., 19: 167, pi. vii, 
fig. 1 : Fundaeion, Colombia. 

1 (U.M.M.Z. 55555) La Fria, Pueblo Nuevo. 
1 (U.M.M.Z. 57398) Sta. Elena, Merida. 
Description. Snout triangular; tongue small, narrow, entire 
and free behind; choanae small, round; eye diameter more or 
less equal to distance between eye and nostril ; interorbital space 
about iy 2 times broader than an upper eyelid ; tympanum very 
indistinct, almost hidden ; parotid triangular, mostly lateral and 
not very distinct ; canthus angular ; loreal vertical, not concave ; 
subarticular tubercles quite distinct and prominent ; fingers free, 
the first slightly shorter than second; fingers and toes distally 
swollen; a conical tubercle on the middle of the tarsus; meta- 
tarsal tubercles moderate, sometimes pointed; toes practically 
free; heel of the adpressed hind limb extends to the eye. Skin 
above, covered with round and elongated warts; a small lateral 
gland on each side between axilla and groin. Belly slightly 
granular. Male with a large subgular vocal sac. 

Color. Above, brown or dull, dark gray with obscure darker 
markings that occasionally follow some of the rows of warts; a 
short, greenish-yellow vertebral stripe from tip of urostyle to 
sacrum or slightly beyond ; limbs spotted or crossbarred. Throat, 
chest and anterior part of the belly infuscate, with a median 
greenish-yellow line that extends to the light-colored hinder por- 
tion of the belly ; this is marked with distinct round spots. The 
ventral surface of the hind limbs may be slightly spotted. 

Measurements. Snout-vent 3 25, 9 28.5 ; head breadth $ 8, 
9 9 ; head length $1,9 7.5 ; femur $ 10, ? 11 ; tibia $ 11, 
9 11.5. 

Additional Localities. Caracas (Boettger, 1892) ; Caripito 
(U.S.N.M. 11792-7) ; El Sombrero (TJ.C.V. 34) ; Espino (U.C.V. 
67); Maraeav (Lutz, 1927); Ocumare del Tuy (U.C.V. 7-8); 
Pie del Cerro (U.S.N.M. 121149) ; Puerto Cabello (Boettger, 
1893) ; Sta. Lucia (U.S.N.M. 121150-1) ; Turgua (Aleman, 1952). 



RIVERO : SALIENTIA OF VENEZUELA 91 

Remarks. In ventral coloration the two specimens from Mara- 
caibo Basin are somewhat intermediate between the typical form 
and the animal that occurs in central and eastern Venezuela. 
The latter are usually smaller and their bellies uniformly colored 
behind. I am uncertain whether the Venezuelan form deserves 
to be separated from E. p. trinitatis, but Netting has examined 
quite a number of specimens and insists on their distinctness. 
Probably the Trinidad frogs are not more divergent from the 
animals of eastern Venezuela than these are from those of the 
west. 

The male specimen measured (U.C.V. 34) is not one of the two 
specimens on which the description was based. Except for the 
darker color of the external vocal sac, this animal is plain below. 
It does not have ventral or vertebral stripes. In some examples 
the venter is smooth while in those from north central Vene- 
zuela the dorsal stripe does not reach the sacrum and may be 
very short or even absent. 

PSEUDIDAE 

Pseudis paradoxus (Linne) 

liana paradoxa Linne, 1758, Syst. Nat., ed. 10: 212: Surinam. 
Pseudis paradoxa Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 39, 44, pi. xi, 
figs. 18, 19. 
No material from Venezuela. 

1 (M.C.Z. 2557) British Guiana. 

2 (M.C.Z. 12135-6) Coast Lands, Demerara, British Guiana. 
Description. Head flat, much broader than long; snout short, 

rounded to subovoid ; tongue broad, rounded, entire or slightly 
nicked and free behind ; vomerine odontoids in two prominent 
transverse groups between the small choanae; eyes small, their 
diameter equal to distance between eye and nostril; interorbital 
space at least twice as broad as the narrow upper eyelid ; tym- 
panum not very distinct, % or more of the eye diameter ; canthus 
flat and indistinct ; loreal sloping, concave ; no prominent supra- 
tympanic fold ; a row of small tubercles along the forearm ; 
fingers free, the first opposed to the others; subarticular tubercles 
of the fingers small, lint larger and more distinct than those of 
the toes; a prominent tarsal fold; a very prominent, almost 
spnrlike (not cornified), inner metatarsal tubercle, the outer 
absent or indistinct; toes broadly webbed to the tips; tips of 
fingers and toes slightly expanded ; heel of the adpressed hind 



92 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

limb extends to the eye. Skin above, smooth, with some glandules 
on the flanks. Below, smooth, including the thighs; a distinct 
fold at the knee and another at the heel. 

Color. Above, yellowish brown or dark brown with peculiar 
elongated spots and occasionally minute points ; limbs with large 
dark spots; flanks and posterior part of the thighs with yellow 
spots. Below, yellow, with scattered, small, brown spots on the 
belly and larger more elongate ones on the throat and chest ; an 
oblique, long, brown line or spot on each side of the breast; 
thighs with pretty, longitudinal spots ; tibiae and tarsi marbled ; 
two innermost toes usually yellow. 

Measurements. Snout-vent $ 57, 9 66 ; head breadth S 20, 
9 24; head length $ 18, 9 19; femur S 31, 9 35; tibia, 

<$ 32, 9 36. 

Habits. According to Lutz, these frogs "rarely come out of the 
water where they are not easily caught, though they can be 
taken by angling. The call of P. paradoxus is a characteristic 
croak." 

Additional Localities. Maracay (Lutz, 1927). According to 
Ditmars (The Making of a Scientist, 1938: 245) tadpoles of this 
species were discovered in "certain lakes of Venezuela." I have 
not found any reference to this collection, but Savage and 
Carvalho (1953: 195) mention eastern Venezuela as part of the 
range of the species. 

Range. Maracay and E. Venezuela. Trinidad and the 
Guianas, south to Paraguay and Argentina. 

Remarks. The genus Pseudis was originally included in the 
Leptodactylidae. In 1935 Parker suggested that, on account of 
its possessing an extra phalanx in each digit (which may be an 
enlarged intercalary phalanx), Pseudis should probably be in- 
cluded in the Hylidae. In a recent paper, Savage and Carvalho 
(1953) have erected a new family for those frogs with an extra 
phalanx in their digits. 

HYLIDAE 

Key to the Genera of Venezuelan Hylidae 

I. Pupil horizontal ; toes webbed. 

A. Female with a dorsal pouch in which the eggs are carried, derm of 
head involved in cranial ossification or, if not, first finger equal 
or longer than second. 

1. Size small (25 mm.) ; dorsal pouch opening by a longitudinal 
slit; fingers free; first finger longer than second Nototheca 



RIVERO : SALIENTIA OF VENEZUELA 93 

2. Size large (50 mm.) ; dorsal pouch opening by two longitudinal 
or crescent shaped slits on the posterior half of the dorsum ; 

fingers webbed Gastrotheca 

B. Female without a dorsal pouch; first finger shorter than second (ex- 
cept in Hyla marahuaquensis) . 

1. Derm of head involved in cranial ossification . Corythomantis 

2. Derm of head free Hyla 

II. Pupil vertical Phyllomedusa 

Key to the Species of Hyla Reported from Venezuela 

I. Choanae large, vomerine odontoids in two groups forming together a 
/ \ or a / \.i 

A. Outer finger with not more than 2 free phalanges. 

1. An external rudiment of pollex. 

a. Outer finger fully webbed (1 phalanx = disk free) ; tym- 
panum % the eye diameter; heel to between eye and nostril 
or tip of the snout ; brown above ; a row of dark spots along 
the flanks ; 75 mm boons 

b. Outer finger with l 1 /* free phalanges; tympanum % the eye 
diameter; heel to the tip of the snout; purple brown with 
dark scribblings above ; lower eyelid with metallic venation ; 
bones green; 75 mm wavrini 

c. Outer finger with 2 free phalanges; tympanum % the eye 
diameter; heel to tip of the snout or beyond; a pair of 
dorsolateral folds from eyes to groin; apple green (when 
alive) or closely dotted with brown (preserved) above; 50 
mm albomarginata 

d. Outer finger with 2 free phalanges ; tympanum *4 the eye 
diameter ; heel to anterior corner of the eye or between eye 
and nostril but not beyond ; no dorsolateral folds ; green 
(when alive) occasionally with peculiar brown spots or yel- 
lowish white (preserved) above; 40 mm granosa 

2. No distinct rudiment of pollex. 

a. Outer finger with l 1 /* to 2 free phalanges; tympanum % the 
eye diameter ; heel to tip of snout ; snout broad but pointed ; 
usually an X-shaped figure on the anterior part of the dor- 
sum; 60 mm geographica geographica 

B. Outer finger with more than 2 free phalanges (a basal or rudimen- 
tary web). 

1. Tympanum distinct, Va or more the eye diameter, 
a. An external rudiment of pollex. 

1. Tympanum at least % the eye diameter; snout mod- 
erate, subovoid ; loreal region moderately oblique; 



i In //. paralitica, 11. juhni and //. platydactyla the vomerine teeth lire very 
slightly oblique and on occasion may be almost transverse. 



94 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

larger disks about V2 the size of the tympanum ; light 
brownish above, usually with a large, diffuse, darker 
spot; flanks and thighs with fine crossbars suv^idQ.iO' ' 

2. Tympanum % the eye diameter; snout long, subellip- 
tical ; loreal region almost vertical; brown, usually 

with transverse bands above 

albopunctata midtifaseiata 

3. Tympanum % the eye diameter; snout short, sub- 
ovoid; largest disks larger than the tympanum; 
marbled with reddish brown and dark gray above ; 
flanks with small, white spots loveridgei 

b. No distinct rudiment of pollex. 

1. Tympanum % the eye diameter; largest disks about 
% the size of the tympanum ; brown, usually vermicu- 
lated with darker above ; a supra-tympanic streak that 
extends for a short distance posteriorly; thighs with 
distinct, broad crossbars raniceps 

2. Tympanum almost as large as the eye; a pair of lon- 
gitudinal ridges on the head (margins of the fronto- 
parietals elevated) ; largest disks almost as large as, 
or larger than, the tympanum ; profusely ( $ ) or 
sparsely ( 9 ) tubercular above; male with two lateral 
vocal sacs ; brown above, sometimes with darker mark- 
ings taurina 

2. Tympanum indistinct or absent; where present, not quite Vi the 

eye diameter (usually %). 

a. Snout almost semicircular; vomerine odontoids very slightly 
oblique ; canthus not well defined ; no distinct supratympanic 
fold; above, whitish, usually with scattered melanophores; no 
light canthal or supratympanic streaks; usually orange 
drab below; 31 mm.; Andes paramica 

1). Snout subovoid ; vomerine odontoids slightly oblique; a supra- 
tympanic fold ; above brownish, closely punctulated with 
darker brown or white (after long preservation) or uniformly 
whitish ; a light canthal, palpebral and supratympanic lines ; 
34 mm. ; Andes jahni 

c. Snout subovoid ; vomerine odontoids slightly oblique ; a 
supratympanic fold; above, purple, closely dotted with 
darker; arms and thighs white, with a stripe of closely set 
dark purple dots; 32 mm.; Andes platydactyla 

d. Snout subovoid ; vomerine odontoids distinctly oblique ; a 

distinct rudiment of pollex; color above brownish with darker 
transverse bands or uniform and with reticulations; ventral 
surfaces orange (when alive) or whitish (preserved) ; 37 

mm. ; Guayanan Cerros bcnitczi 

II. Choanae small or moderate; vomerine odontoids in two groups, trans- 
verse or obliquely directed backwards and inwards. 



RIVERO : SALIENTIA OF VENEZUELA 95 

A. Fingers free. 

1. First finger slightly longer than second. 

a. A light dorsolateral line that meets its opposite in a slight, 
backward notch between the eyes; 27 mm.; Gnayanan Cerros 
mardhuaquensis 

2. First finger shorter than second. 

a. Tympanum % to % the eye diameter; snout long, subovoid ; 
heel to the eye ; above, brown or gray, sometimes with two 
long, longitudinal spots behind the eyes and white spots on 
the anterior and posterior part of the thighs ; 38 mm. . . 
rubra, 

b. Tympanum % the eye diameter ; snout very long, subacumi- 
nate ; heel to between eye and nostril or tip of the snout ; 
brown or dark gray with a triangular spot between the eyes 
and usually other dark markings on the scapular region; 
thighs with alternating black and reddish bars on the an- 
terior and posterior aspects; 40 mm boulcngeri 

c. Tympanum Y? the eye diameter ; snout subovoid ; heel to the 
eye; above, chlorine green (when alive) or yellowish gray 
(preserved) ; anterior and posterior part of the thighs of 
uniform color; 28 mm baitmgardneri 

B. Fingers webbed. 

1. Tympanum absent. 

a. A white or yellow spot below the eye; upper surface of the 
thighs with white or yellow spots; 31 mm. luteocellata 

2. Tympanum present, y 2 or more the eye diameter. 

a. Outer finger fully webbed (1 phalanx = disk free). 

1. Snout almost semicircular; odontoids in an inverted chev- 
ron; forearm and tarsus with a festooned fringe; above, 
marbled with different shades of gray and brown; below 
orange (when alive) or white (preserved) with round, 
black spots; 35 nun m. marmorata 

b. Outer finger not fully webbed. 

1. Tympanum % the eye diameter; snout rounded; loreal 
sloping ; heel to the eye ; skin with scattered glandules 
and tubercles especially in the surroundings of the tym- 
panic region ; male with two vocal sacs at the angle of 
the mouth. 

a. Size very large ; color above more or less uniform ; 
skin very thick and glandular; Maracaibo Basin 
tibiatrix ingens 

b. Size smaller; color above not generally uniform; skin 
not as thick and glandular as above; Venezuela, ex- 
cept Maracaibo Basin tibiatrix tibiatrix 

2. Tympanum % the eye diameter; snout rounded; heel to 
the eye; skin above smooth; above, slate or bronze col- 
ored; anterior and posterior part of the thigh and lower 



96 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

surface of the hind limbs spotted -with white; 42 mm.; 
Andes vilsoniana meridensis 

3. Tympanum % the eye diameter; snout almost semicircu- 
lar; eye diameter greater thnii distance between eye and 
nostril; heel to between eye and nostril; no tarsal fold; 
skin above, smooth ; above, straw colored, generally with 
ornate, light margined markings of irregular shape; no 
dark canthal or supratympanic streaks; a transverse, dis- 
tinct, light line above the anus and another longitudinal 
one at the heel; 26 mm.; Venezuelan Guayana minuta 

4. Tympanum Vj the eye diameter; snout rounded; eye 

diameter greater than distance between eye and nostril ; 
a tarsal fold; heel to between eye and nostril; skin 
above, smooth ; brownish with obscure longitudinal mark- 
ings ; a canthal and a supratympanic streak ; no distinct 
light line above the anus or at the heel; 20 mm.; Coastal 
Eange misera 

5. Tympanum % the eye diameter ; snout pointed and pro- 
jecting; loreal vertical; a slight tarsal fold; heel to 
anterior corner of the eye; skin above, shagreened, yel- 
lowish white in preservation; two horizontal flaps at the 

sides of the anus; 24 mm.; Venezuelan Guayana 

orophila plan icola 

6. Tympanum % the eye diameter; snout rounded; heel to 
between eye and nostril ; skin above, smooth ; above and 
on the ventral surface of throat and limbs distinctly 
dotted with brown; 33 mm.; Coastal Eange battersbyl 

Hyla boans (Linne) 

liana boons Linne, 1758, Syst. Nat., ed. 10: 213: America. 

Hyla maxima Boettger, 1896, Ber. Senckenb. Naturf. Ges., LIY ; Fowler, 

1913, Proc. Acad. Nat. Sci. Phila., 65: 170; Lutz, A., 1927, Mem. Inst. 

Osw. Cruz, 20-. 40, 47. 

1 (U.P.R. 109) La Culebra, 1000 ft., iv.50. 

1 (U.P.R. 110) Cano Chupadero, iii.50. 

1 (U.P.R. Ill) Anaben, Colombia, vi.50. 

1 (M.C.Z. 19918) Orinoco R., below Caroni, i.35. 
Description. Head depressed; snout subovoid ; the nostrils 
somewhat elevated and forming a swollen tip ; tongue subcircu- 
lar, adherent and entire ; vomerine odontoids forming a / \ 
figure behind the large ehoanae ; eye diameter shorter than dis- 
tance between eye and nostril ; interorbital space concave, much 
broader than an upper eyelid ; canthus obtusely angular ; loreal 
sloping, concave ; tympanum % the eye diameter, its postero- 
dorsal margin slight! v hidden under the skin ; a dermal fold 



RIVERO: SALIENTIA OF VENEZUELA 97 

along the forearm ; an external rudiment of pollex ; fingers taken 
in order from first to fourth exhibit the following phalanges 
free of web : 2 to 2^4, 1, 1 to 1%, 1 ; first finger shorter than 
second ; larger disks about % the size of the tympanum ; a tri- 
angular dermal appendage at the heel; two tarsal folds, the 
outer extending to disk of the fifth toe ; no outer metatarsal 
tubercle ; toes fully webbed ; heel of the adpressed hind limb 
extends to between eye and nostril. Skin above finely granular; 
throat smooth or slightly granular. 

Color. Above, brown with darker, irregular markings; flanks 
with short, dark crossbars or spots. Ventral surfaces white or 
greenish white ; thighs posteriorly, dark brown, usually with ob- 
scure darker or lighter mottlings; hind limbs, especially the 
anterior part of the thighs, occasionally with diffuse crossbars. 

Measurements. Snout -vent $ 75, 9 105 ; head breadth $ 28, 
9 40 ; head length $ 27, 9 3(3 ; femur $ 41.5, 9 59 ; tibia $ 42, 
9 58.5. 

Habits. All the U.P.R. specimens were collected at night on 
the trunks of trees on heavily forested areas. Of three stomachs 
examined, two were empty and one had three grasshoppers of 
the same species. 

Additional Localities. Arabopo (U.M.M.Z. 85162 [4], 85163, 
85164 [2], 85165); Auyantepui (A.M.N.H. 39751); Casiquiare 
Canal, nr. Venado (U.S.N.M. 80661-4) ; Cuquenam Valley 
(U.M.M.Z. 85167 [2]) ; Esmeralda (A.M.N.H. 23223) ; Manamo, 
Cano (Fowler, 1913) ; Paulo (U.M.M.Z. 85161) ; Sn. Fernando 
de Atabapo (U.S.N.M. 80652) ; Upper Orinoco (Boettger, 1896). 

Range. The Venezuelan Guayana. A few specimens (U.M.M.Z. 
55558-66) from the Coastal Range determined as Hyla boans 
(unfortunately not examined) may be Hyla albopunctata. Co- 
lombia and Trinidad to Bolivia and northern Brasil (Amazonas). 

Remarks. U.S.N.M. 80652 is a large animal (125 mm.) with 
variegated coloration. It possibly represents some other form. 

Hyla faber and Hyla boans may be conspecific. Specimens of 
the latter with less web than usual are found in Amazonia and 
in the eastern base of the Andes of Colombia. 

Hyla wavrini Parker 

Hyla wavrini Parker, 1936, Bull. Mus. Eoy. Hist. Nat. Belgique, 12: 2: 
Upper Orinoco, Territorio Amazonas, Venezuela. 
No material examined. 
Original Description. Head depressed, as long as broad ; snout 
rather pointed, twice as long as the diameter of the eye ; nostril 



98 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

more than twice as far from the eye as from the tip of the snout ; 
tongue not emarginate and almost entirely adherent behind; 
vomerine odontoids in two series which together form a / \ 
between the very large choanae ; interorbital space as broad as an 
upper eyelid; canthus obtusely rounded; loreal oblique and 
somewhat concave ; tympanum very distinct, % the eye diameter ; 
a supratympanic fold; fingers % webbed, the first with a pro- 
jecting rudiment of pollex, much shorter than the second, which 
is shorter than the fourth ; digital disks nearly as large as the 
tympanum ; toes webbed to the disks ; heel of the adpressed hind 
limb extends to the tip of the snout. Skin above, uniformly 
shagreened ; dermal folds along the outer edges of the forearm 
and tarsus and a quadrangular lappet of skin on the heel. Lower 
surfaces granular, coarsely so on the belly and thighs, finely on 
the throat and chest. 

Color. Pale purple brown above, with a median dark line from 
the snout to the middle of the back; rest of the other surfaces 
with a few irregular, dark scribblings. Limbs crossbarred on 
the exposed surfaces ; their concealed surfaces with black, light 
centered, vertical bars. Lower surfaces uniform white. Lower 
eyelid with metallic venation. Bones green. 

Measurements. 9 snout-vent 75 ; forelimb 45 ; hind limb 128. 

Remarks. The paratype is a male with the same data as the 
type; it agrees in essentials, but is slightly larger (78 mm. in 
snout to vent length), has a more prominent pollex, a vocal sac 
opening by a large slit on each side of the tongue, a brown washed 
gular region and almost uniform dorsal coloring. 

The species is obviously allied to Hyla faber AVied and Hyla 
pardalis Spix, but differs from both in its longer, flatter head, 
shagreened dorsal surfaces and green bones (Parker, op. cit.) . 

Hyla albo margin ata Spix 

Hyla albomarginata Spix, 1 824, Spec. Nov. Testud. Ran.: 33, pi. viii: 
Bahia; Lutz, A., 1927, Mem. Inst, Osw. Cruz, SO: 38, 43. 
1 (M.C.Z. 15369) Maracaibo, v.06. 
Description. Snout subovoid ; tongue oval, adherent, slightly 
nicked behind : vomerine odontoids forming a / \ figure be- 
tween the large, oval choanae ; eye large, its diameter slightly 
shorter than distance between eye and nostril ; interorbital space 
about twice as broad as an upper eyelid ; canthus moderate ; 
loreal oblique, not concave ; tympanum distinct, about V2 the 
eye diameter; a longitudinal fold from elbow to hand; an ex- 
ternal rudiment of pollex ; first finger shorter than second ; 



RIVERO : SALIENTIA OF VENEZUELA 99 

fingers taken in order from first to fourth exhibit the following 
phalanges free of web : all, 2, 2i/ 2 , 2 ; heel with a short dermal 
appendage; a tarsal fold; one flat, inner metatarsal tubercle, 
no outer; toes exhibiting the following free phalanges: iy 2 , 
1, iy 2 , 2y 2 , iy s ; disks of the toes smaller than those of the 
fingers; heel of the adpressed hind limb extends to slightly 
beyond the tip of the snout. Skin above and on the sides of 
the head finely granular; a pair of dorsolateral folds from 
posterior corner of the eye, along the flanks, to the groins. 

Color. The color of the only Venezuelan specimen available 
is faded and the animal presents a creamy white appearance 
throughout. Brasilian specimens are light brown, closely punc- 
tuated with darker brown. Noble (1918: 342) describes the liv- 
ing animal as green, with bluish throat, lemon yellow belly, 
bluish legs, yellow feet and hands, and orange web. 

Measurements. S snout-vent 51 ; head breadth 18 ; head length 
16.5; femur 27; tibia 29. 

Range. Maracaibo Basin. Central America, Colombia and 
British Guiana to southern Brasil. No records available from 
Ecuador, Bolivia and Pern. 

Remarks. I do not find much difference between the Vene- 
zuelan example and specimens from Pernambuco, Brasil. 

Lutz bases the inclusion of this species in the Venezuelan fauna 
on a specimen which lie says was collected by Robinson. I have 
not found this species mentioned in Stejneger 's report on Lyon 
and Robinson's collection but the specimen reported here un- 
doubtedly belongs to Hyla albomarginata. 

Hyla granosa Boulenger 

Hyla granosa Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 358, pi. 
xxiv, figs. 1, 2: Demerara Falls: Santarem; interior of Brasil and 
Canelos, Ecuador. 

8 (U.P.R. 118-25) La Culebra, 1000 ft., v.50. 
1 (U.P.R. 199) Anaben, Colombia, vi.50. 
Description. Head depressed; snout rounded; nostrils some- 
what elevated and forming a truncate tip ; tongue rounded, ad- 
herent and entire ; vomerine odontoids in two oblique groups 
between the choanae, their anterior extremities convergent; eye 
diameter slightly shorter than distance between eye and nostril ; 
interorbital space about twice as broad as an upper eyelid ; 
canthus indistinct ; loreal strongly sloping, somewhat concave ; 
tympanum not very distinct, about y 2 the eye diameter; an 



100 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

external rudiment of pollex ; subarticular tubercles small, round, 
somewhat depressed ; first finger shorter than second ; fingers 
taken in order from first to fourth exhibit the following phalanges 
free of web : all, 1% to 2, 2y 2 , 2 ; a tarsal fold ; one small, oval 
inner metatarsal tubercle ; toes with the following free phalanges : 
1% to 1%, 1 to iy 3 , 1 to 1%, 1% to 2, iy 2 ; heel of the adpressed 
hind limb extends to the anterior corner of the eye or slightly 
beyond. Skin above and on the sides of the head and body 
minutely granular ; a strong fold across the chest. Male with an 
external, subgular vocal sac. 

Color. Above, green (yellowish white in alcohol) ; occasionally 
a distinctive light brown spot between the eyes or on the back or 
legs ; minute brown points under a lens. Ventral surfaces white. 

Measurements. $ Snout-vent 40 ; head breadth 15 ; head length 
13; femur 19; tibia 16.5. 

Habits. The species was heard in bushes along the banks of 
the Orinoco from a little beyond Sanariapo to the upper reaches 
of Rio Cunucunuma. Sometimes one may walk for several miles 
without hearing the peculiar marimba-like call of granosa until 
attracted by it to a bush that is full of frogs. Usually the area 
occupied does not cover more than a few square yards. Similar 
conditions were observed for several other Hyla in South 
America. 

Range. The Venezuelan Guayana. British Guiana and north- 
ern Brasil, ?Ecuador. 

Remarks. I think it would be advisable to restrict the locality 
of this species but I prefer to leave the problem to someone hav- 
ing access to the typical material. My impression is that Boulen- 
ger's "g" specimen from Canelos may be Hyla punctata, which 
has been found in Ecuador by Peracca (1904). This, of course, 
is not conclusive evidence and the fact that H. granosa has been 
collected in Rio Uaupes somewhat detracts from it ; consequently, 
it is as well not to restrict the type locality until these sugges- 
tions can be verified. 

Melin (1941: 21) has described a new subspecies from the 
Uaupes River in northern Brasil. He characterizes the new form 
by the different coloration (greenish yellow), absence of claw-like 
pollex, more slender build, less webbing and narrower limbs. 
Perhaps Melin ascertained from the British Museum to which of 
Boulenger's specimens the figure corresponds but I do not find 
anything in the literature in this respect. It is even possible that 
the figured specimen came right from the type locality of Melin 's 
Hyla granosa gracilis. 



RIVERO : SALIENTIA OF VENEZUELA 101 

Since Boulenger had preserved material, the first differential 
character of H. granosa gracilis should not be taken into con- 
sideration, while the third and last, which were based on Boulen- 
ger 's figure, might be due to individual differences or to the 
different conditions of preservation. A small spinule is felt in 
the pollex of the males from Venezuela and British Guiana 
(specimens examined: A.M.N.H. 39734, 39985, 45752, 46234, 
49260, 49262; U.S.N.M. 118053-4, all from British Guiana), and 
in all there is some individual variation in regard to the amount 
of webbing. Consequently, I have preferred to retain the bi- 
nomial. 

Hyla granosa resembles Eyla punctata but can be distinguished 
by its laterally situated eyes (in punctata they look somewhat 
to the front), granular skin, large disks, more sloping loreal re- 
gion, smaller tympanum and more extensive webbing between 
the toes. This species is found in British Guiana and in all 
probability occurs in Venezuela. 

Hyla ornatissima Noble is, I believe, a synonym. The ornate 
coloration of the type (vide) is found, though to a lesser extent, 
in several of the specimens of Hyla granosa reported here. 

Hyla geographica geographica Spix 

Hyla geographica Spix, 1824, Spec. Nov. Test. Ran.: 39, pi. xi, fig. 1: 
Teffe River, Amazonas, Brasil; Gimther (part or all?), 1858, Cat. Batr. 
Sal. Brit. Mus. : 99. 

1 (Senckenb. Mus. 2432) Iguapo, Upper Orinoco, v.1895. 

Description. Head as broad as long; snout broad, subovoid; 
tongue oval, adherent and nicked behind; vomerine odontoids 
forming a / \ figure between the large choanae ; ; eye diameter 
equal to distance between eye and nostril; interorbital space 
broader than an upper eyelid ; canthus curved, rounded but well 
defined ; loreal oblique ; tympanum % the eye diameter ; no dis- 
tinct supratympanic fold ; an outer fold between elbow and disk 
of fourth finger ; no apparent rudiment of pollex ; fingers taken 
in order from first to fourth show the following phalanges free 
of web ; all, 1%, 2\< 2 , 2 ; first finger shorter than second ; disks of 
fingers not more than 14 the size of the tympanum ; a pointed 
lappet at the heel ; a distinct tarsal fold ; no apparent metatarsal 
tubercles; toes exhibiting the following free phalanges: 1, 1, iy 2 , 
2y 2 , 1 ; disks of the toes smaller than those of the fingers ; hind 
limbs slender; heel of the adpressed hind limb extends to the 
middle of the snout. Skin above, shagreened. 



102 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Color. Above, light brown, with an irregular, darker marking 
that starts behind the eyes with two horns which unite and 
broaden posteriorly into a more extensive spot ; a dark vertebral 
line from tip of the snout to scapular region; a not too distinct 
spot behind the sacrum; flanks with dark transverse striations; 
thighs with narrow transverse bands and reticulations ; anterior 
and posterior aspect of thighs darker than dorsolateral portions ; 
rest of the limbs crossbarred with alternating broad and narrow 
stripes. Below, brownish. 

Measurements. $ ? snout-vent 40; head length 15; head 
breadth 15; femur 21; tibia 21. 

Additional Localities. Venezuela (Gunther, 1858) ; ?Iguapo 
(Senckenb. Mus. 2590 = Boettger, 1896, II. punetatissima). 

Range. Venezuelan Guayana. According to Parker (1935) : 
Cis- Andean Ecuador and Peru, Amazonas, Para, Guianas and 
Trinidad. 

Remarks. Parker (1935) does not mention the differences be- 
tween Hyla (j. (jeographica and H. g. punetatissima and Boulen- 
ger (1882) had both races in his H. appendicidata. 

For comparison with the only specimen from the north, I had 
material from Sta. Catharina, Brasil, Robore and Buena Vista, 
Bolivia. The Brasilian animals seem to have a straighter canthus 
and a little more web in the fingers. The snout is longer (eye 
diameter shorter than distance between eye and nostril) and in 
most of the larger specimens, broader. Instead of the transverse 
vermiculations of the flanks, the Sta. Catharina animals have 
profuse dark dotting, and except for some ill-defined transverse 
bars on the posterior part of the dorsum of some specimens, or 
occasional black spots, there is no other dorsal pattern. The 
thigh bars are closer together and do not surround white areas 
or spots, and transverse limb bars are not present in several 

specimens. 

The Bolivian specimens show a pattern similar to the Vene- 
zuelan example but the large spots are more abundant and there 
is sometimes profuse dotting of the dorsal surfaces. The limb 
bars are usually very well defined but the flanks sometimes show 
white, black-margined spots instead of bars or dark spotting. In 
M.C.Z. 12869 and 29849 the venter is covered with brown, equi- 
distant and irregularly-shaped spots. The canthus in these speci- 
mens is not well defined and the eyes do not seem to be as large 
or protuberant as in either Brasilian or Venezuelan examples. 
Thev are as different from the northern as from the southern 



RIVERO : SALIENTIA OP VENEZUELA 103 

Brasilian animals and further studies will probably indicate them 
to be racially distinct. 

The juvenile form of H. crepitans is probably punctate above, 
as is that of //. gcographica. 

Hyla crepitans Wied 

Hyla crepitans Wied, 1824, Abbild. Naturg. Bras. : (4) 5, pi. v, fig. 1 : Bahia ; 
Peters, 1877, Monatsb. Akad. Wissensch. Berlin: 460; Boulenger, 
1882, Cat. Batr. Sal. Brit. Mus., ed. 2 : 352 ; Boettger, 1892, Kat. Batr. 
Saniru. Mus. Senckenb.: 40; 1893, Ber. Senckenb. Naturf. Ges.: 40; 
1896, Ber. Senckenb. Naturf. Ges.: LIV; Stejneger, 1902, Proc. U.S. 
Nat. Mus., 24: 181; Boulenger, 1903, Ann. Mag. Nat. Hist., (7) 11: 
481; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 38, 43, pi. xv, figs. 35, 
35a, 36, 36a, 36b; Parker, 1936, Bull. Mus. Boy. Hist. Nat. Belgique, 
12: 2; Schmidt, K. P., 1932, Zool. Ser. Field Mus. Nat. Hist., 18: 160; 
Shreve, 1947, Bull. Mus. Comp. Zool., 99: 536; Rohl, E., 1949, Fauna 
Descr. de Ven., ed. 2: 400, fig. 180; Aleman, 1952, Mem. Soc. Cienc. 
Nat. La Salle, 12: 27. 
Hyla pardalis Gtinther (part or all ? ), 1858, Cat. Batr. Sal. Brit. Mus.: 99. 
6 (M.C.Z. 25978-83) Pauji. 
1 (M.C.Z. 26142) Riecito. 
Description. Head broader than long; snout broad-subovoid ; 
tongue oval or subcircular, adherent, entire or emarginate be- 
hind; vomerine odontoids forming a / \ figure between the 
large, oval choanae ; eye diameter equal or slightly shorter than 
distance between eye and nostril ; interorbital space more or less 
equal to an upper eyelid ; canthus rounded ; loreal moderately 
oblique, not concave in fresh specimens; tympanum % the eye 
diameter ; a supratympanic fold ; an outer, sometimes indistinct, 
fold from elbow to disk of fourth finger ; an external rudiment 
of pollex ; fingers taken in order from first to fourth exhibit the 
following phalanges free of web : all, 1% to 2, 2 to 2^2> 2% to 
2!/2 ; nrs t finger shorter than second ; disks of the fingers not 
more than y 2 the size of the tympanum ; a slight tarsal fold ; 
outer metatarsal tubercle absent ; toes exhibiting the following 
free phalanges : 1 to iy 2 , 1 to IVi, 1V4 to iy 2 , 2y 2 to 2%, 1 to 
1V4 ; disks of the toes smaller than those of the fingers ; heel of 
the adpressed hind limb extends to the tip of the snout or slightly 
beyond. Skin above, smooth. Male with a subgular vocal sac. 

Color. Above, light brown or creamy, generally with a large, 
central, diffuse spot ; flanks lighter, with narrow transverse lines 
on the posterior half; thighs with narrow transverse bars that 
sometimes broaden and become more diffuse above ; tibial segment 



104 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

uniform or with irregular, diffuse spots. Below, yellowish gray 
or white, the throat usually speckled with dark. 

The color of the living animal was described by Kugler 
(Shreve, 1947): "Above white, almost silver white, with large 
black eyes, whose grayish eyelids have a fine black ring around 
the base ; finger tips and belly light yellow. One pair taken in 
embrace had brownish markings on a yellow ground, another 
mating pair exhibited brown marblings exactly the shade of the 
soil." 

Measurements. Snout-vent $ 62, 9 65; head breadth $ 22, 
9 22 ; head length S 20.5, 5 21 ; femur S 33, 9 31 ; tibia S 33, 
9 33. 

Habits. Of three stomachs examined two were empty and one 
contained a cricket and a beetle. 

In the M.C.Z. Pauji group (iv-xi.44), one female is full of eggs. 

Additional Localities. Albarregas, Rio (A.M.N.H. 10521-4) ; 
Arabopo (U.M.M.Z. 85192-4, 85196[2]) ; Baruta (Aleman, 
1952) ; Bejuma (U.M.M.Z. 55564) ; Boca de Rio (U.S.N.M. 

128790) ; Calabozo (Peters, 1877) ; Caracas (U.S.N.M. 55329-30, 
120091, 129264; U.C.V. 821; Giinther, 1858; Boettger, 1882); 
Caicara (A.M.N.H. 16907-10; U.S.N.M. 36376); Caserio Silva 
(U.M.M.Z. 55565) ; Chama (A.M.N.H. 10632-3) ; Cuquenan Val- 
ley (U.M.M.Z. 85189[2]) ; El Limon (U.S.N.M. 12159-61); El 
Periquito (U.C.V. 91-5); El Valle (C.N.H.M. 26188; U.S.N.M. 

128791) ; Espino (U.C.V. 53, 521) ; Giiiria, Peninsula de Paria 
(M.C.Z. 23024-8) ; Kunana, Perija Mts. (Aleman, 1953) ; La Cruz 
Pruviera (U.S.N.M. 72757-8) ; La Guaira (U.S.N.M. 22543-4 
[= Stejneger, 1902] ; Los Canales (U.S.N.M. 128787-9) ; Macuto 
(U.C.V. 79); Maracay (Lutz, 1927); Merida (M.C.Z. 2525; 
C.N.H.M. 3569-70; Boulenger, 1903) ; Ocumare del Tuy (U.C.V. 
49); Palma Sola (U.M.M.Z. 55563); Parmana (U.C.V. 50); 
Paulo (A.M.N.H. 39756) ; Petare (U.S.N.M. 121162) ; Pie del 
Cerro (U.S.N.M. 121154-7) : Puerto Cabello (Boettger, 1893) ; 
Sn. Esteban (U.M.M.Z. 55558-62) ; Sn. Juan de los Morros 
(U.S.N.M. 72756) ; Sierra Maestra, 1,260 m. (U.S.N.M. 129575) ; 
Sosa (U.C.V. 87); Sta. Lucia (U.S.N.M. 121152-3); Turgua 
(Aleman, 1952) ; Upper Orinoco (Boettger, 1896 ; Parker, 1936) ; 
Urama (C.N.H.M. 26185, 29184-5) ; Yumarito (U.M.M.Z. 55.166) ; 
Venezuela (C.N.H.M. 28115-6; Giinther, 1858); Zaraza (T T .C.V. 
30). 

Range. Probably all the physiographical provinces of Vene- 
zuela including the Andes to 1,600 m., Central America, Colom- 



RIVERO : SALIENTIA OF VENEZUELA 105 

bia, Trinidad and the Guianas to southern Brasil. No records 
available from Ecuador, Bolivia, and Peru. 

Remarks. I do not find much geographic variation in this 
species from Panama to Minas Gerais, although there is much 
individual variation everywhere. A dark vertebral line is occa- 
sionally present in specimens from any locality and the throat is 
profusely infuscated in fresh males. In specimens from the 
Magdalena Valley, Colombia (M.C.Z. 21475-83), the head is 
narrower than usual and there is fine granulation on the an- 
terior third of the body. In M.C.Z. 2204 from Trinidad the 
snout is very broad and flat while M.C.Z. 26655 from Sao Paulo, 
Brasil, has dark dots over the dorsal surface. Apparently this 
last animal is not identical with H. crepitans. The specimens 
from La Cruz Pruviera reported here show broad bars that bifur- 
cate on the posterior part of the thighs. 

Specimens from Casa de Julian (U.P.R. 130-1) are provision- 
ally included here although I have little doubt that they belong 
to some other form. The tympanum is about the same size as 
the eye (as it is in some H. crepitans), the heel extends to between 
eye and nostril and the color alive was distinctly different from 
that described by Kugler and others for the living H. crepitans. 
The dorsum in both specimens was dark green, the flanks light 
green, transversely striped with brown ; loins blue ; ventral sur- 
face of the limbs greenish blue and the throat yellow. Both 
specimens were collected at night upon being guided by their 
short, sporadic "crah crah." One of them was on the ground 
near a shallow stream; the other was sitting on the leaf of a 
Taro plant. These specimens agree in almost all structural 
characters with II. crepitans but their size is considerably smaller 
(the larger 46.5 mm.) than any adult males of crepitans observed 
and even the color in preservation is different. The type of Hyla 
indris Cope which is supposed to be very similar to crepitans has 
not been found. 

Several specimens from Venezuela and British Guiana that 
were determined as Hyla pardalis in various museums have 
proven to be Hyla crepitans. 

Hyla albopunctata multifasciata Giinther 

Hyla multifasciata Giinther, 1858, Cat. Batr. Sal. Brit. Mus.: 101, 146, pi. 

viii, fig. I) : Para. 
?Hyla rostrata Peters, 1863, Monatsb. Akad. Wissensch. Berlin: 466. 
2 (U.M.M.Z. 85178) Arabopo. 



106 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

2 (U.M.M.Z. 85179) Arabopo. 

1 (U.M.M.Z. 85180) Cuquenam Valley. 

Description. Snout long, subelliptical ; tongue narrow, oval, 
adherent, entire or emarginate behind ; vomerine odontoids in a 
/" ~\ series between the large, oval choanae ; eye diameter 
shorter than distance between eye and nostril ; interorbital space 
broader than an upper eyelid; canthus distinct; loreal almost 
straight, concave ; tympanum about % the eye diameter ; a supra- 
tympanic fold from posterior corner of the eye to above the 
shoulder or farther back along the flank; a longitudinal fold 
along the forearm; a not very prominent rudiment of pollex; 
subarticular tubercles very prominent; first finger shorter than 
second, practically free; others with a short basal web; disks 
narrow, much smaller than the tympanum ; hind limbs very long 
and slender ; a slight tarsal fold ; toes taken in order from first to 
fifth exhibit the following phalanges free of web : 1%, 1, 1%, 2y 2 
to 3, 1 to l!/4 ; heel of the adpressed hind limb extends to the tip 
of the snout or beyond. Skin above smooth; a fold across the 
chest. Male with a subgular, external vocal sac. 

Color. Above, brown, usually with light-margined, transverse 
bands and a dark vertebral line ; apparently a canthal streak ; a 
transverse light glandular ridge line above the anus ; limbs gen- 
erally crossbarred; flanks and posterior part of the thighs uni- 
formly brown. Below, light brownish. 

Measurements. Snout-vent <$ 47, 9 49 ; head breadth S 13.5, 
9 14 ; head length $ 15.2, 9 17 ; femur <$ 24, 9 27 ; tibia $ 26, 

9 31. 

Range. The eastern Venezuelan Guayana. The species extends 
from British Guiana to southern Brasil. The race appears to 
stop in the vicinity of latitudes 5° and 10°. 

Remarks. As far back as 1900 Andersson pointed out that Hyla 
maxima Laurenti was a synonym of Rana boans Linne and that 
Hyla boans Daudin should receive another name, for which he 
proposed Hyla albopunctata Spix. Apparently overlooking this, 
Ahl (1939: 318) suggested that the name Hyla boans Daudin 
be changed to Hyla albopunctata, due to preoccupation by 
Calamita boans Schneider, 1799 (= Hyla tibiatrix Laurenti, 
1768). This is not the case, however, and in 1940, Mertens (p. 
195) reemphasized that boans is only applicable to Hyla maxima 
Laurenti and not to Hyla tibiatrix Laurenti. Confusion still per- 
sists in some museums where the two very different species Hyla 
albopunctata Spix and Hyla boans (Linne) are included under 
the same name. 



RIVERO : SALIENTIA OF VENEZUELA 107 

Besides the 5 Venezuelan specimens, I have examined 36 from 
British Guiana (A.M.N.H., U.M.M.Z.) and 4 from Paraguay and 
Brasil (M.C.Z.). The northern and southern animals show a 
decided difference in the coloration of the dorsum, flanks and 
limbs. In the Brasil-Paraguay group, both the flanks and the 
posterior part of the thighs are spotted with white (as described 
by Spix) and the dorsum is not crossbanded (except in one case, 
where the bands are somewhat diffuse and the design quite dif- 
ferent from the condition found in the northern animals). The 
5 Venezuelan frogs (one indistinctly) and most of those from 
British Guiana are banded above and the flanks and hinder part 
of the thighs of all of them are of a plain brown color. It is 
interesting to note that Daudin, most of whose specimens were 
apparently from Surinam, and Boulenger, who also had the 
greater number of specimens from the north, do not describe any 
white spots on the flanks and limbs. Dumeril and Bibron (1841) 
make no mention of the white spots and they had animals from 
''Surinam, Cayenne et du Bresil." Cochran (personal communi- 
cation) and Ruthven (1919) have also spoken of the uniform 
coloration of the posterior part of the thighs and flanks in speci- 
mens from British Guiana. 

The frog from Rio Bermejo named Hypsiboas raniceps by Cope 
(1862) was thought later to be identical (Cope, 1863) with Rein- 
hardt and Luetken's Hyla oxyrhina described as having white 
spots. Dr. Cochran, however, tells me (letter, 16.10.51) that these 
two species should be kept distinct. In a recent paper (1955 : 80) 
she puts H. oxyrhina in the synonymy of H. albopunctata and 
H. spegazzini in the synonymy of H. raniceps. Hyla multifas- 
ciata Giinther, 1858, and Hyla boons Bauman, 1912, refer to 
specimens from Para, none of which had the white spots. Parker 
(193"i) does not say anything of his British Guiana specimens 
but (1928) describes the white spots in animals from Brasil. 

Shreve (1935:211) pointed out the difference between Hyla 
boans Daudin and Hyla albopunctata Spix but the specimen of 
"boons" he had was Hyla raniceps (erroneously det. as boans 
in M.C.Z.) from Pernambuco. This apparently led him to con- 
elude that H. raniceps (spegazzini auct.) could be a southern 
representative of H. boans. 

As the present distribution and the morphological characters 
of the species only suggest subspecific differentiation between 
the northern and the southern form, the name Hyla albopunctata 
multifasciata is suggested for the animal occurring north of the 
Amazon. 



108 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



Although I have examined considerable material from the 
Coastal Range of Venezuela, no specimen from that region agrees 
with Peter's description of Hyla rostrata. Hyla albopunctata 
multifasciata agrees in many important details with this descrip- 
tion and in all probability, rostrata is a synonym of this species. 

Hyla loveridgei sp. n. 

Figure 8 

Type. Museum of Comparative Zoology No. 28565, a $ from 
Pico Culebra, Mt. Duida, 3000 ft., Territorio Amazonas. Coll. 
J. A. Rivero, 4 May 1950. 






Fig. 8. Hyla loveridgei sp. n. Type M.C.Z. 28565. 

Diagnosis. A medium sized Hyla with broad, adherent tongue ; 
tympanum y 2 the eye diameter; outer finger with three free 
phalanges; a "blunt pollex; disks of the fingers larger than the 
tympanum; disks of the toes much smaller than those of the 
fingers ; heel extending to between eye and nostril and marbled 
coloration above. 



RIVERO: SALIENTIA OF VENEZUELA 109 

Description. Snout rounded ; tongue very broad, adherent and 
nicked behind ; vomerine odontoids forming a /~ \ figure be- 
tween the large, oval choanae ; eye diameter very slightly greater 
than distance between eye and nostril ; interorbital space more 
or less equal to an upper eyelid ; canthus moderate ; loreal slightly 
oblique, concave ; tympanum V2 the eye diameter, its dorsal mar- 
gin covered by a supratympanic fold that runs to the shoulder ; 
a row of closely set tubercles along the forearm ; a blunt, ex- 
ternal rudiment of pollex ; first finger shorter than second ; sub- 
articular tubercles moderate; disks large, the largest larger than 
the tympanum ; outer three fingers with a rudiment of web ; a 
very distinct transverse fold at the heel ; a slight tarsal fold ; 
two metatarsal tubercles, the outer small, round and indistinct; 
toes taken in order from first to fifth exhibit the following 
phalanges free of web : 2, V/ 2 , V/2, 3, 1% ; disks of the toes much 
smaller than those of the fingers ; heel of the adpressed hind limb 
extends to between eye and nostril. Skin above, smooth. Male 
with a brown rugosity on the pollex and inner finger but no 
apparent external vocal sac. 

Color. Above, reddish brown marbled with black ; flanks with 
numerous small whitish spots ; limbs crossbarred. Below, immacu- 
late. 

Measurements. $ Snout-vent 42; head breadth 15; head 
length 14.5 ; femur 20 ; tibia 22. 

Habits. The only specimen was taken at about 3 P.M. as it 
sprang from beneath (or near?) a rock in a treeless, rocky area 
of Mt. Duida. 

Remarks. Like Hyla callipleura, this species has white spots 
on the flanks but these are small and less distinct than in the 
Bolivian animal which also has a completely different coloration 
above, less webbed fingers, and different vomerine teeth. 

Hyla raniceps (Cope) 

Eypsiboas raniceps Cope, 1862, Proe. Acad. Nat. Sei. Phila., 14: 353: Rio 
Vermejo, Paraguay. 

1 (A.M.N.H. 13055) Cucuhy R. 
Description. Snout subovoid ; tongue broad, cordiform, slightly 
free and nicked behind; vomerine odontoids forming a /~ ~\ 
figure between the moderate choanae ; eye diameter equal to 
distance between eye and nostril ; interorbital space more or less 
equal to an upper eyelid ; canthus well defined, curved ; loreal 
almost vertical, slightly concave ; tympanum oblique, % the eye 



110 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

diameter ; a dorsolateral fold from posterior corner of the eye to 
behind the shoulder; no external rudiment of pollex; a slight 
fold along the forearm; one elongated but flat and indistinct 
inner metacarpal tubercle; fingers with a short basal web (all, 2, 
2%, 2%) ; disks small, the largest about % the size of the tym- 
panum ; a slight tarsal fold ; a triangular dermal appendage at 
the heel; a small, oval, inner, and a minute, rounded, outer 
metatarsal tubercle ; toes fully webbed with the exception of the 
fourth, where two phalanges are left free ; heel of the adpressed 
hind limb extends to between eye and nostril. Skin above, very 
finely granular. 

Color. Above, brown, with distinctive darker vermiculations ; 
loreal region darker than the rest of the head; a dark supra- 
tympanic streak continues with the fold for about % of the 
length of the flank ; posterior part of the thighs yellowish brown 
with broad, dark brown crossbars that become diffuse and less 
distinct above and then darker again on the anteroventral por- 
tion ; rest of the hind limbs with diffuse spots that sometimes form 
crossbars; ventral surfaces yellowish brown. 

Measurements. $ Snout-vent 70 ; head breadth 23 ; head length 
21 ; femur 34.5 ; tibia 38. 

Habits. Collected on ground in open grass during the day. 

Range. Only known from this specimen in Venezuela. Para 
and S. Venezuela to Chaco Argentine 

Remarks. Comparison of this specimen with seven examples 
from Paraguay (M.C.Z. 24811-G, Dept. Villeta) reveals these 
last to be smaller (larger, 63 mm.), having a more pointed snout, 
occasionally a vertebral stripe and always a longer hind limb, 
the heel of the adpressed hind limb extending to the nostril or tip 
of the snout. Some males in this group have moderate, subgulo- 
pectoral vocal sacs. 

Cochran (1955:96) has discovered that Hyla spegazzini Bou- 
lenger is a synonym of Hyla raniceps (Cope). 

Hyla taurina (Steindachner) 

Ostcocephalus taurinus Steindachner, 1862, Archiv. per. la Zool., S: 11, pi. 

vi, figs. 1-3: Rio Negro, Brasil; Boulenger, 1882, Cat. Batr. Sal. Brit. 

Mns., ed. 2: 363. 
Hi/la taurina Boettger, 1896. Ber. Senekenb. Naturf. Ges.: LIT. 

1 (U.P.R. 113) Tapara, v.50. 

2 (U.P.R. 114-5) Mt. Marahuaca, v.50. 

1 (U.P.R. 116) La Culebra, 1000 ft., vi.50. 
1 (U.P.R. 117) Pto. Ayacucho, vi.50. 



RIVERO: SALIENTIA OF VENEZUELA 111 

Description. Head depressed ; skull rugose ; snout rounded in 
the female, less so in the male ; tongue adherent, subcircular or 
broadly oval, entire and slightly free behind ; vomerine odontoids 
ina/^ ^\figure between the oval, oblique ehoanae ; eye diam- 
eter about % the distance between the eye and nostril ; interorbital 
space much broader than an upper eyelid, with two bony ridges 
of varied prominency that extend back to the occipital region ; 
canthus strong; loreal concave, sloping in the female, almost 
straight in the male ; tympanum distinct, almost as large as the 
eye ; a distinct supratympanic fold ; no distinct rudiment of 
pollex ; subarticular tubercle of the outer finger double ; first 
finger shorter than second; taken in order from first to fourth, 
the fingers exhibit the following phalanges free of web : all, 1%, 
2% to 2%, 2y 2 to 2% ; larger disks almost as large as the tym- 
panum ; a small, round, outer and a prominent, inner metatarsal 
tubercle ; fourth toe leaving from 2 to 2y 2 free phalanges, others 
completely webbed ; disks of the toes smaller than those of the 
fingers ; heel of the adpressed hind limb extends to the nostril. 
Skin above, with small, flat tubercles in the female, distinctly 
tubercular in the male. Male with a pair of black vocal pouches 
behind the angle of the jaw and a brown rugosity in the inner side 
of the first digit. 

Color. Above, brown with irregular and generally obscure 
darker blotches ; flanks occasionally spotted with white ; limbs 
crossbarred, the hind part of the thighs uniformly brown. Lower 
surfaces grayish white, generally reticulated with brown. 

Measurements. Snout-vent $ 73, 9 86; head breadth £ 24, 
9 28; head length & 24, 9 26.5; femur J 36, 9 41; tibia 
<S 37.5, 9 44. 

Habits. All specimens were taken at night on the lower 
branches of trees or palm fronds, usually near water. The voice 
is a deep croak similar to that of Hyla boans. U.P.R. 117 was 
full of eggs. 

Additional Localities. Caracas (Boulenger, 1882) ; Esmeralda 
(A.M.N.H. 23174) ; Pescado, Caho (A.M.N.H. 27777) ; Yapacana 
(U.S.N.M. 83947) ; Upper Orinoco (Boettger, 1896). 

Range. The Venezuelan Guayana. British Guiana, Ecuador, 
Bolivia, N. Brasil and probably southeastern Colombia. 

Remarks. The male of this species is quite different from the 
female (snout less broad, cranial ridges less prominent, dorsum 
distinctly tubercular), and rather similar to some males of Hyla 



112 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

tibiairix, from which it can be immediately distinguished by the 
position of the vomerine teeth. 
See remarks under Hyla tibiatrix. 

Hyla paramica sp. n. 

Type. Museum of Zoology University of Michigan No. 59016, 
a $ from Escorial, Venezuela. 

Diagnosis. A small Hyla with an almost semicircular snout ; 
long, almost transverse vomerine teeth situated immediately be- 
hind the choanae ; adherent tongue ; small, indistinct tympanum ; 
outer finger with 2*4 free phalanges; heel extending to the eye 
and smooth dorsal surfaces. 

Description. Snout short, almost semicircular ; tongue broad, 
entire and adherent ; vomerine odontoids in two long, scarcely 
oblique, almost continuous groups, slightly behind the moderate 
choanae, their converging anterior extremities extending to the 
level of the inner margin of the latter ; eye diameter equal to 
distance between eye and nostril ; interorbital space about IV2 
times broader than an upper eyelid ; canthus rounded, much 
curved ; loreal slightly oblique, not concave ; tympanum small 
and indistinct, not quite y 2 the eye diameter; a blunt rudiment 
of pollex ; one oval, inner metacarpal tubercle ; subarticular tu- 
bercles small ; first finger shorter than second ; fingers taken in 
order from first to fourth exhibit the following phalanges free 
of web: all, 2, 2 1 /), ^Vi . ; disks small, the largest slightly smaller 
than the tympanum; no tarsal fold: inner metatarsal tubercle 
very indistinct, outer absent; toes exhibiting the following free 
phalanges: 2, 1V>, 1%, 3, IV2; heel of the adpressed hind limb 
extends to anterior corner of the eye. Skin above, smooth. 

Color. Above, yellowish white with scattered, small melano- 
phores ; anterior and posterior part of the thighs with a slight 
reddish tinge. Below, yellowish. 

Measurements. $ Snout-vent 31; head breadth 11.5; head 
length 10; femur 14.5; tibia 15.5. 

Remarks. The following specimens are designated as para- 
types : 

C.N.H.M. 3567, a 9 from Escorial. Snout -vent 32; head 
breadth 16; head length 11; femur 15; tibia 16.5. Tympanum 
slightly less distinct than in the type and pollex considerably 
more prominent but always blunt and unhooked. 

C.N.H.M. 3568, a 9 from Escorial. Snout-vent 34; head 
breadth 12; head length 10.5; femur 17; tibia 17.5. Pollex 
more prominent than in the type ; a definite orange tinge on the 



RIVERO: SALIENTIA OF VENEZUELA 113 

sides of the head, throat, chest and underside of the hind limbs. 

A.M.N.H. 10636, 6 , La Culata, 3000 m. Snout-vent 35 ; head 
breadth 11.5; head length 11 ; femur 15.5; tibia 16.5. A rounded 
pollex and a slight external vocal sac; throat, loins and thighs 
below, orange drab. 

A.M.N.H. 10639, 9 , La. Culata, 3000 m. Snout-vent 28 ; head 
breadth 11; head length 10; femur 14; tibia 14.5. Orange drab 
on the dorsum, back of the head and on the belly. 

This species is somewhat similar to //. jahni from which it can 
be distinguished by the rounded snout, less defined canthus, lack 
of supratympanic fold and absence of canthal, palpebral and 
supratympanic lines. 

Hyla jahni sp. n. 

Type. Museum of Zoology University of Michigan No. 46465, 
a $ from Escorial, Venezuela. 

Diagnosis. A small Hyla with subovoid snout; adherent 
tongue ; slightly oblique vomerine teeth behind the choanae ; hid- 
den or indistinct tympanum ; flat fingers and toes ; outer finger 
with 2 1 /? free phalanges ; heel extending to the anterior corner 
of the eye; punctuated dorsal surfaces and light line along the 
canthus and supratympanic fold. 

Description. Snout subovoid; tongue circular, adherent and 
very slightly nicked behind; vomerine odontoids in two slightly 
oblique groups behind the choanae, their converging anterior 
extremities extending to the horizontal of the inner margin of 
the latter ; eye diameter more or less equal to distance between 
eye and nostril ; interorbital space about twice as broad as an 
upper eyelid ; canthus distinct, curved ; loreal almost vertical, 
not concave; tympanum hidden; a very distinctive fold from 
posterior corner of the eye to slightly behind the axilla; no dis- 
tinct external rudiment of pollex ; a prominent oval, inner meta- 
carpal tubercle ; first finger shorter than second ; fingers and toes 
flattened ; taken in order from first to fourth, the fingers exhibit 
the following phalanges free of web : all, 2, 2y 2 , 2y 2 ; subarticular 
tubercles small ; one flat and indistinct inner metatarsal tubercle 
but no outer ; toes exhibiting the following free phalanges : 1%, 
1%» 1%> 2%, 1 ; heel of the adpressed hind limb extends to an- 
terior corner of the eye. Skin above, smooth. 

Color. Above, brown, closely punctuated with darker brown ; 
canthus, margin of upper eyelid, and probably the supratym- 
panic fold with a light line ; anterior and posterior part of the 
thighs plain brownish. 



114 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Measurements. S Snout-vent 34.5; head breadth 11.5; head 
length 11; femur 15.5; tibia 16.5. 

Remarks. The following specimens have been designated as 
paratypes : 

U.M.M.Z. 59016, a 9 from Escorial. Snout -vent 29; head 
breadth 11; head length 9.5; femur 15; tibia 16. Tongue entire; 
vomerine teeth juxtaposed ; tympanum present but indistinct, 
not quite y 2 the eye diameter ; disks larger than the tympanum ; 
toes exhibiting the following phalanges free of web : 1%, 1^4, l 1 /^ 
2%, IV2 ; nee l extending to between eye and nostril ; color uniform 
yellowish above and below. 

U.M.M.Z. 46465, a 9 from Escorial. Snout-vent 25; head 
breadth 8.5 ; head length 8 ; femur 12.5 ; tibia 12.5. Tympanum 
hidden or almost so; color above whitish with dark points that 
are scattered and less distinct than in the type. Canthal line 
distinct. 

M.C.Z. 2523, a $ from Merida. Snout-vent 24.5; head 
breadth 11.5; head length 7.5; femur 12; tibia 12. Tympanum 
indistinct ; color above and below whitish ; white supratympanic 
line present. 

A.M.N.H. 10637, a 9 from La Culata, 3,000 m. Snout-vent 
26 ; head breadth 8 ; head length 7.5 ; femur 12.5 ; tibia 13. Tym- 
panum hidden ; fingers appear to be shorter relatively than in 
the type; punctuation above, white instead of dark; canthal, 
palpebral and supraorbital lines distinct ; ground color above 
and below, yellowish white. 

A.M.N.H. 10638, a $ from La Culata, 3,000 m. Snout-vent 
31.5; head breadth 11; head length 10.5; femur 14.5; tibia 16. 
Tympanum very indistinct, almost hidden; rudiment of pollex 
more distinct than in the type and other specimens; supratym- 
panic fold extending for some distance along the flank ; canthal, 
palpebral and supratympanic lines distinct ; ground color above 
yellowish, with white punctuation ; hind limbs below, with an 
orange drab. 

A.M.N.H. 10640, a S from La Culata, 3,000 m. Snout-vent 
31.5; head breadth 11: head length 10.5; femur 15; tibia 16. 
Tympanum almost hidden; ground color brownish; canthal, pal- 
pebral and supratympanic lines golden ; color above and below 
brownish with dark punctuation above. 

The dark punctuation of the dorsal surface may become light 
or even disappear on preservation. When present, this species 
looks somewhat like Hyla albomarginata but can be distinguished 
from that form by its smaller size and many other characters. 



BIVERO : SALIENTIA OF VENEZUELA 115 

Hyla jahni appears to be restricted to the high elevations of the 
Andes where Hyla albomarginata does not exist. 

Specimens of Hyla jahni have been also confused with Hyla 
punctata but although this form, or a race of it, may be found in 
Venezuela, it is unlikely to be in the Andes as high as 2,000 m. 
Hyla punctata has large, well defined tympana that are about 
% the eye diameter, the eyes are partially directed forwards and 
the small spots above are scattered and always white in preserva- 
tion. 

Hyla paramica differs from Hyla jahni in its rounded snout, 
less defined canthus and in lacking supratympanic fold, canthal, 
palpebral and supratympanic lines and closely set dark dots 
above. 

The species is named after Dr. Alfredo Jahn, great student of 
the Venezuelan Paramos. 

Hyla platydactyla Boulenger 

Hyla platydactyla Boulenger, 1905, Ann. Mag. Nat. Hist., (7) 16: 183: 
Merida, Andes of Venezuela ; Lutz, A., Mem. Inst. Osw. Cruz, 20 : 39, 44. 

No material examined. 

Original Description. Head moderate, much depressed, a little 
broader than long; snout rounded, hardly as long as the orbit; 
nostrils near the tip of the snout ; tongue subcircular, entire, 
slightly free behind ; vomerine odontoids just behind the level of 
the moderately large choanae, in two slightly oblique series con- 
verging forwards ; interorbital region a little broader than an 
upper eyelid ; canthus rostralis distinct ; loreal region very 
oblique, concave ; tympanum rather indistinct, % the eye diame- 
ter ; fingers much flattened, with a short basal web which extends 
as a fringe to the disks ; toes short, much flattened, with a deeply 
notched web extending to the disks ; latter about % the eye diame- 
ter; subarticular tubercles feeble; no tarsal fold; heel of the 
adpressed hind limb extends to the eye. Skin above, smooth. 
Below, belly and thighs granular. Male with a feebly developed, 
subgular vocal sac. 

Color. Above, purplish, closely dotted with darker ; upper sur- 
faces of the arms and thighs white, with a median stripe of 
closely set, dark purple dots. Below, white. 

Measurements. Snout-vent 32. 

Remarks. This species seems to be very distinctively colored. 
It appears to be of a uniform light color, densely mottled with 
darker ; between mottlings, the light ground color can be seen 



116 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

especially on the flanks and limbs. On the hand and foot, the 
dark color is limited to the outer digits, giving the impression 
of the situation in Phyllomedusa. 

Hyla platydactyla is of approximately the same size as Hyla 
jahni and Hyla paramica and occurs in the same general area as 
these two species. It differs from Hyla jahni in the shorter snout 
and less defined canthus, absence of a light canthal and supra- 
tympanic line, narrower interorbital space, more vertical loreal 
region and different coloration. From Hyla paramica it can be 
differentiated by its longer snout, more oblique loreal region, 
absence of a pollex rudiment (not described or shown in photo- 
graph) and different coloration. It is also said to be allied to 
Hyla vilsoniana Cope (Boulenger, 1905). 

I owe to Mr. J. C. Battersby the comparison of a photograph 
of Hyla jahni with the type of Hyla platydactyla and to Miss 
A. G. C. Grandison the photographs of the type specimen shown 
here. All remarks are based on this photograph and on the de- 
scription of the species. 

Hyla benitezi sp. n. 
Figure 9 

Type. Museum of Comparative Zoology No. 28564, a $ from 
Cafio Wanadi, Mt. Marahuaca, Territorio Amazonas. Coll. J. A. 
Rivero, 20 May 1950. 

Diagnosis. A medium sized Hyla with adherent tongue, vo- 
merine teeth forming a / \ figure between the choanae ; tym- 
panum small, indistinct; an external rudiment of pollex; outer 
finger with 2% free phalanges ; largest disks larger than the tym- 
panum and orange ventral coloration in life. 

Description. Snout subovoid; tongue circular, adherent and 
nicked behind; vomerine odontoids forming a / \ figure be- 
tween the moderate, oval, choanae, their anterior extremities 
extending to about the middle of the oblique inner margin of 
the latter; eye diameter more or less equal to distance between 
eye and nostril; interorbital space slightly broader than an 
upper eyelid; canthus indistinct; loreal sloping, slightly non- 
cave ; tympanum very small and indistinct, % the eye diame- 
ter; an external rudiment of pollex; first finger shorter than 
second ; outer three fingers with a short basal web ; disks moderate 
but larger than the tympanum; subarticular tubercles small, 



BIVERO : SALIENTIA OF VENEZUELA 



117 



rounded; an elongated, flat and indistinct inner metatarsal tu- 
bercle but no outer ; toes fully webbed with the exception of the 
fourth where about two phalanges are left free ; disks of the toes 
smaller than those of the fingers; heel of the adpressed hind 
limb extends to the middle of the eye. Skin above, smooth ; below, 
finely granular. Male with a slight subgular vocal sac. 

Color. Above and on the limbs brown, with darker brown 
transverse bars of varying width. Ventral surfaces including 
the limbs bright orange (in life) or white (alcohol after forma- 
lin). The inner fingers and toes and the webs were also orange 
above. 






Fig. 9. Eyla benitezi sp. n. Type M.C.Z. 28564. 



Measurements. $ Snout-vent 37; head breadth 13.5; head 
length 12; femur 16; tibia 18.5. 

Habits. Both type and paratype were collected on the leaves 
of a dwarf palm that thrives on the stones of mountain stream 
rapids. The voice is heard sporadically at night but due to the 



118 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

uproar of the water, their localization is very difficult and their 
collection in the inaccessible places from which they call gen- 
erally impossible. 

Remarks. This form is quite different from anything known 
in Venezuela or from material of northern South America avail- 
able to the author for comparison. 

Morphologically, the only paratype is almost identical with 
the type but differs from it in coloration. 

U.P.R. 145, a $ from Cano Wanadi, Mt. Marahuaca. Coll. 
J. A. Rivero, May, 1950. Snout-vent 36 ; head breadth 13.5 ; head 
length 12.5 ; femur 16 ; tibia 19. Color above, grayish brown with 
very obscure, slightly darker spots on the body and reticulations 
on the head. The tympanum is probably less distinct than in the 
type and the openings of the Eustachian tubes smaller. 

Hyla marahuaquensis sp. n. 
Figure 10 

Type. Museum of Comparative Zoology No. 28566, a $ (?) 
from Cafio Caju, Mt. Marahuaca, Territorio Amazonas. Coll. 
J. A. Rivero, May 1950. 

Diagnosis. A small Hyla with physiognomy of Eleutherodac- 
tylus; transverse vomerine teeth between the ehoanae ; fingers 
webbed; first finger longer than second; heel extending to the 
nostril; very peculiar dorsal coloration. 

Description. Snout rounded ; tongue oval, slightly free and 
nicked behind ; vomerine odontoids in two juxtaposed, transverse 
groups between the moderate, oval ehoanae ; eye diameter equal 
to distance between eye and nostril ; interorbital space slightly 
convex, as broad as an upper eyelid ; eanthus moderate ; loreal 
moderately oblique, concave ; tympanum large, i/> the eye diame- 
ter ; an indistinct supratympanic fold from lower eyelid to 
shoulder; fingers long and slender, free, the first longer than the 
second but shorter than fourth; two small metacarpal tubercles; 
subarticular tubercles well defined, rounded; largest disks about 
V2 the size of the tympanum ; a small inner metatarsal tubercle 
but no outer; toes taken in order from first to fifth exhibit the 
following phalanges free of web : 2, 2, 2, 3%, 2*4 ; hind limbs 
long and slender ; heel of the adpressed hind limb extends to the 
nostril. Skin above smooth ; granular on the loreal and temporal 
regions ; rugose on the upper eyelids. Ventral surfaces of the 
body and thighs finely granular. 



RIVERO : SALIENTIA OF VENEZUELA 



119 



Color. Above, brown, this color bounded by a pair of dorso- 
lateral lines that cross the black upper eyelids and meet in a 
posteriorly directed notch in the interorbital space ; snout lighter 
than the dorsal color enclosed by the dorsolateral lines ; a small 
black spot behind the nostril ; upper lip with light vertical bars ; 
three dark spots on the dorsum in front of the sacrum, one in 
the middle of the back and one on each side just below the dorso- 
lateral line; behind these, other more diffuse spots occur in 
succession; limbs yellowish brown streaked with darker brown; 
flanks of the same color as the limbs but uniform below the 
already described spots. Ventral surfaces dirty white. 






Fig. 10. Hyla marahuaquensis sp. n. Type M.C.Z. 28566. 



Measurements. $ ? Snout-vent 27; head breadth 10; head 
length 10.5; femur 14; tibia 15.9. 

Remarks. This specimen was thought to be an Eleutherodac- 
tylus until its true affinities were revealed by the position of its 
vomerine teeth, the expanded sacral diapophysis, the presence 
of claw-shaped terminal phalanges and intercalary disks. It is, 



120 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

however, a very peculiar frog whose allocation to the genus Hyla 
may be considered provisional. 

It differs from all Venezuelan Hyla in having the first finger 
longer than the second and in this respect it agrees more with 
Gastrotheca, a genus in which it will perhaps eventually be 
placed. 

Hyla rubra Laurenti 

Hyla rubra Laurenti, 1768, Syn. Kept.: 35: America; Boettger, 1892, Kat. 
Batr. Samm. Mus. Senckenb. : 45; 1896, Ber. Seuckenb. Naturf. Ges. : 
LIV; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 39, 43. 

1 (U.P.R. 152) Sn. Fdo. de Atabapo, vi.50. 

Description. Snout long, subovoid ; tongue oval, adherent and 
indistinctly nicked behind ; vomerine odontoicls in two short, 
transverse groups between the moderate, oval, choanae ; eye 
diameter shorter than distance between eye and nostril ; inter- 
orbital space as broad as an upper eyelid ; canthus rounded, in- 
distinct ; loreal moderately oblique, slightly concave ; tympanum 
distinct, about % the eye diameter ; a flat, indistinct supratym- 
panic fold ; no external rudiment of pollex ; subarticular tubercles 
small ; fingers free, the first shorter than second ; larger disks 
about i/j the size of the tympanum ; two small metatarsal tu- 
bercles ; toes taken in order from first to fifth exhibit the follow- 
ing phalanges free of web : 2, 1, iy 2 , 2, 1 ; heel of the adpressed 
bind limb extends to the eye. Skin above, with small granules 
on the head and scapular region ; flanks granular ; throat granu- 
lar. Male with a slight, subgular vocal sac. 

Color. Above, very dark gray, almost black ; limbs lighter than 
the body color ; flanks, anterior part of the thighs, and ventro- 
posterior part of the tibiae obscurely mottled with light ; a broad 
but short longitudinal white line along the posterior part of the 
thigh. Ventral surfaces dirty white. 

Measurements. $ Snout-vent 41; head breadth 12.5; bead 
length 14; femur 16; tibia 20. 

Additional Localities. Caracas (Boettger, 1892) ; Upper Ori- 
noco (Boettger, 1896). 

Range. Tbe Venezuelan Guayana and the Coastal Range. 
Central America through Colombia to Bolivia on the west and 
British Guiana to Bahia on the east. Boulenger (1882) records 
it from Uruguay, and Cei (1956) cites Mirando Ribeiro as in- 
cluding Argentina within the range of the species. Cocbran 
(1955) does not include it in the fauna of S. Brasil. 



RIVERO: SALIENTIA OF VENEZUELA 121 

Remarks. The San Fernando specimen is large for the species 
and its very dark color is different from any specimen of Hyla 
rubra I have seen. It agrees with //. rubra, however, in all struc- 
tural characters and the markings that are usually shown on the 
limbs of II. rubra are present, though obscure, in the San Fer- 
nando example. The white line on the thighs appears to be a 
longitudinal expansion of a spot and perhaps does not have any 
importance. 

An M.C.Z. specimen from Central America, and another from 
Para, Brasil, exhibit very peculiar spotting all over the dorsum. 
A group from Barana Dist., British Guiana, shows rounded snouts 
(in cross-section) which are much elevated between the eyes and 
in the intercanthal region. Probably H. rubra as known today 
represents a composite of several related forms. 

In several museums, specimens of Hyla Columbiana Bttgr. are 
confused with Hyla rubra. 

Hyla rubra x-signata Midler, 1927, H. rubra, hubneri Melin, 
//. rubra inconspicua Melin and H. rubra duartei Lutz, B., have 
been described from different localities. I have preferred to use 
the binomial until the whole species is studied and its components 
seuTeo-ated. 



J o J 



Hyla boulengeri (Cope) 

Scytopis boulengeri Cope, 1887, Bull. U.S. Nat. Mus., 32: 12: Nicaragua. 

?Hyla acuminata Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 39, 43. 

Hyla boulengeri Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 40. 

.'Hyla palpebrogranulata Lutz, A. (not Andersson), 1927, Mem. Inst. Osw. 

Cruz, 20: 39, 44, pi. XI, fig. 20, 20a. 
Hyla boulengeri Shreve, 1947, Bull. Mus. Comp. Zool., 99: 537. 

1 (M.C.Z. 12888) Bejuma, nr. Rio Montero, ii.20. 

1 (M.C.Z. 25984) Pauji, iv.45. 

1 (U.M.M.Z. 55828) Aroa. 

1 (U.M.M.Z. 55829) Rio Bejuma. 
Description. Snout subacuminate, projecting beyond the 
mouth; tongue cordiform, free and nicked behind; vomerine 
odontoids in two short, transverse groups between the large, oval 
choanae ; eye diameter shorter than distance between eye and 
nostril; interorbital space about l 1 /? times broader than an upper 
eyelid ; canthus indistinct ; loreal oblique and concave ; tympanum 
% the eye diameter ; no rudiment of pollex ; subarticular tu- 
bercles rather small ; fingers free, their disks transversely oval 
and about V. the size of the tympanum; no tarsal fold; meta- 
tarsal tubercles small ; toes taken in order from first to fifth 



122 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

exhibit the following phalanges free of web : almost all, 1, 1*4, 
2^2 to 3, 1 ; head of the adpressed hind limb extends to the tip 
of the snout. Skin above, mostly smooth. Male with a subgular 
vocal sac. 

Color. Above, brown with a triangular spot between the eyes 
that is occasionally light margined; a few, small, dark spots on 
the anterolateral part of the dorsum ; thighs with a longitudinal 
brown band above and black transverse bars in front and behind 
it ; on reaching the longitudinal band, the transverse bars assume 
a much lighter color ; space between the bars pinkish red (alive) ; 
hidden part of the tarsus and upper part of the foot rose and 
black spotted ; rest of the limbs uniform or with wide transverse 
spots. Below, grayish or dirty white; vocal sac generally in- 
fuscate. 

Measurements. Snout-vent $ 34, 9 58.5 ; head breadth $ 11, 
9 16; head length $ 12.5, 9 16; femur $ 18, 9 24; tibia 
S 21.5, 9 30. 

Habits. The Puerto Ayacucho specimens were collected while 
calling from tall grass in an inundated area. 

Additional Localities. Maracay (Lutz, 1927) ; Orinoco R., be- 
low mouth of Caroni (M.C.Z. 19919) ; Orinoco R., nr. mouth 
Cano Ilorobopo (?) (M.C.Z. 19920); Puerto Ayacucho (U.P.R. 
132-8). 

Range. The Venezuelan Guayana and the Coastal Range. 
Central America, Colombia and British Guiana. 

Remarks. It is possible that the specimens from northern Vene- 
zuela are different from those of Central America. A topotypical 
specimen examined (M.C.Z. 4896) has distinct tubercles on the 
head and anterior part of the body, the heel extends to between 
eye and nostril and the bars on the thighs cross this limb segment 
above without losing their color or being interrupted by the 
median longitudinal band. There are also black spots (and prob- 
ably reddish ones) on the groins and posterior part of the flanks. 
The material from Puerto Ayacucho also looks different from 
that of northern Venezuela and Nicaragua but possibly this is 
because all of the Puerto Ayacucho specimens are males while 
the others arc mostly females. They are very rugose above ; the 
heel reaches between eye and nostril ; the thigh bars are very 
well denned and not confluent and irregular as in the individuals 
from the north ; the head looks much narrower ; the toes are 
slightly less webbed and the color above is dark gray, without 
dorsolateral spots. 



BIVERO : SALIENTIA OF VENEZUELA 123 

I have placed Hyla palpebrogranulata Lutz in the synonymy 
of this species pending further investigation. According to An- 
dersson, who is the author of Hyla palpebrogranulata, this species 
' ' agrees most nearly with Hyla phrynoderma Boulenger and with 
Hyla verrucigera Werner and the related species of this one, H. 
buckleyi Gthr. and H. leprcurii Blgr." As the last two forms 
are different and unrelated to H. phrynoderma, the comparison 
makes one feel uncertain as to the appearance of this frog. Mr. 
Shreve has called to 1113' attention that H. phrynoderma Blgr., 
1889, is probably a synonym of H. acuminata Cope, 1862 ; and 
H. acuminata Blgr., 1882, a synonym of II. boulengeri. 

It also appears that Hyla acuminata Lutz, which was perhaps 
identified by using Boulenger 's catalogue, is also a synonym of 
Hyla boulengeri. However, I have placed it in the synonymy with 
some misgivings, for in referring to that species, Lutz speaks of 
two vocal sacs in the male, a peculiarity that is only found in 
Hyla tibiatrix among the Hyla reported from the Coastal Range. 
I doubt, however, that Lutz could have confused H. tibiatrix with 
H. acuminata. 

Hyla palpebrogranulata Andersson also seems to be a synonym 
of H. acuminata Cope, but the figure of this species shown by 
Lutz apparently belongs to H. boulengeri. The animal is repre- 
sented Avith a pointed snout tip such as is found in Miranda 
Ribeiro's Garbeana garbei, a form which is probably conspecific 
with H. boulengeri. 

Lutz also mentions Hyla boulengeri in his report, but this is 
based on material that was not examined by him. The specimens 
he collected were apparently given only two names : acuminata 
and palpebrogranulata. 

Hyla baumgardneri sp. n. 

Type. Museum of Comparative Zoology No. 28563, a $ from 
Casa de Julian, Territorio Amazonas. Coll. J. A. Rivero, 26 
May, 1950. 

Diagnosis. A small Hyla of the rubra group with flat head ; 
almost transverse vomerine teeth between the choanae ; adherent 
tongue ; tA'mpanum y 2 the eye diameter ; fingers free ; no rudi- 
ment of pollex ; short hind limbs and chlorine green color in 
life. 

Description. Snout flat above, subovoid ; tongue oval, nicked 
and very slightly free behind ; vomerine odontoids in two short, 
transverse groups between the small choanae ; eye diameter equal 



124 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

to distance between eye and nostril; interorbital space a little 
broader than an upper eyelid ; canthus rounded, indistinct ; loreal 
scarcely oblique, flat ; tympanum small but distinct, about y 2 the 
eye diameter ; a slight supratympanic fold ; metacarpal tubercles 
indistinct; no external rudiment of pollex; fingers free, their 
disks almost as large as the tympanum ; a small, oval inner and 
an indistinct outer metatarsal tubercle ; toes taken in order from 
first to fifth exhibit the following phalanges free of web : almost 
free, 1%, IV2, 2%, 1; heel of the adpressed hind limb extends 
to the eye. Skin above, smooth except on the head, snout, and 
upper eyelids where it is sparsely granular; a fold across the 
chest. A large subgular vocal sac. 

Color. Above, chlorine green (in life) or yellowish gray (alco- 
hol after formalin) ; canthus and supratympanic fold slightly 
darker than the rest of the head but with no well defined streak. 
Below, greenish yellow (in life) or grayish white. 

Measurements. $ Snout-vent 29 ; head breadth 9 ; head length 
9.5; femur 11.5; tibia 13.5. 

Habits. The type and U.P.R. 184 were collected at night while 
calling from shrubs at about 5 ft. from the ground. There was a 
stream running over a granite bed (see Leptodactylus rugosus) 
at some twenty feet away and as it was raining considerably at 
that moment, most of the ground under the shrubs was inun- 
dated. Most of the La Culebra specimens came from tall grass 
(savanna) in an inundated area but one was taken in a thatched 
roof at about 1 :00 A.M. The noise made by the frog was so an- 
noying that those trying to sleep beneath the thatch united to 
catch the intruder. The voice is a cicada-like "crrrr" that may 
continue without interruption for several minutes. 

Two pairs of specimens were in amplexus on grass blades about 
two feet from the water. 

Remarks. Except for a slight variation in the amount of web 
and other characters mentioned below, the paratypes agree in all 
essentials with the type. 

Anion:? the paratypes there are two females (U.P.R. 149, 179) 
both collected at La Culebra, 1,000 ft., and having a snout-vent 
length of 32 and 29.5 mm., respectively. The other paratypes 
were taken at Casa de Julian (U.P.R. 184), La Culebra (U.P.R. 
150-1, 176-8, 181-2) and Puerto Ayacucho (U.P.R. 180). Except 
for specimen No. 180 which is smaller than usual (22.5 mm.) and 
has a more pointed snout, there is no great variation among the 
members of the group. A triangular interorbital spot may be 
present or absent. 



RIVERO: SALIENTIA OF VENEZUELA 125 

This species is very similar to the Panamanian 77. altae Dunn 
(= 77. staufferi, fide Taylor, 1952) and both are so similar to 77". 
rubra that they might be mistaken for juveniles. 77. baumgard- 
neri is, however, distinguished from the type of Hyla altae by 
its flatter, granular snout which does not project beyond the 
mouth, probably larger size (type of altae, 23 mm.), different 
coloration, and probably different voice. As in 77. rubra, II. altae 
has two curved longitudinal lines on the dorsum from behind the 
eyelids to groin, and the basic color is light brown. A cross- 
section of the snout of 77. altae would be semicircular (as in 
British Guianan 77. rubra) and the voice, as described by Dunn, 
is a harsh "whark." Dr. Dunn explains, however (letter, 
14.4.51), that the Panamanian frogs were calling with so many 
others, that there is a distinct possibility the cries may have 
been confused. 

From Hyla rubra, Hyla baumgardneri differs in its smaller 
size, shorter snout, smaller tympanum, darkened eyelids and 
different coloration. 

Hyla luteocellata Roux 

Hyla luteo-ocellata Eoux, 1927, Verhandl. Naturf. Ges. Basel., SS: 260: El 
Mene, Prov. Falcon, Venezuela. 

No material examined. 

Original Description. Head somewhat broader than long ; snout 
short, scarcely longer than the orbital diameter; nostrils near the 
tip of the snout ; tongue oval, broader than long, entire and 
slightly free behind ; vomerine odontoids in two small groups 
between the choanae ; eyes large, protuberant ; interorbital space 
twice as broad as an upper eyelid ; canthus moderate ; loreal high, 
slightly oblique; tympanum hidden; second finger shorter than 
fourth; digits webbed at the base only; disks large; hind limbs 
long ; an outer, oval, metatarsal tubercle ,- subarticular tubercles 
rounded and rather small; toes % webbed; last two phalanges 
of the fourth toe free ; disks of the toes equal to those of the 
fingers. Skin above, smooth. Below, throat and breast smooth; 
a fold across the chest; belly reticulate and widely granular. 

Color. Above, grayish brown merging into light gray on the 
sides and sprinkled on the trunk and head with small, black 
spots ; a more or less distinct brown band between the eyes ; a 
narrow light line, commencing at the tip of the snout, follows 
the canthus rostralis and continues behind the eye, bordering 
the light dorsolateral region below ; loreal region with a dark 



126 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

brown band that extends to the side of the body passing over the 
forelimb ; a white spot on the hind part of the flanks ; a white 
(yellow in the living animal?) spot below the eye is surrounded 
with brown and extends to the lower jaw; inner aspect of the 
thigh occupied by a large, elongated-oval, white-margined, black 
spot ; upper surfaces of the thighs light brown with more or less 
rounded white or yellowish spots ; tibia grayish above, showing a 
light elongated spot on its lower surface near the articulation 
with the thigh ; tarsum speckled with white and gray. Belly 
whitish ; throat with some brown spots. 

Measurements. $ Snout-vent 31 ; hind limb 32. 

Remarks. The species is only known from the two male indi- 
viduals described by Roux. It is well characterized, however, and 
not likely to be confused with any other Venezuelan HyJa. 

Hyla marmorata marmorata (Laurenti) 

Bufo mannoratus Laurenti, 1768, Syn. Sept.: 29: Surinam. 

4 (U.P.R. 126-9) Sn. Fdo. de Atabapo, vi.50. 

Description. Snout short, almost semicircular, equal or slightly 
longer than the eye diameter ; tongue subcircular, a little free 
and nicked behind ; vomerine odontoids in two short, transverse 
and juxtaposed groups between the small, round choanae ; inter- 
orbital space more or less equal to an upper eyelid ; canthus 
rounded ; loreal little oblique and convex ; tympanum distinct, 
V2 to % the eye diameter ; a loose fold of skin from elbow to 
flanks and an external fringe from elbow to disk of last finger; 
no rudiment of pollex ; subartieular tubercles flat ; fingers de- 
pressed, exhibiting the following phalanges free of web : l!/->> 1, 
2, 1 ; the web extends to the disks as distinct lateral fringes ; 
disks almost as large as the tympanum ; a festooned, lateral fringe 
from knee to disk of outer toe ; two metatarsal tubercles, the 
outer, small and inconspicuous, the inner oval and prominent; 
toes fully webbed with the exception of the fourth where IV2 
to 2 phalanges are left free ; disks of the toes smaller than those 
of the fingers ; heel of the adpressed hind limb extends to the 
eye. Skin above, mostly smooth. Male with an external sub- 
gular vocal sac. 

Color. Above, grayish brown, marbled with darker tones of 
brown ; a light, hour-glass marking between the eyes, broadened 
anteriorly at the snout ; anterior and posterior parts of the 
thighs orange (whitish in alcohol) with a few dark, well defined 
spots; a longitudinal streak of different shades of brown along 



RIVERO : SALIENTIA OF VENEZUELA 127 

the upper part of the thighs ; tibial segment with dark, transverse 
spots on a lighter brown background. Below: throat, belly and 
ventral surface of thighs orange, with numerous small, black 
spots; lower lip dotted with white; tibiae, tarsi and toes black; 
web of fingers and toes orange and black. 

Measurements. $ Snout-vent 35; head breadth 11; head 
length 10; femur 15; tibia 11. 

Habits. The four males were collected, while calling, from the 
leaves of a bush near a rain pool. Surrounding area, savanna. 

Range. The Venezuelan Guayana. The species extends from 
the Guianas to Brasil (Angra dos Reis), Peru and Bolivia. Hyla 
melanargyrea and Hyla senicula, now considered races of mar- 
morata, were described from Matto Grosso (headwaters of 
Xingu) and Rio Napo or Upper Maranon, respectively. 

Remarks. The color recorded was based on observations made 
on a living frog at night with the aid of a flashlight. It is pos- 
sible that the orange may be actually red. 

Hyla tibiatrix tibiatrix Laurenti 

Hyla tibiatrix Laurenti, 1768, Syn. Kept.: 34 (type loe. not cited). 

Hyla venulosa Boettger, 1893, Ber. Senekenb. Naturf. Ges.: 40; Stejneger, 

1902, Proc. U.S. Nat. Mus., 24: 180; Lutz, A., 1927, Mem. Inst. Osw. 

Cruz, 20: 39, 44, pi. xiv, figs. 33, 33a, b, c, 34; Eohl, E., 1948, Fauna 

Descr. cle Ven., ed. 2 : 401, fig. 181. 
Hyla nigromaculata Boettger, 1895-6, Ber. Senekenb. Naturf. Ges.: LIV. 
Phrynohyas zonata Duellman, 1956, Misc. Publ. Mus. Zool. Univ. Micli., 

No. 96: 35, pi. v, figs. 1, 2; pi. vi, figs. 1, 2. 

2 (U.S.N.M. 22545, 27797) La Guaira, vii.02. 
1 (U.S.N.M. 128792) Los Canales, v.39. 
1 (U.S.N.M. 80612) nr. Carapa, x.29. 

1 (U.S.N.M. 36377) Caicara, v.05. 

2 (U.S.N.M. 117098-9) Caripito, 42. 

Description. Snout short, rounded; tongue cordiform, slightly 
nicked and free behind; vomerine odontoids in two transverse 
groups between the moderate choanae ; eye diameter shorter than 
distance between eye and nostril ; interorbital space more or less 
equal to an upper eyelid ; canthus rounded ; loreal moderately 
sloping, concave ; tympanum % to % the eye diameter, its dorsal 
margin covered by a glandular fold that runs to the shoulder; 
a number of rounded glandules behind the angle of the mouth ; 
no external rudiment of pollex ; subarticular tubercles not largo 
but prominent; first finger shorter than second; taken in order 



128 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

from first to fourth the fingers exhibit the following phalanges 
free of web: 2, V/ 2 to 1%, 2y 2 to 3, 2 ; larger disks about the 
same size as the tympanum; a slight tarsal fold; toes exhibiting 
the following free phalanges: 1% to 1%, 1 to V/ 4 , 1, 2 to 2y 2 , 
2 to 2V4 ; heel of the adpressed hind limb extends to the eye. 
Skin above, smooth, or more usually with scattered tubercles and 
warts; flanks glandular. Male with two vocal pouches at the 
angle of the mouth and a brown rugosity on the inner side of 
the first digit. 

Color. Above, light brown, marbled, spotted or with undulating 
broad bands that form variable patterns; hind limbs crossbarred, 
mottled or dotted with dark brown ; the crossbars, when present, 
are usually broad, with concave ends that enclose an almost cir- 
cular light spot. 

Measurements. Snout-vent 6 60.5, 9 76.5; head breadth $ 
20.5, 9 25.5; head length $ 18.5, 9 21.5; femur $ 28.5, 9 34; 
tibia $ 29.5, 9 37. 

Habits. Robinson (in Stejneger, 1902) describes the voice of 
this species as "A monotonous note like the bleating of a goat and 
fully as loud." 

Additional Localities. Maracay (Lutz, 1927) ; La Guaira (Stej- 
neger, 1902 [= U.S.N.M. 22545, 27797]); Maturin (Boettge'r, 
1893). 

Range. The Coastal Range, the eastern part of the Llanos 
(Caripito), the Delta and the Venezuelan Guayana. Central 
America to Bolivia (Procter, 1921) and from Trinidad and the 
Guianas to \ T . Brasil. Cochran (1955: 55) mentions its occur- 
rence in Argentina; Peracca (1904) has reported the species 
from Paraguay. Duellman (1956) considers those from Minas 
Gerais, Paraguay and Argentina to represent Phrynophyas 
liebes (Hyla hebes) . 

Remarks. In 1956, in a comprehensive study of the frogs in- 
cluded under the name Hyla venulosa, Duellman suggests that 
the neotropical hylid frogs with two lateral vocal sacs situated 
behind the angle of the jaw, without a frontoparietal fontanelle, 
with free cranial skin and bifid maxillary teeth should be placed 
in the genus Phrynohyas of Fitzinger. He has split Hyla reint- 
losa auct. into a number of species, two of which are represented 
in Venezuela, lie uses the name Phrynohyas inf/ens for the form 
that occurs in the Maracaibo Basin and Phrynohyas zonata for 



RIVERO: SALIENTIA OF VENEZUELA 129 

the one that occurs in the Coastal Range and elsewhere in north- 
ern South America. He considers Hyla tibiatrix Laurenti a 
nomen dubium for the following- reasons: 1 

1. Hyla tibiatrix Laurenti is based on Seba I. 71. 1-2 and this 
drawing may represent any of the forms comprised by //. 
venulosa auct. 

2. The type locality given is America. 

3. The figure in Seba shows no color pattern and it might 
possibly show a casqued head. 

There does not seem to me to be much doubt regarding the 
identity of Seba's picture with Hyla venulosa auct. I believe 
the casque would have been mentioned if it existed and no refer- 
ence is made to it; to me it does not appear that a casque is 
represented in the drawing. 

[n their paper of June 1951, Dunn and Stuart express them- 
selves regarding the restriction of type localities. Re-wording 
articles 29 and 31 of the Laws of Nomenclature, they state that 
". . . if a species is divided into two or more restricted species, 
its valid name must be retained for one of the restricted species" 
and further on ". . . the division of a species into two or more 
restricted species may automatically involve a concomitant selec- 
tion or restriction of type localities as well as of type speci- 
mens ..." 

I believe it would have been much simpler and more 
convenient, to restrict the type locality rather than to consider 
Hyla tibiatrix a "nomen dubium." As there are only three forms 
of the presently discussed Phrynohyas in South America, and of 
these, at least one is probably a race of another (according to 
Duellman, hebes, of S. Brasil, Argentina and Paraguay is per- 
haps a race of zonata of the north of South America), the third 
being restricted to the Maracaibo Basin, the probabilities are that 
Seba's drawing represents zonata. If we declare "nomina, dubia" 
all the doubtful forms whose type locality is America, very many 
well known names of frogs and other animals will have to dis- 
appear from our nomenclature. 

In trying to relate Hyla taurina with the Phrynohyas and 
other related groups, I ran into a few difficulties. H. taurina has 
bicuspid teeth, but this character is shared with H. boans, H. 
raniceps, H. septentrionalis and according to Noble (1931: 123) 



i In Bull. Zool. Nomencl., 12 (5), 1956, Duellman makes a form request to the 
International Commission to declare //. tibiatrix Laurenti a nomen dubium. 



130 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

with most metamorphosed Amphibia. Cope (1865b: 108) in- 
cluded Hyla taurina (as Osteocephalus) among the frogs with a 
frontoparietal fontanelle, but in his "Batrachia of North Amer- 
ica" (1889: 322) he rearranged his key and " Osteocephalus" 
fell among the hylids with no fontanelle. I have examined one 
specimen of this species, two of H. boans (H. maxima auct.) and 
one each of H. raniceps, H. crepitans, H. rubra, H. acuminata 
(H. phrynoclerma auct.), H. mesophaea, H. buckleyi, H. boulen- 
geri, H. leprieurii, H. imitatrix and H. vilsoniana. All the frogs 
having two lateral vocal sacs 1 (taurina, mesophaea, buckleyi, lep- 
rieurii, imitatrix) lacked the frontoparietal fontanelle, but there 
are four species, without lateral vocal sacs (boans, rubra, acumi- 
nata, boulengeri) in which a fontanelle could not be detected. 
H. taurina agrees with H. venulosa auct. (Duellman's Phryno- 
hyas) in having a glandular skin (males) but in this respect it 
differs from Hyla mesophaea and most H. imitatrix, both of which 
on the other hand, have lateral vocal sacs and lack the fronto- 
parietal fontanelle. If the characters of the skin and teeth are 
not considered for characterizing Phrynohyas, we are left with 
the lateral sacs, and the lack of fontanelle and of skin ossification 
to distinguish the group from other hylids. 

H. taurina agrees with H. leprieurii (with which it is very 
closely related) and H. buckleyi in the character of the vomerine 
teeth (/~ \-shaped in the first two, / \-shaped in the latter), 
a character they share with H. boans, H. geographica and H. crepi- 
tans but not with H. venulosa auct., H. imitatrix and H. meso- 
phaea. Is it possible that there are two groups of frogs with 
lateral vocal sacs and solid skull roof, each one of them having 
a different origin? This is only a possibility but one that cannot 
be discarded. Liu has expressed the opinion that "the evolution 
of the vocal sac does not always parallel the steps of the evolu- 
tion of species or other groups, the exceptions being due to 
independent origin of the vocal sacs or to acceleration or retarda- 
tion of evolution in this character" (1935 : 29) . 

This discussion does not, by any means, prove that Duellman's 
conclusions are incorrect. It does prove, however, that a detailed 
study of related forms is an absolute necessity before establishing 
any generic or even subgeneric boundaries. Until such a study is 
made, the name Hyla tibiatrix is maintained for the species un- 
der consideration. The type locality is hereby restricted to Suri- 
nam. 

i Following the classification of Liu, 1935. 



RIVERO: SALIENTIA OF VENEZUELA 131 

The lack of frontoparietal fontanelle in H. boulengeri, H. 
rubra, H. phrynoderma and perhaps in other Hyla of the rubra 
group suggests the need for further and more detailed study of 
these frogs, which are also characterized by lacking any web 
between the fingers. 

Dr. Mertens informs me that the Hyla nigromaculata men- 
tioned by Boettger in his 1895-6 report is really Hyla tibiatrix. 

According to Rohl (1949) the common name of this frog in 
Venezuela is "rana lechera." 

Hyla tibiatrix ingens (Duellman) 

Phrynohyas ingens Duellmaai, 1956, Misc. Publ. Mus. Zool. Univ. Mich., 
No. 96: 22, pi. ii, fig. 2. 

1 (C.N.H.M. 2604) Orope, 08. 

4 (U.M.M.Z. 55567-70) La Fria. 

1 (U.M.M.Z. 57397) Sabana de Mendoza. 

Description. Hyla tibiatrix ingens is distinguished from the 
typical form by having a more or less uniform color above (brown 
with darker brown dots, especially on the posterior half) and by 
its greater size and thicker, more glandular skin. 

Measurements. 9 snout-vent 105 ; head length 30 ; head 
breadth 33.5; femur 49.4; tibia 54.3. 

Habits. The following paragraph is quoted from Duellman, 
who quotes from Baker : ' ' Two pairs of these big frogs were copu- 
lating in a peccary wallow pond near a big tree. The male makes 
a loud noise like dropping a rock into a vessel partially filled 
with water. After heavy rains, eggs on surface (of water) formed 
films two or three feet across." 

Range. The Maracaibo Basin. 

Remarks. There is a 112.5 mm. female specimen at U.M.M.Z. 

Hyla vilsoniana meridensis ssp. n. 

Hyla wilsoniana, Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 40. 

Type. Museum of Comparative Zoology No. 2527, a 9 from 
Merida, 1630 m. 

Diagnosis. Hyla vilsoniana meridensis differs from the typical 
form in having white spots or marbling on the posterior part 
of the thighs, flanks, axillae, underside of the hind limbs, hidden 
portions of the tibiae and tarsus and dorsal part of the foot. 

Description. Snout short, rounded ; tongue broad, % free and 
nicked behind ; vomerine odontoids in two transverse groups be- 
tween the round or oval choanae ; eye diameter greater than 



132 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

distance between eye and nostril, but shorter than the snout; in- 
terorbital space slightly broader than an upper eyelid; canthus 
rounded, indistinct ; loreal moderate^ oblique, not concave ; tym- 
panum !/2 the eye diameter ; a slight supratympanic fold ; two 
faint metacarpal tubercles; one flat inner metatarsal tubercle 
but no outer ; subarticular tubercles of fingers and toes moderate 
but rather flat; fingers with a short basal web, the first shorter 
than second ; larger disks about the same size as the tympanum ; 
toes taken in order from first to fifth exhibit the following 
phalanges free of web: 1, 1, 1%, 3, 1 ; heel of the adpressed hind 
limb extends to the middle of the eye. Skin above, smooth. Be- 
low, with large and very distinctive flat granules; a transverse 
pectoral fold. 

Color. Above, slate or bronze colored (said to be green, alive) ; 
axillae, groins, anterior and posterior aspects of the thighs, 
ventral portions of the tibiae, hidden portions of the tibiae and 
tarsus and dorsal part of the foot spotted with white. Below, 
yellowish orange (in life?) or slate, the granules lighter. 

Measurements. 9 Snout-vent 42 ; head breadth 13 ; head length 
12.5; femur 18; tibia 20. 

Additional Localities. Chama (U.S.N.M. 71114; U.M.M.Z. 
3574; C.N.H.M. 3573-4) ; La Culata (A.M.N.H. 10671-3) ; Vene- 
zuela (A.M.N.H. 10666-7, 10669). 

Range. The Subtropical Zone of the Merida Andes. 

Remarks. The eye is equal to distance between eye and nostril 
in a few specimens and in some the interorbital space is equal to 
an upper eyelid. The web may be 1*4, 114, IV2, 2%, 1V4- 

Some frogs (A.M.N.H. 10671-3) are labeled "La Culata, N. W. 
Ecuador." My reasons for including the specimens here are as 
follows : 

1. No place with such a name has been found in Ecuador. 

2. Hyla vilsoniana has not been reported otherwise from Ecua- 
dor. 

3. If Hyla vilsoniana does occur in Ecuador it is not likely 
to be on the western side of the Andes. 

4. There is a place by the name of La Culata in A'enezuela. 

5. The specimens agree in all details with Venezuelan animals 
rather than with those from Colombia. 

6. Specimens with approximately the same date and received 
from the same dealer were collected in La Culata, Vene- 
zuela. 



RIVERO : SALIENTIA OF VENEZUELA 133 

Dunn (1944b: 72) has placed H. vilsoniana Cope in the synon- 
omy of H. labialis Peters, 1863. Whether or not this is cor- 
rect I am not prepared to say, but I am well aware of a large, 
vilsoniana-like frog that occurs in Gutierrez, Colombia. In some 
museums, specimens of this species are determined as H. labialis, 
while in others it is known as H. gularis. Three alternatives are 
possible : 

1. That H. gularis is a distinct species while H. vilsoniana is 
a synonym of H. labialis. 

2. That H. gularis is a synonym of H. labialis while H. vil- 
soniana is a distinct species. 

3. That H. labialis is a distinct species while H. gularis is 
a synonym of H. vilsoniana. 

As both forms are very similar and it is not possible to come 
to any conclusion without examining the types, I have decided 
for the present to use the name vilsoniana on the basis that in two 
cases out of three this should be the proper name to use. 

All the 12 Venezuelan specimens examined show the charac- 
teristic spotting of the limbs that characterize the race. It is 
possible that differentiation of H. v. meridensis occurs at or near 
the Depression of San Cristobal in Venezuela. Colombian speci- 
mens available for comparison (A.M.N.H. 16738-54, Bogota, 
5456-61, 53822-4, cotypes, nr. Bogota; M.C.Z. 7299-7308, 20936-7, 
24196, Bogota and San Pedro, Antioquia) do not show any limb 
or flank spotting, except perhaps A.M.N.H. 53822, where there is 
some indication of it. 

Hyla minuta Peters 

HyJa minuta Peters, 1872, Monatsb. Akad. Wissensch. Berlin: 680: New 
Freiberg, Brasil. 

14 (U.P.R. 153-66) Pto. Ayacucho, vi.50. 
1 (U.P.R. 167) Sanariapo, vi.50. 
Description. Snout short, almost semicircular, little longer 
than the eye diameter ; tongue oval, slightly nicked and free 
behind ; vomerine odontoids in two short, slightly oblique series 
between the round choanae, their posterior extremities directed 
inwards ; eye diameter greater than distance between eye and 
nostril ; interorbital space much broader than an upper eyelid ; 
canthus rounded ; loreal almost vertical, slightly convex ; tym- 
panum indistinct, partly covered by a loose supratympanic fold, 
about !/2 the eye diameter ; no external rudiment of pollex ; 
subarticular tubercles small and indistinct ; fingers taken in order 



134 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

from first to fourth exhibit the following phalanges free of web : 
almost all, 1% to IV2, 2%, 2 to 2^4 ; disks small, about the same 
size as the tympanum ; no tarsal fold ; toes depressed, fully webbed 
except in the third and fourth where the membrane does not 
extend to the disks ; disks of toes not much more expanded lat- 
erally than the width of the phalanges; heel of the adpressed 
hind limb extends to between eye and nostril. Skin above, 
smooth ; a fold across the chest ; male with a very large, subgular 
vocal sac. 

Color. Above, straw colored or yellowish brown with definite, 
light-margined, darker spots of irregular form and size. The 
spots are generally saddle or chevron-shaped, but hour-glass, 
triangular, or round ones are often found. The marginal lines 
are fine and of a golden or cream color, but spots without the 
marginal lines are sometimes present although apparently not 
in the same individual having the margined ones. Very few of 
the specimens are uniformly brownish or grayish. A small trans- 
verse line at the anus and a short, longitudinal one along the 
heel seem to be constant characters. The thighs are uniformly 
colored, the rest of the hind limbs generally marked with light 
margined spots ; forelimbs equally marked. Ventral surfaces 
immaculate. 

Measurements. Snout-vent $ 23, 9 26 ; head breadth $ 8, 
9 8.9 ; head length $ 7.5, $ 8 ; femur $ 11, 2 11 ; tibia $ 
12.5, $ 12. 

Habits. Hundreds of these frogs were seen in a shallow inun- 
dated area between Puerto Ayacucho and Sanariapo. All were 
males, however, and upon returning to the place two nights 
later, the animals had deserted and no eggs were found. Most 
of the frogs were calling from grass blades or from the water, 
which was not deeper than their bodies. The pool was actually 
in a grassland but there was high forest some yards to the east 
and west. 

In Marahuaca, collecting was carried out in a similar situation 
but the forest at this place was heavy and very humid. 

Additional Localities. Temiche (U.P.R. 169-75, 260-262). 

Range. The Venezuelan Guayana. Trinidad and the Guianas 
to southern Brasil. 

Remarks. The specimens from Temiche are smaller, much 
darker, and lack the ornamental coloration of the lowland group. 
Probably they should be given a racial name. 

Specimens from Rio de Janeiro are also smaller and probably 
slightlv less webbed than the animals from Puerto Ayacucho. 



RIVERO : SALIENTIA OP VENEZUELA 135 

Hyla misera Werner 

Ilyla misera Werner, 1903, Zool. Anz., 26: 2:12: Caracas, Venezuela; Nieden, 
1923, Das Tierreich. Anura, I: 263; Lutz, A., 1927, Mem. Inst. Osw. 
Cruz, 20: 39, 43, pi. xi, fig. 16 and pi. xv, fig. 37. 

1 (M.C.Z. 26150) Cerro Cosine, x.39. 

Description. Head as broad as long; snout short, rounded, 
slightly longer than the eye diameter; tongue rounded, nicked 
and slightly free behind; vomerine odontoids in two short, 
slightly oblique groups between the moderate choanae, their pos- 
terior extremities directed inwards; eye diameter greater than 
distance between eye and nostril ; interorbital space broader than 
an upper eyelid; canthus indistinct; loreal slightly oblique, not 
concave ; tympanum not very distinct, little less than y 2 the eye 
diameter; subarticular tubercles of fingers and toes not very 
distinct ; fingers short, taken in order from first to fourth they 
exhibit the following phalanges free of web : all, iy 2 , 2y 2 , y 2 ; 
a tarsal fold ; one small, oval inner, but no outer, metatarsal 
tubercle ; toes fully webbed, except for the third and fourth in 
which the web does not extend beyond the penultimate articula- 
tion ; disks of the toes smaller than those of the fingers ; heel of 
the adpressed hind limb extends to between eye and nostril. 
Skin above smooth. Male with a subgular vocal sac. 

Color. Above, brown with obscure longitudinal markings of 
darker color and dark punctuations under a lens ; a dark canthal 
streak. Below, light brown. 

Measurements. $ Snout-vent 20; head breadth 6.2; head 
length 6.2; femur 10; tibia 11. 

Additional Localities. Caracas (Nieden, 1923 = Werner, 
1903?); Caracas and Maracay (Lutz, 1927); Kunana, Perija 
Mts. (Aleman, 1953). 

Range. The Falcon Region and the Coastal Range, and the 
Parija Mts. 

Remarks. This form is rather similar to Hyla minuta, from 
which it can be distinguished by its smaller size, straighter loreal 
region, distinct tarsal fold and less ornate coloration as well as 
by the lack of the anal and heel lines that are present in minuta. 

The Venezuelan specimen differs from the original description 
and from Colombia (M.C.Z. 16077, Tucurinca; M.C.Z. 13751-3, 
Sevilla) and Trinidad material (M.C.Z. 17809-17900, St. Augus- 
tine) in lacking the very distinct darker coloration of the loreal 
region and flanks. The Trinidad and Colombian animals, which 
look identical to me, have a white line that extends from tip of 



136 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

the snout, over the outer margin of the upper eyelid to the sacral 
region or beyond. This may have existed in the Venezuelan 
example but its presence is not evident in the present condition 
of the animal. 

Hyla misera is closely related to Hyla microcephala, from 
which it is distinguished by lacking the well defined dorsal pat- 
tern of the latter (present but very faintly indicated in preserved 
II. misera). The dark lateral coloration and the dorsolateral 
white line that divide the dorsal and ventral colors of misera are 
present in microcephala, but the lateral color does not seem to 
extend as far down ventrally as in misera. 

In northeastern Colombia, //. misera is found as far west as 
west of Sierra de Santa Marta, and it has been recently collected 
by Aleman at the Perija Range of Venezuela. It seems that H. 
microcephala meets misera somewhere in the Magdalena Valley 
and that the latter is only a race of microcephala. Until a more 
complete examination of material is made, I have preferred to 
continue using the binomial. 

Several races of Hyla microcephala have been described from 
Central America. 

Hyla orophila planicola Lutz and Lutz 

Hyla (Splioenoliyla) planicola Lutz, A. and B., 1938, Ann. Acad. Bras. Sci., 
10 : 182 : Recreio dos Bandeirantes, Federal District, Brasil. 
1 (M.C.Z. 19917) Orinoco R., below Barrancas, ii.35. 
Description. Head small, flat ; snout short, rounded at the tip 
and projecting beyond the mouth ; tongue cordiform, slightly 
indented and free behind ; vomerine odontoids almost imper- 
ceptible, probably in two round groups behind and between 
the small, round choanae ; eye small, not protruding, its diameter 
equal to distance between eye and nostril ; interorbital space 
about twice as broad as an upper eyelid ; canthus apparently well 
defined; loreal vertical, not concave; tympanum not very dis- 
tinct, V2 the eye diameter; no projecting external rudiment of 
pollex, but a fairly distinct tubercle at the base of the first finger ; 
subarticular tubercles indistinct; first finger much shorter than 
second ; taken in order from first to fourth, the fingers exhibit 
the following phalanges free of web : 2, 1, 2, IV2 ; largest disks 
about as large as the tympanum ; an oval, inner metatarsal tu- 
bercle ; toes fully webbed, with the exception of the fourth where 
two phalanges are left free ; heel of the adpressed hind limb 
extends to the anterior corner of the eye. Skin above, shagreened ; 



BIVERO: SALIENTIA OP VENEZUELA 137 

a pair of horizontal flaps at the sides of the anus. Male with 
an external vocal sac. 

Color. Above and below, yellowish white; a light brown can- 
thai streak. 

Measurements. $ snout-vent 24; head breadth 7; head length 
6.8; femur 11.5; tibia 12. 

Range. In Venezuela known only from this specimen. British 
Guiana to Bolivia and southern Brasil. Hyla orophila orophila 
occurs in the mountains of southern Brasil while Hyla orophila 
planicola is restricted to the lowland. These as well as Hyla 
aurantiaca surela Cochran, from Parana, are considered separate 
species by Goin (1957). 

Remarks. The Venezuelan individual has slightly more web- 
bing on the toes than animals from southern Brasil. The snout 
is also blunt, instead of triangular and pointed, but this is prob- 
ably because it has received a blow in front. The almost total 
absence of vomerine teeth in this specimen is probably an abnor- 
mality. Since it has a distinct tympanum it cannot be considered 
as a specimen of Hyla hdbra (Goin). 

Hyla aurantiaca Daudin, 1803, is preoccupied by Hyla auran- 
tiaca Laurenti, 1768. This latter is based on Seba /, plate 71.3 
which was used for the description of Hyla boans. Therefore, I 
use the name Hyla orophila Lutz and Lutz for the species hith- 
erto known as Hyla aurantiaca Daudin. 

Goin, 1957, erects the genus Sphoenohyla 1 for the group of 
frogs comprising Hyla orophila, on the basis of a different vocal 
sac, a backward projecting process in the ischium, and probable 
different habits. In trying to put boundaries to the genus, I 
dissected a few frogs, not only from South America, but also 
the Old World H. arborea, the genotype of Hyla, and H. aurea 
from Australia. These two show a rounded ischium, more or 
less as figured for the North American H. cinerca (Goin, 1957: 
14, fig. 2), but Hyla rubra and to a greater extent Hyla crepitans 
show a dorsal, posterior projection as figured for Sphoenohyla 
aurantiaca (Goin, op. cit.). In Hyla tibiatrix and //. brunnea. 
(Jamaica) the ischium is more extensive posteriorly than in the 
European or Australian frogs, but there is no notch or indenta- 
tion to divide the bone into dorsal and ventral projections. If 
the character of the ischium is to be taken as generic, perhaps all 
the South American Hyla can be separated from the others into 
a different genus. 

i Created as a subgenus by Lutz and Lutz, 1938. 



138 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Liu, 1935, classifies the vocal sacs into median subgular (in- 
ternal or external), paired subgular (internal or external), and 
paired lateral (internal or external). According to him (op. tit., 
pp. 33-35), all these, except the paired subgular external, are 
represented in the genus Hyla. In two species, the vocal sacs are 
totally absent. Hyla aurantiaea auct. was not studied, but Hyla 
nana, considered by Goin of the Sphoenohyla group, shares a 
median, subgular external sac with 59 of the 88 species men- 
tioned. In comparing Sphoenohyla with Hyla, Goin does not 
mention what species he has compared. However, considering 
the different vocal sacs that are known to occur within the genus 
as presently known, I do not think it wise to accept the separation 
of Sphoenohyla, especially since the character of the ischium is 
not apparently an exclusive character of the group. 

Hyla battersbyi sp. n. 
Figure 11 

Hyla leucomelas Giinther (not Dumeril and Bibron), 1858, Cat. Batr. Sal. 
Brit. Mus. : 110: Caracas. 

Type. British Museum of Natural History No. 53.2.4.165, a 6 
from Caracas, Venezuela. 

Diagnosis. A Hyla with transverse vomerine odontoids ; well 
defined tympanum, about ^2 the eye diameter ; no external rudi- 
ment of pollex; % webbed fingers and grayish color minutely 
dotted with reddish brown on all the upper and lower surfaces 
except the forelimbs and belly. 

Description. Snout rounded; tongue circular, entire and ad- 
herent ; vomerine odontoids in two transverse, juxtaposed groups 
on level with the posterior margins of the small, round choanae ; 
eye diameter equal to distance between eye and nostril ; inter- 
orbital space somewhat broader than an upper eyelid ; canthus 
moderate ; loreal very slightly oblique, concave ; tympanum well 
defined, about y 2 the eye diameter ; a supratympanic fold ; no 
external rudiment of pollex ; first finger shorter than second ; 
subarticular tubercles rounded, distinct ; fingers taken in order 
from first to fourth exhibit the following phalanges free of web : 
practically all, 1%, 2y 2 , 2y 2 ; larger disks as large as the tym- 
panum ; an almost imperceptible tarsal fold ; one internal, oval 
metatarsal tubercle ; toes exhibiting the following free phalanges : 
1%, iy 5 , 1V4, 2 to 2y 2 , iy 2 ; heel of the adpressed hind limb ex- 
tends to between eye and nostril. Skin above smooth, below 



RIVERO : SALIENTIA OF VENEZUELA 



139 



granular on the belly and thighs ; a transverse pectoral fold ; 
male with a slight subgular vocal sac. 

Color. Above, light brown with numerous small but sharply 
defined reddish brown dots that appear more profusely on the 
top and sides of the head, flanks and limbs; a dark canthal and 
supratympanic streak ; loreal whitish with dark dots ; limbs 
crossbarred. Belly, ventral surface of forelimbs and granular 
ventral part of the hind limbs uniformly light brownish, the 
rest of the hind limbs and the throat minutely dotted ; a pair 
of callous, white spots on the buttocks. 




c=^ 



Fig. 11. Ilyla battersbyi sp. n. Type BM (NH) 53.2.4.165. 



Measurements. $ Snout-vent 33, head breadth 10.5 ; head 
length 10.5; femur 17; tibia 17. 

Remarks. In the dotting of its body surfaces this species agrees 
somewhat with Hyla rufopunctata Andersson, Hyla aluminiata 
Andersson (which may be the young of something else), Hyla 
palmeri Blgr., Hyla festae Peracca, Hyla albida Melin, Hyla 
phyllognatha Melin, Hyla columbiana Bttgr. and the young of 



140 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

Hyla geographica Spix. From the first two and the young Hyla 
geographica it differs in the well exposed tympanum and in the 
different position of the vomerine odontoids, from H. palmeri 
and H. festae in the form of the odontoids, slighter inclination 
of the loreal region, amount of webbing, smoothness of the skin, 
and in having shorter legs, smaller size and different coloration 
than festae. From H. albida it can be rapidly distinguished by 
the visible tympanum, adherent tongue, lack of external rudi- 
ment of pollex, etc., and from H. phyllognatha by the size of 
the tympanum, position of the vomerine odontoids, amount of 
webbing, lack of external rudiment of pollex and other charac- 
ters. With Hyla columbiana Boettger, this species agrees in 
several characters but it can also be readily differentiated by its 
well defined tympanum, completely adherent tongue and dif- 
ferent coloration of the ventral surfaces. While H. battersbyi 
is profusely dotted on the throat and ventral surface of the hind 
limbs, Hyla columbiana is uniformly colored on these regions 
or more usually, distinctly variegated or marbled on the throat 
and belly. 

I believe Hyla variabilis Boulenger is a synonym of Hyla 
colvmbiana Boettger. 

CORYTHOMANTIS VENEZOLANA MertenS 

Ilyla nigrornaculata Boettger, 1896, Ber. Senckenb. Naturf. Ges. : LIY. 
Corythomantis brunoi Mertens, 1926, Senckenbergiana, 8: 137. 
Corythomantis venezolana Mertens, 1950, Senckenbergiana, 31: 1: San Fer- 
nando de Atabapo, Upper Orinoco. 

No material examined. 

Original Description. Skull forming a flat, bony helmet, with 
a rougher dorsal side and rather projecting rims; snout pro- 
jecting over the mouth opening ; pupil roundish, horizontal ; 
tongue heart-shaped, slightly free, clearly notched behind; vo- 
merine odontoids in two small groups between the choanae ; no 
palatine or parasphenoid teeth ; canthus distinct ; a vertical bony 
ridge in front of the eye ; tympanum about % the eye diameter ; 
fingers with a short web ; toes % webbed ; disks of the fingers y 2 
the size of the tympanum, of the toes, % ; subarticular tubercles 
larger in fingers than in toes, mostly single ; inner side of first 
finger with a large, blunt tubercle ; a rather large, outer and a 
small inner metatarsal tubercle ; heel of the adpressed hind limb 
extends to the anterior corner of the eye. Skin above, smooth. 
Belly granular. 

Color. Middle of the back bright brown with darker brown 
marblings; thighs yellowish, with brown network; upper side 



RIVERO: SALIENTIA OF VENEZUELA 141 

of limbs transversely banded; ventral side, gray, without pattern. 

Measurements. £ ? snout -vent 58; anterior limb 24; posterior 
limb 83. 

Range. Only known from San Fernando de Atabapo. 

Remarks. This species can be distinguished from Corythoman- 
tis greeningi Boulenger by the webbing in the fingers, the nar- 
rower and more protuberant snout, the heart-shaped tongue, the 
not indented posterior margin of helmet and the smaller disks. 

Carvalho (1941) pointed out that the specimen collected by 
Hubner at Sn. Fernando de Atabapo and discussed by Mertens 
as Corythomantis brunoi (1926) probably represented a juvenile 
of Trachycephalus nigromaculatus Tschudi. His conclusion was 
based on the fact that of the helmeted hylids comprising the 
genera Diaglena, Triprion, Pternohyla, Corythomantis, Aparas- 
pkenodon and Trachycephalus, 1 only the latter was described 
with webbed fingers and barred hind limbs. 2 The total absence 
of palatine and parasphenoid odontoids, a character which Mer- 
tens' specimen shares only with Corythomantis, is apparently 
attributed to immaturity of the example, which however, was 
58 mm. in length. 

In 1950 Mertens described Corythomantis venezolana on the 
basis of his former Corythomantis brunoi. I believe it is safe to 
conclude that Mertens re-examined his specimen in the light of 
Carvalho 's comments and that although it was webbed and 
barred, he was not convinced of the identity of the animal with 
the genus Trachycephalus. Like Aparasphenodon, the latter is 
described as having palatine teeth, while Corythomantis lacks 
both palatine and parasphenoid teeth. 

Examination of the genera mentioned before reveals relation- 
ships between some of them. The pupils have been described as 
horizontally oblong in Aparasphenodon, "transversely oval when 
expanded and subquadrangular when contracted" in Diaglena, 
a transverse slit in Trachycephalus, rhomboidal in Corythoman- 
tis, a vertical slit in Triprion and round in Tetraprion? I do not 
consider pupil shape of generic significance if not accompanied 
by other characters. If this character is not considered valid 
for separation, two forms can probably be ranked together: 
Diaglena and Triprion. 4 The form of their casques is very similar 



i In writing about these helmeted frogs, Carvalho does not mention Pternohyla. 

^Diaglena, Corythomantis, Aparasphenodon and Pternohyla actually have short 
webs between the fingers. . ... _. , , 

STetraprion Steineger and Test, 1891, was synonymized with Diaglena dj 
Boulenger 1891 and with Triprion by Giinther, 1900, but revived by Myers. 1942. 

4 And probably Tetraprion. which I have not had the opportunity to examine. 



142 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

and with little doubt derived one from the other, perhaps Trip- 
rion from Diaglena by further specialization of the skull structure 
and disappearance of the palatine odontoids. These, and 
Tetraprion, are the only helmeted frogs possessing parasphenoid 
odontoids and they also have characteristic vomerine odontoids 
and general physiognomy in common. 

The relationship of the other forms is not as clear. Aparas- 
phcnoclon is said to have lateral vocal sacs (Cochran, 1955 -AT), 1 
a character it more or less shares with Trachycephalus 2 and mem- 
bers of the Hyla tibiatrix group. The vomerine odontoids are 
more or less like those of Hyla taurina and Trachycephalus but 
different from the other helmeted frogs here described. The 
palatine odontoids are shared with Trachycephalus and with the 
Central American Diaglena, Triprion and Ptemohyla. For the 
present it is perhaps more convenient to keep Ptemohyla, Trachy- 
cephalus and Aparasphcnoclon in separate genera, although the 
relationships of the latter with Triprion (including Diaglena), 
seems to be quite close. 

Through the courtesy of Dr. Ernest Williams, I was able to 
examine the skeleton of one of the only two specimens of Triprion 
at M.C.Z., and examine the sacral diapophyses of one of each of 
Diaglena, Ptemohyla, Aparasphcnoclon, Corythomantis, Gar- 
beana and a few other hylids. Dr. Doris Cochran kindly made 
available a specimen of Trachycephalus nigromaculatus from 
Brasil. 

In Diaglena, Triprion, Corythomantis, Ptemohyla and 
Aparasphenoilon, the sacral diapophyses are considerably dilated, 
and there is in addition a large cartilaginous alar expansion on 
their free sides. Furthermore, the ilium, instead of ending at 
the diapophysis, is not apparently firmly attached to it and con- 
tinues anteriorly under the alar expansion for a length that 
varies according to the genus. In Corythomantis the ilium ex- 
tends forward of the diapophysis for only a short distance, but 
in the other four genera the anterior projection may be quite 
long. In this respect, Corythomantis may be the most primitive, 
but the examination of only one specimen is not enough to permit 
any conclusion. It should be remembered, however, that Cory- 
thomantis is also the only one of the four genera not having 
palatine or parasphenoid teeth. 

i A male specimen from Rio de Janeiro (M.C.Z. 25694) shows a subgnlar vocal 
sac which is perhaps more expanded on the sides. 

2 Although it does not quite show this character, as in the latter the sacs are 
posterior to the angle of the jaw, while in Aparasphenodon they seem to he 
subgular. 



RIVERO : SALIENTIA OF VENEZUELA 143 

In Trachyccphalus, the alar expansion of the diapophysis is 
not too well developed and the ilium does not extend forward 
beyond its anterior margin. If considered to belong to the 
Triprion group, Trachycephalus may be thought to be the most 
primitive in ilium development, but it has palatine odontoids 
and two distinct anal flaps that it shares with the Brasilian 
Hyla langsdorffi, a large Brasilian frog with free cranial derm 
but. great similarity with T. nigromaculatus. These forms may 
or may not be within the main line of descent of the Triprion 
group. It appears, however, that the genera Triprion (inc. 
Diaglena), Corythomantis, Ptemohyla, Aparasphenodon and 
Tetraprion constitute a natural group characterized by having 
the skin of the head involved in the cranial ossification and an 
ilium that extends beyond the anterior margin of the sacral 
diapophysis. Ptemohyla has undergone considerable specializa- 
tion and may be an early offshoot from the main line. 

Incidental to the study of the helmeted frogs of the aforemen- 
tioned groups, I had the opportunity of examining specimens of 
Hemiphractus braconnieri and Garbcana garbei from Ecuador. 
Prom the description of the latter and examination of the Ecua- 
dorian specimen (M.C.Z. 19662), I gather it is conspecific with 
Hyla boulcngeri. As in //. braconnieri, the sacral diapophyses of 
G. garbei are but slightly dilated, there is no alar expansion, and 
the ilia do not extend forward beyond the diapophyses. 

Key to the Species of Gastrotheca Reported from Venezuela 

I. Derm of head involved in cranial ossification ovifera 

II. Derm of head free from cranial ossification williamsoni 

Gastrotheca ovifera (Weinland) 

Notodelphys ovifera Weinland, 1854, Archiv. Anat. Physiol.: 473, pi. xvii- 

xix: Venezuela; Lichtenstein and Martens, 1856, Nomencl. Rept. Am- 

phib. : 36. 
Ophistodelphia ovifera Ernst, 1877, Flora y Fauna de Ven. (Caracas) : 

281. 
Nototrema ovifcrum Nieden, 1923, Das Tierreich. Anura I: 323; Lutz, A., 

1927, Mem. Inst. Osw. Cruz, 20: 39, 44. 
Hyla vogli Miiller, L., 1938, Zool. Anz., 121 : 284. 

4 (U.S.N.M. 121164-7) Colonia Tovar. 
Description. Skull rugose ; derm of head involved in cranial 
ossification; two transverse ridges on the occipital region and 
sometimes two spines on the supratympanic ridge ; snout short, 



144 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

rounded ; tongue circular, about Vi f ree an d nicked behind ; 
vomerine odontoids in two transverse groups between the small, 
round choanae ; eye diameter equal to distance between eye and 
nostril ; interorbital space much broader than an upper eyelid ; 
canthus ridged, inclined downwards toward the front ; loreal 
moderately oblique, concave, rugose ; tympanum y 2 the eye diame- 
ter; a fold that arises on the supratympanic ridge runs for a 
short distance posteriorly; one oval, inner metacarpal tubercle; 
first finger slightly shorter than second, free; other fingers with 
a short basal web ; largest disks as large as, or larger than, the 
tympanum ; tarsal fold slight or absent ; toes taken in order from 
first to fifth exhibit the following phalanges free of web : 2, V/ 2 , 
1% to 2, 3, iy 2 ; heel of the adpressed hind limb extends to be- 
tween eye and nostril . Skin above, shagreened ; a pair of 
crescent-shaped folds on the posterior part of the body. Male 
with a large, subgular vocal pouch. 

Color. Above, gray with scattered dark spots; usually a pair 
of dark lines that begin on the occipital ' ' horns ' ' and extend pos- 
teriorly to the dorsal folds; a series of black spots along the 
flanks and on the anterior and posterior parts of the thighs where 
they sometimes form crossbars ; between the spots, the thighs are 
rose or salmon. Below, gray, darker on the throat (6 $ ?). 

Measurements. S snout-vent 50 ; head breadth 20 ; head length 
17 ; femur 28 ; tibia 28. 

Additional Localities. Cerro Avila (U.C.V. 29) ; Hacienda Los 
Venados, Avila (Miiller, L., 1938) ; Puerto Cabello (Licht. and 
Mart., 18.16 [= Weinland?] ) ; Vene/.uela (Nieden, 1923; Lutz, 
1927 [= Weinland?]). 

Range. The Coastal Range. ^Central America to N. Venezuela. 

Remarks. U.C.V. 29 is very dark gray all over the body; the 
lateral spots are very obscure and the spots on the thighs form 
well defined crossbars. No reddish or salmon color is visible. 
The tympanum is % the eye diameter. 

The Central American Opisthodelphis ovifera of Giinther, 
1858, should, according to Taylor, probably be referred to Ano- 
theca coronata (Stejneger). 

From the description it is evident that Hyla vogli Miiller is a 
synonym of this species. 

Gastrotheca williamsoni Gaige 
Gastrotheca iclUiamsonl Gaige, 1922, Oee. Papers Mus. Zool. U. Mich., no. 
107: 1-3: San Esteban, Venezuela; Lutz, A., 1927, Mem. Inst. Osw. 
Cruz, 20: 40; Rohl, 1949, Fauna Deser. de Yen., ed., 2: 402, fig. 182. 

1 (U.M.M.Z. 55559, type) Sn. Esteban. 



RIVERO : SALIENTIA OP VENEZUELA 145 

Description. Derm of head free from cranial ossification ; snout 
rounded ; tongue large, rounded, deeply notched behind ; vo- 
merine odontoids in two transverse groups between the choanae ; 
eye diameter equal to distance between eye and nostril ; canthus 
distinct, curved ; loreal concave ; tympanum obliquely oval, % 
the eye diameter ; a supratympanic fold to the shoulder ; a strong 
fold between elbow and wrist ; first finger as long as second ; 
fingers taken in order from first to fourth exhibit the following 
phalanges free of web : all, 1%, 2*4, 2 ; a faint tarsal fold ; a large, 
inner metatarsal tubercle ; subarticular tubercles well developed ; 
toes fully webbed, their disks slightly smaller than those of the 
fingers and as wide as the horizontal diameter of the tympanum ; 
heel of the adpressed hind limb extends to between tympanum 
and eye. Skin above, shagreenecl; pouch opening almost square 
and attached across the anterior end so it is open to the pouch 
only on the sides. 

Color. Above, reddish gray obscurely spotted with dusky; a 
darker spot on the back just behind the head ; an angular, dark, 
light-edged mark in front of the pouch with the edge directed 
forwards, and another faint, angular mark in front of the anus 
and within the pouch opening ; lower lip with indistinct, vertical 
bars ; a row of tiny, dark-edged tubercles connecting the eyelids 
across the interorbital space; flanks light, with dark vertical bars 
which slant slightly forward ; limbs banded with black ; back of 
the thighs dark, with small, light spots. Throat yellowish ; belly 
and lower surfaces of limbs greenish yellow. 

Measurements. Snout-vent 53; head breadth 20; head length 
17.5; femur 24; tibia 28.5. 

Range. San Esteban. 

Remarks. According to Gaige this specimen is closer to G. 
longipes Boulenger, from which it may be distinguished by the 
difference in coloration, shorter legs and greater webbing of 
fingers and toes. 

The color notes above are taken from the original description, 
for when I examined the type most of the markings described 
had faded. 

Nototheca pygmaea (Boettger) 

Nototrema pygmaeum Boettger, 1893, Ber. Senckenb. Naturf. Ges. : 40: 
Puerto Cabello ; Zool. Gart., 34 : 129. 
1 (U.M.M.Z. 55698) Quebrada Grande, Sn. Esteban. 
Description. Derm of head free from cranial ossification ; snout 



146 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

short, rounded ; tongue broad, adherent and slightly nicked 
behind ; vomerine odontoids in two small, very slightly oblique 
groups between the round ehoanae ; eye diameter equal to dis- 
tance between eye and nostril ; interorbital space as broad as or 
a little broader than an upper eyelid ; canthus moderate, curved ; 
loreal little oblique, faintly concave ; tympanum % the eye 
diameter ; a short supratympanic fold ; a rather flat, inner meta- 
carpal tubercle ; subarticular tubercles fairly prominent ; fingers 
free ; the first slightly longer than second ; larger disks as or 
slightly larger than the tympanum ; a row of tubercles along the 
tarsus ; an oval, inner metatarsal tubercle but no outer ; toes 
taken in order from first to fifth exhibit the following phalanges 
free of web : 2, 1% to 2, 1% to 2, 3y 2 , 2 ; heel of the adpressed 
hind limb extends to between eye and nostril. Skin above, smooth, 
slightly shagreened on the head ; female with a longitudinal fold 
that ends on the crevice-like opening of the breeding pouch 
posteriorly. 

Color. Above, head and anterior part of the back carmine, 
with blackish spots and designs, one of which forms a posteriorly- 
pointing triangle between the eyes ; sides of the head and throat 
with small, white flecks and dots ; limbs with blackish crosslines ; 
thighs gray, speckled with gray brown ; tibiae of female carmine ; 
feet gray. Below, yellowish, the throat and chin being of a 
weaker color; hind limbs speckled with gray brown (modified 
from Boettger). In preservation, specimens are more or less 
of a plain brown color. 

Measurements. Snout -vent 23; head breadth 8.5; head length 
8.5; femur 11.2; tibia 13. 

Additional Localities. Rancho Grande (U.C.V. 20). 

Range. The Coastal Range. 

Remarks. Nototheca pygmaea can be distinguished from all 
Venezuelan HyJa, with the exception of H. marahuaqurnsis, by 
having the first finger longer than the second. 

Bokerman (1950) has separated the Gastrotheca species with 
non-adherent head skin and longitudinally opened breeding 
pouch as the genus Nototheca (Genotype, Codonotiis fissilis 
Miranda Ribeiro). For the species whose breeding pouch opens 
through a supra-anal orifice, he preserves the name Gastrotheca, 
while for those "breeding pouch frogs" having the derm of the 
head involved in the cranial ossification he suggests tbe name 
Fleetonotus Miranda Ribeiro. 

This arrangement will leave Gastrotheca boliriana in that 



RIVERO : SALIENTIA OF VENEZUELA 147 

genus but the animals so far known as G. pygmaea and G. fitz- 
geraldi, will go into Nototheca, while G. ovifera, G. helenae, G. 
nice fori and probably G. marsupiata and G. monticola will be 
included in the genus Fleet onot us. The latter two have ossified 
head skin, but the derm is not actually involved in the cranial 
ossification. G. williamsoni and G. argenteoventris, which have 
free cranial derm, do not have, however, a longitudinally opened 
pouch, so that they will be difficult to allocate under this arrange- 
ment. 

Although recognizing the probability of a different origin for 
the frogs of the G. pygmaea group, and the convenience of sepa- 
rating them into another genus, I have preferred to maintain the 
other Venezuelan breeding pouch frogs in the genus Gastrotheca, 
until a more complete study of the "marsupial" frogs is made. 

Nototheca fitzgeraldi (Parker) may prove to be conspecific 
with N. pygmaea. A specimen from Tobago at M.C.Z. (27784) 
carries five large eggs on its back. 

Key to the Species of Phyllomedusa Reported from Venezuela 

I. First and second toes equal bicolor 

IT. First toe longer than second. 

A. Vomerine teeth present; parotids distinct; green above; spotted 
white on the flanks burmeisteri trinitatis 

B. Vomerine teeth absent; parotids indistinct; purple above; striated 
on the flanks h. hypocondrialis 

Phyllomedusa bicolor (Boddaert) 

Farm bicolore Boddaert, 1772, Epist. de Rana bicolore: 1, pi. iv, figs. 1-5: 
Surinam. 

2 (U.P.R. 185-6) Sn. Fdo. de Atabapo. 
Description. Head much broader than long; snout truncate; 
tongue entire, free behind ; vomerine odontoids in two short, 
slightly oblique series between the large choanae ; eye diameter 
shorter than distance between eye and nostril ; interorbital space 
almost twice as broad as an upper eyelid ; canthus angular ; loreal 
straight, excavated; tympanum large, its posterodorsal margin 
hidden by the parotid, % the eye diameter; parotid long, ex- 
tending almost to the groin but much more prominent and dis- 
tinct on the supratympanic region ; a large oval, inner metacarpal 
tubercle; subarticular tubercles large; fingers free, with a slight 
but thick lateral edge ; first finger shorter than second, which is 
shorter than fourth ; disks large, about % the size of the tym- 
panum ; hind limbs feeble for the body size ; no tarsal fold ; 



148 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

subarticular tubercles of toes large, rounded ; a slight web be- 
tween third and fourth toe; other toes free, with a thick lateral 
edge ; first toe as long as the second, second shorter than fifth ; 
disks of the first three toes smaller than the other two ; heel of 
the adpressed hind limb extends to the shoulder ; upper surfaces 
studded with bony deposits. Male with a brown rugosity on the 
inner side of the first digit but no apparent external vocal sac. 

Color. Above, bright green (blue green in preserved speci- 
mens) ; posterior and anterior surfaces of thighs and inguinal 
region dirty white with purple reticulations or milky white, 
purple-edged spots ; lower eyelid marked with purple-edged, 
white spots. Ventral surfaces of the body and limbs white ; mar- 
gin between the green upper and the white ventral color marked 
by purple-edged spots or by a white, purple-edged line on the 
fore limbs and distal segments of the hind limb ; fingers white, 
except the last two, which might be particolored or with green 
spots ; first three toes white, immaculate or with spots ; outer 
two, green ; disks of fingers and toes blue green, sometimes with 
white spots; lower lip with white, purple-edged elongated spots; 
a pair of white, purple-edged spots on the sides of the anal open- 
ing and two others on the breast, at the base of the forelimbs. 

Measurements. Snout-vent $ 103, 9 112 ; head breadth $ 
35.5, 9 39 ; head length s 32, 9 33 ; femur $ 30, 9 31 ; tibia 
$ 36, 9 40. 

Habits. The pair of specimens reported here were collected at 
night, during amplexus, on the ground, about 20 feet from a 
shallow pool. The voice is a trumpet-like "trrrbp." 

Range. The Venezuelan Guavana. The Guianas to Peru. 

Remarks. The two Venezuelan specimens have much smaller 
disks than either the Dutch Guiana or Peruvian individuals with 
which they have been compared ; the first two toes seem to be also 
more reduced, the canthus more angular and the loreal region 
more vertical. There is also some difference in the markings be- 
tween the upper and lower colors in the body of the Peruvian 
animal (M.C.Z. 4766). 

Phyllomedusa burmeisteri trinitatis Mertens 

Phyllomedusa trinitatis Mertens, 1926, Senckenbergiana, 8: 14o: Port of 
Spain. 

3 (C.N.H.M. 29181-3) Urama. 
Description. Head broader than long ; snout subovoid, truncate 



RIVERO : SALIENTIA OF VENEZUELA 149 

as seen from above ; tongue oval, entire or nicked behind ; vo- 
merine odontoids in two short, slightly oblique and posteriorly 
convergent groups between the large, oval choanae ; eye diameter 
equal or scarcely greater than distance between eye and nostril ; 
interorbital space about iy 2 times broader than an upper eyelid; 
canthus well defined, slightly curved; loreal slightly oblique, 
concave ; tympanum y 2 to % the eye diameter ; parotid extending 
to the sacrum or farther back, but more prominent on the supra- 
tympanic region ; an oval, well-defined inner metacarpal tubercle ; 
subarticular tubercles large; fingers free; the first shorter than 
the second, which is shorter than the fourth ; largest disk about 
y 2 the size of the tympanum ; a flat, oval, inner and a small in- 
distinct outer metatarsal tubercle ; toes free, with a thick lateral 
edge ; first toe longer than second ; disks of first three toes smaller 
than the others; heel of the adpressed hind limb extends to the 
posterior corner of the eye. Skin above studded with bony de- 
posits on the posterior half. Male with a brown rugosity on the 
inner side of the first digit but no apparent external vocal sac. 

Color. Above green, the flanks with white spots ; ground color 
on the posterior part of the flanks purplish, with white spots; 
lower eyelids margined with white ; lower lip distinctly margined 
with white ; a transverse white line above the anus. Ventral 
surfaces of the throat, chest and limbs purplish, with a few white 
spots near the chest and thighs, a pair of white spots under the 
thighs on each side of the anus; limbs green above, yellowish or 
purplish beneath, the two colors separated on the posterior part 
of the humerus and tarsal segment by a definite purple-margined 
line ; on the anterior part of the hind limb and also on the pos- 
terior part of the thighs and tibiae the green color changes to 
purple before assuming the yellowish color of the underparts; 
a series of white spots appear in this purple area from groin to 
inner toes ; three inner toes white, sometimes with purplish spots, 
but only the disks of the two inner toes are white ; three outer 
toes and also part of the fourth white, but the disk of the fourth 
is colored. A number of white spots at the base of the forearms. 

Measurements. Snout-vent S 65, 9 72 ; head breadth $ 22, 
9 25 ; head length $ 20.5, 9 23 ; femur $ 30, 9 32 ; tibia $ 
30.5, 9 34. 

Localities. El Periquito (U.C.V. 90). 

Range. The Coastal Range. The species extends from Trinidad 
to Argentina. The race is apparently limited to Trinidad, N. 
Venezuela, the Guianas and perhaps N. Brasil. 



150 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Remarks. Phyllomedusa burmeisteri burmeisteri from southern 
Brasil has very well defined and relatively large, white spots on 
the thighs, and some specimens are beautifully spotted with white 
on the throat. The specimens available to me for comparison (3) 
are considerably smaller than the Venezuelan and Trinidad 
specimens. One of the latter is 83 mm. long and the anterior and 
posterior parts of the thighs are purple, with whitish, very ob- 
scure spots. 

Phyllomedusa hypocondrialis hypocondrialis (Daudin) 

Hyla hypocondrialis Daudin, 1803, Hist. Rainette: 29 (hypocliondrialis on 
pi. x, fig. 1 ) : Surinam. 

2 (U.P.R. 187-8) Pto. Ayacucho, vi.50. 

Description. Snout short, truncate, slightly longer than the 
eye diameter ; tongue narrow, entire and free behind ; vomerine 
odontoids absent ; eye diameter greater than distance between 
eye and nostril; interorbital space broader than an upper eyelid; 
canthus moderately angular, somewhat curved ; loreal almost 
straight, concave ; tympanum fairly distinct, its dorsoposterior 
margin hidden by the parotid ; parotids indistinct ; male with a 
brown rugosity in the inner side of the first digit ; fingers free, 
the first shorter than the second, which is shorter than the fourth, 
fourth extending to the disk of the third ; disks of the fingers 
about % the diameter of the tympanum ; toes free ; first toe much 
longer than second, fourth shorter than fifth ; disks of the second 
and third toe smaller than the others ; heel of the adpressed hind 
limb extends to the tympanum. Skin above smooth; underside 
of forearms granular. Male with a small external subgular vo- 
cal sac. 

Color. Above and on the sides of the head, bluish purple ; upper 
lip and flanks white, the latter with purplish, vertical bars; hu- 
merus white, with cross reticulations; forearm purplish; thighs 
white, with purplish crossbars and a longitudinal purplish brown 
stripe along its dorsal surface ; tibial and tarsal segments 
purplish ; concealed surfaces of the limbs with crossbars ; ventral 
surfaces yellowish white. 

Measurements. Snout-vent $ 3fi, 9 44 ; head breadth $ 12, 9 
13.1 ; head length $ 11, 9 12.1 ; femur $ 10, 9 15 ; tibia $ 13, 
9 16. 

Habits. Collected at night in the tall grass of an inundated 
area. The female specimen is filled with very large, yellow eggs. 



RIVERO : SALIENTIA OP VENEZUELA 151 

Range. The Venezuelan Guayana. British Guiana to N. Brasil. 
Phyllomedusa hypocondrialis azure a Cope is found in Paraguay 
and Argentina. It is easily distinguished from the northern form 
by its much smaller disks, which are not broader than the width 
of the fingers. Five Pernambuco specimens at the M.C.Z. seem 
to be intermediate in disk size. The race is not, however, recog- 
nized by Funkhouser (1957). 

Remarks. The spelling hypocondrialis has been preferred to 
hypochondrialis not only because Daudin used it in the text of 
the original description but also because he repeated it in another 
paper (Hist. Rept., 1803b, 8: 60). 

CENTROLENIDAE 

Key to the Species of Cochranella Reported from Venezuela 

I. Fingers practically free, the first shorter than second; above, purple 

or bluish buchleyi 

II. Fingers webbed, the first equal or slightly longer than second ; above, 
green (in life) or yellowish white. 

A. Tympanum present; melanophores large and abundant 

Species indeterminata 

B. Typanum absent; melanophores not distinct fleishmanni 

Cochranella buckleyi (Boulenger) 

Hylella buckleyi Boulenger, 1882, Cat. Batr. Sal. Brit, Mus., ed. 2: 420, pi. 

xxv, fig. 5 : E. Ecuador. 
Centrolenella lucMeyi Noble, 1920, Bull. Amer. Mus. Nat. Hist., 42: 442. 

1 (M.C.Z. 2526) Merida. 
Description. Snout semicircular, almost as long as the eye 
diameter; tongue rounded, about x /2 free, nicked behind; vo- 
merine odontoids absent ; choanae small, round ; interorbital 
space broader than an upper eyelid ; canthus indistinct ; loreal 
oblique, concave ; tympanum indistinct ; a fold from elbow to 
disk of fourth finger ; a flat outer and a small, oval, inner meta- 
carpal tubercle ; subarticular tubercles flat ; fingers depressed, 
practically free, the first shorter than second ; disk of the fourth 
finger extending to the base of the disk of the third ; disks large, 
truncate, larger than the tympanum ; no humeral or tarsal 
folds ; a flat inner metatarsal tubercle ; toes taken in order from 
first to fifth exhibit the following phalanges free of web : 1%, 
l 1 /!* 1%) 3, 1% ; disks of the toes smaller than those of the fingers ; 
heel of the adpressed hind limb extends to the nostril. Skin 
above, smooth. 



152 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Color. Above, bluish purple ; a broad bluish purple band above 
the arm and another narrow one along the dorsal surface of the 
thigh, tibia and tarsus; remaining parts of the limbs yellowish. 
Ventral surfaces yellowish. 

Measurements. 9 Snout-vent 32.8; head breadth 11; head 
length 9; femur 16; tibia 17. 

Range. The Venezuelan Andes and probably the Coastal 
Range. Eastern face of the Andes of Ecuador through Colombia 
to Venezuela. 

Remarks. The specimen described here had been previously 
studied and dissected by Noble, who erroneously refers to this 
specimen as collected in Ecuador by Rosenberg (Noble, 1920: 
442). 

The genus Centrolenella, proposed by Noble (1920: 441) was 
found untenable by Dunn (1931: 399). Taylor (1951) on the 
basis of absence of vomerine teeth has proposed the name Coeh- 
ranella for most of the species that were included under the name 
Centrolenella (the genotype was found to be a Centrolene). 

COCHRANELLA Sp. 

1 (M.C.Z. 28569) Temiche, Mt. Marahuaca, 4050 ft., v. 50. 

Description. Snout semicircular, more or less as long as the 
eye diameter; tongue round, mostly adherent and indistinctly 
nicked behind ; vomerine odontoids absent ; choanae round ; inter- 
orbital space as broad as an upper eyelid; canthus very indis- 
tinct; loreal oblique, slightly concave; tympanum small and 
indistinct, about % the eye diameter ; no humeral spine ; first 
finger a little longer than second ; outer two fingers webbed, the 
web leaving about two free phalanges on the third and 1% on 
the fourth finger; a flat, inner metatarsal tubercle; toes taken 
in order from first to fifth exhibit the following phalanges free 
of web : 2, 1^4, 1, 2*4 to 2%, 1 ; heel of the adpressed hind limb 
extends to the nostril. Skin above, smooth. 

Color. Above, green with yellow mottles (in life) or yellowish 
white and with a relatively great profusion of melanophores 
(alcohol after formalin) ; upper eyelid golden. Below, whitish. 

Measurements. 9 1 Snout-vent 20 ; head breadth 7 ; head length 
6.5; femur 11; tibia 11. 

Remarks. This form is very similar to Cochranella fleishmanni 
from which it is distinguished by the presence of a tympanum 
and by the great profusion of melanophores on the dorsal surface 
of body and limbs. 



BIVERO: SALIENTIA OF VENEZUELA 153 

The animal was about to be named when, due to some unfortu- 
nate circumstance, the alcohol in the jar evaporated and the 
specimen dried out considerably. It was slightly restored with 
sodium triphosphate, but not sufficiently to bring out its original 
characters. Although there is a photograph of it, I prefer not 
to describe it as a species. 

COCHRANELLA FLEISHMANNI (Boettger) 

Uylclla fleishmanni Boettger, 18!)3, Ber. Senckenb. Naturf. Ges. : 251: San 
Jose, Costa Eica. 

2 (U.S.N.M. 117523-4) Los Canales. 

5 (U.S.N.M. 128789-93) R. Chacaito, viii, ix.38-39. 

3 (U.S.N.M. 128796-8) Los Canales. 

1 (U.S.N.M. 128800) Colonia Tovar, i.39. 

1 (U.S.N.M. 128801) Camino de Galipan, R. Cotiza, iii.39. 

Description. Snout semicircular, equal to the eye diameter; 
tongue rounded, % free and slightly nicked behind; vomerine 
odontoids absent ; choanae small and round ; interorbital space 
slightly broader than an upper eyelid; eanthus absent or very 
indistinct ; loreal oblique, not concave ; tympanum hidden ; no 
humeral spine ; first finger more or less equal to second ; only two 
outer fingers webbed, the web leaving two free phalanges on the 
third and iy 2 to 1% on the fourth finger; fingers flat, disks 
truncate ; a small inner metatarsal tubercle ; toes taken in order 
from first to fifth exhibit the following phalanges free of web : 
iy 2 , 1, \ x /-2i 2%> 1%; heel of the adpressed hind limb extends to 
between eye and nostril. Skin above, slightly shagreened. 

Color. Yellowish white throughout; no apparent chromato- 
phores ; eyelids white. 

Measurements. Snout-vent 25 ; head breadth 9 ; head length 
7.5 ; femur 13.5 ; tibia 13. 

Range. The Coastal Range and probably the lower Andean 
slopes. Central America to Venezuela. Dr. Dunn informed me 
that the species has been collected at the eastern base of the Andes 
of Colombia. 

Remarks. In some specimens the tongue is entire and in one 
the heel reaches the tip of the snout. 

DENDROBATIDAE 

Key to the Genera of Dendrobatidae Reported from Venezuela 
I. Upper jaw toothed. 

A. Toes webbed; ground color generally black or gray . Prostherap-ia 



154 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

B. Toes free or with a rudiment of web; ground color generally brown 

or tan Phyllobates 

II. Upper jaw toothless. 

A. Toes webless Dendrobates 

Key to the Species of Prostherapis Reported from Venezuela 
I. Toes y% webbed or more (not more than two free phalanges on the fifth 
and third and 1% on the second toe). 

A. Toes not fully webbed. 

1. A black band across the chest collaris 

2. No black band across the chest shrevei 

B. Toes fully webbed, with the exception of the fourth; above spotted 
with white ; no dark band across the chest dunni 

II. Toes not more than % webbed. 

A. A black band across the chest. 1 

1. Lower surface of dorsolateral band ending anteriorly at the 
corner of the eye; orbito-tympanic space white or whitish; loreal 
region usually with dark scribblings; chest band broad and 
well defined t. trinitatis 

2. Lower surface of dorsolateral band ending anteriorly at or near 
the lower margin of the eye; orbitotympanic space black; loreal 
region usually of uniform white color; chest band usually nar- 
row and ill defined ( 6 $ ) neblina 

B. No black band across the chest. 

1. Snout short and rounded; first finger shorter than second; most 

or all of the ventral surfaces dark, with white dots 

alboguttatus 

2. Snout truncate; first finger shorter than second; above, greenish 
silvery with an irregular vertebral and two lateral bands ; ventral 
surfaces light with obscure darker spots . trinitatis mandelorum 

Prostherapis collaris (Boulenger) 

Phijllobates trinitatis Boulenger, 1903, Ann. Mag. Nat. Hist., (7) 11: 482. 
Hylixalus collaris Boulenger, 1912, Ann. Mag. Nat. Hist., (8) 10: 190: 

Merida, 5,200 ft. and Rio Albarregas, 11,300 ft. 
Hyloxalus collaris Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20 : 40. 

3 (M.C.Z. 3887, 10723-4) Rio Albarregas, Merida. 
8 (U.M.M.Z. 58909) Rio Albarregas, Merida. 

1 (U.M.M.Z. 51267) Rio Albarregas, nr. Merida. 

2 (C.N.H.M. 3558-9) Rio Albarregas, Merida. 
1 (U.S.N.M. 118678) Merida. 

4 (A.M.N.H. 10512-15) Rio Albarregas, nr. Merida. 

3 (A.M.N.H. 10690, 10695-6) Merida. 

i May be absent in occasional specimens of P. t. trinitatis or P. neblina. 



RIVERO : SALIENTIA OF VENEZUELA 155 

Description. Snout subovoid, not truncate ; tongue oval or 
spatulate, free, entire or slightly emarginate behind ; eye diameter 
greater than distance between eye and nostril but a little shorter 
than the snout ; interorbital space broader than an upper eyelid ; 
canthus rounded ; loreal almost straight, slightly concave, if at 
all; tympanum indistinct or hidden, about y 2 the eye diameter; 
two flat, not distinct, metacarpal tubercles; fingers free, the first 
shorter than second ; an oblique, distinct tarsal fold ; two small 
metatarsal tubercles ; toes taken in order from first to fifth ex- 
hibit the following phalanges free of web : iy 2 , iy 2 , 2, 3y 2 to 3%, 
2 ; the web extends to the disks as distinct lateral fringes ; heel 
of the adpressed hind limb extends to the eye. Skin above and 
below, smooth. 

Color. Above and on the sides of the head and body grayish 
brown ; loreal region probably lighter than rest of the head ; 
flanks sometimes witli white spots in and near the groins. Ventral 
surfaces yellowish white, with some infuscation on the limbs and 
a broad brown band across the chest ; posterior part of the thighs 
obscurely marbled with light. 

Measurements. Snout-vent £ 27, 9 32 ; head breadth S 10, 
9 12 ; head length <5 9, 9 10.5 ; femur S 12, 9 15 ; tibia & 
12.5, 9 15. 

Range. The subtropical zone and above in the Merida Andes. 

Remarks. In 1940 Hellmich "corrected" Boulenger's descrip- 
tion of Prostherapis coUaris on the basis of differences found be- 
tween his 656 specimens and the original description. All his 
specimens were from the Coastal Range where P. collaris does not 
exist, an indication that what he had were P. trinitatis trinitatis. 

Prostherapis shrevei sp. n. 

Type. Museum of Comparative Zoology No. 28567, a $ from 
Mt. Marahuaca, 5000-6000 ft. Coll. J. A. Rivero, May 1950. 

Diagnosis. A Prostherapis with truncate snout, tympanum 
present, first finger shorter than second, a short tarsal fold, toes 
exhibiting the following phalanges free of web: 1, 1, 1%, 3%, 
1% ; brown dorsal color, having one interorbital, one or two 
scapular, two sacral and one preanal dark spots and crossbarred 
limbs. 

Description. Snout truncate ; tongue narrow, indistinctly 
nicked behind ; eye diameter greater than distance between eye 
and nostril, almost as long as the snout ; interorbital space broader 



156 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

than an upper eyelid ; canthus angular ; loreal vertical, not con- 
cave ; tympanum not very distinct, y 2 the eye diameter, dorsally 
hidden by a flat skin fold ; a flat, round, outer and a smaller, 
oval, inner metacarpal tubercle ; first finger shorter than second ; 
subarticular tubercles oval, not very prominent ; disks smaller 
than the tympanum ; a short but distinct fold at the distal half 
of the tarsus ; metatarsal tubercles small, the outer round, the 
inner elongated and more prominent ; toes taken in order from 
first to fifth exhibit the following phalanges free of web : l 1 /^, 
1^4, 2V4, 3V4, 1% ; web extended to the disks as distinct lateral 
fringes; heel of the adpressed hind limb extends to the anterior 
corner of the eye. Skin above, smooth, somewhat rugose behind 
the eyes and on the flanks. Belly finely granular under a lens. 

Color. Above, dark brown with an interorbital spot, another 
dark spot on the scapular region, two others in front of the 
sacrum and a single one behind these ; a number of white, radiat- 
ing lines from lower eyelid to upper lip ; one of these extends 
posteriorly to the shoulder; flanks very dark brown, with white 
spots that sometimes fuse with the ventral color ; a line of white 
dots from groin to about middle of the flank ; limbs crossbarred, 
the bars being oblique and very distinct on the thighs ; fingers and 
toes barred brown and white. Below, pure white. 

Measurements. S Snout -vent 21.5; head breadth 8; head 
length 8 ; femur 10.5 ; tibia 11. 

Habits. Collected in the proximity of streams or on stones 
within streams. 

Remarks. Except for specimen No. U.P.R. 232, the paratypes 
are not in very good condition and U.P.R. 234 and 235 are 
juveniles. 

U.P.R. 140, Mt, Marahuaca, 5000-6000 ft, Coll. J. A. Rivero, 
May 1950. Snout-vent 30.5; head breadth 7; head length 7; 
femur 10; tibia 11. Very similar to type in all respects but the 
snout is shorter and blunter, probably because it has received a 
blow from the front. Toes exhibiting the following phalanges 
free of web : iy 2 , 1 V>, 2, 3y 2 , 1%. 

U.P.R. 232, 9, Mt. Marahuaca, 5000-6000 ft. Coll. J. A. 
Rivero, May 1950. Snout-vent 36 ; head breadth 9 ; head length 
9 ; femur 12.5 ; tibia 12. Toes exhibiting the following phalanges 
free of web: iy 2 , IV?, 2Vo, 3, 1%. Almost solid brown above, the 
scapular and sacral spots, the loreal bars and the crossbars of the 
limbs being very obscure. Throat, chest and anterior part of the 
belly with peculiar chocolate spotting or speckling. 



RIVERO : SALIENTIA OP VENEZUELA 157 

U.P.R. 233, $ ?, Mt. Marahuaca, 5000-6000 ft. Coll. J. A. 
Rivero, May 1950. Snout-vent 20 ; head breadth 7.5 ; head length 
7 ; femur 11 ; tibia 11.5. Toes exhibiting the following phalanges 
free of web: iy 2 , IV2, 2, 3, 2y 2 . Dark brown above; spots ob- 
scure ; venter with brown mottling on the throat, chest and an- 
terior part of the belly. 

U.P.R, 234-5, Mt. Marahuaca, 5000-6000 ft. Juvenile speci- 
mens agreeing in all characters with the type. 

U.P.R. 236, Mt. Duida, about 1500-2000 ft. Coll. J. A. Rivero, 
May 1950. Snout-vent 17.5; head breadth 6.5; head length 6; 
femur 8 ; tibia 9. Except for its lighter loreal region and smaller 
size, this Mt. Duida specimen is almost identical with the type 
from Marahuaca. 

This species agrees best with Prostherapis bocagei (Espada) of 
which it is perhaps a race. It differs from this form in its slightly 
less webbed toes, more slender build, distinct crossbanding of the 
limbs and in its apparently persistent dark spotting above. Spot- 
ting may also occur in P. bocagei but it does not show the same 
pattern in this species as in P. shrevci. Apparently, Prostherapis 
shrevei represents another high altitude, Guayanan frog with 
Andean relationships. 

The species is named after Mr. Benjamin Shreve of the Mu- 
seum of Comparative Zoology. 

Prostherapis dunni sp. n. 

Type. Chicago Natural History Museum No. 35987, a 9 from 
above Caracas, D. F. Coll. Padre Cornelio Vogl, 1940. 

Diagnosis. A Prostherapis with a short rounded snout, equal 
to the eye diameter ; first finger shorter than second ; fully webbed 
toes ; dorsum spotted with white ; hind limbs banded with brown 
and yellow. 

Description. Snout short, subtriangular but with a blunt tip ; 
tongue small, oval, entire and free behind ; eye diameter as long 
as the snout ; interorbital space slightly broader than an upper 
eyelid ; canthus angular ; loreal vertical, not concave ; tympanum 
not very distinct, % the eye diameter ; two metacarpal tubercles ; 
first finger shorter than second ; disks relatively large, short : an 
indistinct, oblique tarsal fold ; two small metatarsal tubercles ; 
subarticular tubercles of fingers and toes small ; toes fully webbed 
with the exception of the fourth, which has two phalanges free ; 
heel of the adpressed hind limb extends to the anterior corner of 
the eye. Skin above and below, smooth. 



158 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Color. Above, brown with whitish and yellowish spots of ir- 
regular size and form; a dark brown band that begins at the 
posterior corner of the eye continues for a short distance pos- 
teriorly and then merges with rest of the body color; loreal 
region below the level of the nostrils light with dark scribblings ; 
a light line extends from anterior corner of the eye, around the 
tip of the snout, to the other eye ; another whitish, ill-defined line 
crosses the tympanic disk and extends to the shoulder; a rela- 
tively broad, irregularly margined line from groin toward shoul- 
der; posterior part of the thighs with an irregularly margined, 
longitudinal, yellow band; thiglis above, and rest of the limbs, 
with distinct brown crossbands that are separated by yellow 
areas. Throat yellowish, with some slight infuscation on the mar- 
gins of the jaw; belly white; hind limbs yellow. 

Measurements. 9 Snout-vent 24; head breadth 8 ; head length 
8; femur 13; tibia 12.5. 

Remarks. The 8 paratypes (C.N.H.M. 67379-67384) have the 
same data as the type and differ from it only in minor details. 
No. 67379 is a male, with the following measurements : snout- 
vent 20; head breadth 7.5; head length 8; femur 11; tibia 11. 
The largest specimen is No. 67384 with 25 mm. in snout-vent 
length and the smallest, No. 67380 with 18 mm. In some speci- 
mens there is more yellow and less brown on the thighs as 
compared witli the type while in others the dorsal spots are 
larger and the crossbars on the limbs better defined. 

This peculiar Prostherapis does not have any close relative in 
Venezuela. Prostherapis alboguttatus from the Mericla Andes 
has a more rounded snout, much less webbing and different ven- 
tral and dorsal coloration. 

Prostherapis trinitatis trinitatis (Garman) 

PhylloMtes trinitatis Garman, 1887, Bull. Essex Inst., 19: 13: Trinidad; 

Stejneger, 1902, Proe. U.S. Nat. Mus., 24: 179; Lutz, A., Mem. Inst. 

Osw. Cruz, 20: 40, 4fi ; Schmidt, K. P., 1932, Zool. Ser. Field Mus. Nat. 

Hist., 18: 160; Shreve, 1947, Bull. Mus. Comp. Zool., 99: 537; Aleutian, 

1952, Mem. Soc. Ciene. Nat. LaSalle, 12: 29, fig. 3. 
Prostherapis hrrminae Boettger, 1893, Ber. Senckenb. Naturf. Ges. : 37. 
Byloxalus coUaris Hellmich, 1940, Bol. Soc. Ven. Ciene. Nat., 6: 318; Zool. 

Anz., 131: 119. 

6 (M.C.Z. 26147-9 + 3 dupl.) Cerro Cosme. 
Description. Snout short, truncate ; tongue oval, free and 
slightly nicked behind; eye diameter longer than distance be- 
tween eye and nostril but shorter than the snout; interorbital 



RIVERO : SALIENTIA OF VENEZUELA 159 

space broader than an upper eyelid ; canthus angular ; loreal 
vertical, on occasions slightly concave ; tympanum i/> the eye 
diameter, its posterodorsal margin concealed under a skin fold ; 
metacarpal tubercles indistinct ; fingers free, their disks smaller 
than the tympanum ; first finger slightly shorter or equal to 
second; a short oblique fold at the distal half of the tarsus, 
metatarsal tubercles small ; two inner toes with a rudiment of 
web, the others with the following phalanges free of web : 2, 3, 
all; heel of the adpressed hind limb extends to the middle of 
the eye or between eye and nostril. Skin above, smooth or tuber- 
cular ; usually tubercular beyond the sacrum ; ventral surfaces 
smooth. 

Color. Above, dark grayish brown ; loreal region light with 
dark scribblings; humerus with an anterior dark longitudinal 
line ; a white lateral line from groin to about the middle of the 
flank ; other white dots are usually present on the flanks ; pos- 
terior part of the thighs obscurely marbled; hind limbs indis- 
tinctly crossbarred. Ventral surfaces Avhite with a distinct, but 
narrow, brown band across the chest. 

Measurements. Snout-vent $ 23, 9 26 ; head breadth <$ 8, 
9 9; head length $ 7.5, 9 9; femur «J 11, 9 11 ; tibia $ 11.."), 
9 12.5. 

Additional Localities. Baruta (Aleman, 1952) ; Camino de 
Galipan, R. Cotiza (U.S.N.M. 128782-3) ; Caracas (Hellmich, 
1940) ; Cerro Avila (U.C.V. 96-8, 2016-7) ; Colonia Tovar 
(U.S.N.M. 128781, 117540, 121135-43) ; Cordillera Padre Yogi 
(Hellmich, 1940); Cueva del Guacharo (U.C.V. 1) ; La Guaira 
(U.S.N.M. 27792 [= Stejneger, 1902]); Laguna de Valencia 
(Hellmich, 1940) ; Los Canales (U.S.N.M. 128900, 128777-80) ; 
Macuto (C.N.H.M. 5650) ; Maracay (Lutz, 1927) ; Pie de Avila 
(U.C.V. 109, 111) ; Pie del Cerro (U.S.N.M. 121144-5) ; Puerto 
Cabello (Boettger, 1893); Rancho Grande (M.C.Z. 26953-9; 
U.C.V. 23-27, 39-40, 63, 70-5, 84) ; Rio Chacaito (U.S.N.M. 
117538-9); Sn. Esteban (U.M.M.Z. 55687-9, 55546-9 [6] ) : Sn. 
Julian (U.S.N.M. 27808 [ = Stejneger, 1902]); Turgua (Ale- 
man, 1952) ; Mt. Turumiquire 17790 [10 = Schmidt, 1952]). 

Ranqe. The Coastal Range and the northeastern Falcon Re- 
gion. Trinidad. 

Remarks. The specimens described differ from other Vene- 
zuelan material and from the cotypes in their larger, stouter 
body, longer and more truncate snout, distinctly tubercular dor- 
sal surfaces (irrespective of sex) and brown coloration in which 
the dark lateral band is not generallv distinguishable. 



160 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

There seems to be considerable variation in Prosthcrapis t. 
trinitatis in general and this lias led to the inclusion within the 
species of several sibling forms that probably deserve specific 
distinction. Dr. Test has just detected one such ease in his 
P. neblina, but others may be found. 

M.C.Z. 26959 and less distinctly C.N.H.M. 5650 are all black 
above and below, the first only having a whitish area on the 
ventral aspect between the thighs and under the tibiae. In 
M.C.Z. 26953-8 the limbs are much lighter than the body color 
and the thighs are very distinctly crossbarred, the bars being 
separated by very light areas. The two Pie de Avila individuals 
have a pair of light bands extending backwards from behind 
the eyes. Although present in many P. t. trinitatis these bands 
are much more distinct and striking in the two Avila specimens 
as they continue inwards on the sides of the dorsum. 

U.C.V. 2016 and 2017 from Cerro Avila are of enormous size 
(32.5 and 30.5 mm.) and have white spots on the dorsal surfaces 
and a broader, less tapering snout. U.C.V. 96, 98 and 109 repre- 
sent this same form. 

Three males carrying tadpoles on their backs have been ex- 
amined. In every instance the adult frog is a much lighter gray 
than is normal for P. t. trinitatis. U.C.V. 74, also of very light 
grayish color, is probably a male that has recently shed its load 
of tadpoles. 

Prostiterapis trinitatis mandelorum (Schmidt) 

Phyllobates mandelorum Schmidt, K. P., 193:2, Zool. Ser. Field Mus. Nat. 
Hist., 18: 160: Mt. Turumiquire, 8,000 ft., Venezuela. 

2 (C.N.H.M. 17788-9, type + paratype) Mt. Turumiquire. 

8000 ft. 

Description (from original). Distinguished from the typical 
species by its dorsal and ventral coloration. The dorsum, includ- 
ing the top of the head is greenish silvery, sharply distinct from 
the chocolate colored sides; a dark irregular vertebral band 
of the same color as the sides begins with an expansion be- 
tween the eyes and extends nearly to the hind limbs; this band 
widens and encloses a light spot opposite the shoulders ; the dark 
color of the sides extends around the snout ; a silvery band on 
the upper jaw connects with the same color on the upper arm ; 
border of upper jaw dark; a light line extends forward from the 
frroin in the dark lateral band, which is not reached by tin 1 broad 
dark bars of the upper and posterior surfaces.- tibia barred. 



RIVERO: SALIENTIA OF VENEZUELA 161 

Ventral surfaces light with obscure darker spots; chest without 
trace of transverse dark band. 

Measurements. Snout-vent 26; head breadth 8.5; head length 
8.5 ; femur 12 ; tibia 11.5. 

Range. Mt. Turumiquire. 

Remarks. The type of P. /. mandclorum has a striking simi- 
larity to some of the cotypes of P. t. trinitatis. In some of the 
latter the chest band is very extensive and does not really form 
a band, while in others it is not very distinct. The similarity is, 
of course, to be expected as Mt. Turumiquire is relatively close 
to the type locality of P. t. trinitatis. 

Prostherapis neblina Test 

Prostherapis neblina Test, 1956, Oce. Pap. Mus. Zool. Univ. Mich., No. 577: 
2 : Portachuelo Pass, Baiicho Grande. 

2 (U.M.M.Z. 103011, 103014) Portachuelo Pass, Rancho 
Grande, 900-1100 m., 51. 

Description. Snout short, truncate; tongue pyriform, y 2 free 
and nicked behind ; eye diameter as long as the snout ; interorbi- 
tal space as broad as an upper eyelid ; canthus angular ; loreal 
vertical ; tympanum 1/2 the eye diameter, its posterodorsal margin 
concealed under a skin fold ; a round, prominent palmar and a 
smaller, oval, inner metacarpal tubercle ; fingers free ; first finger 
equal to second when adpressed in the middle ; disks smaller than 
the tympanum ; a distinct, oblique tarsal fold ; metatarsal tu- 
bercles small but prominent and distinct ; subarticular tubercles 
rounded ; toes with a short basal web ; taken in order from first 
to fifth, with the following phalanges free of web : 2, 2, 3, practi- 
cally all, practically all ; the web continues to the disks as con- 
spicuous lateral fringes; disks of toes approximately the same 
size as that of the tympanum; heel of the adpressed hind limb 
extends to between eye and nostril. Skin above, smooth or 
slightly shagreened, occasionally with sparse tubercles, or more 
closely set ones behind the sacral region. 

Color. Grayish brown, with two dark brown lateral bands 
from tip of the snout to inguinal region ; behind the eye, the 
lower margin of the band starts at about 7:00 o'clock (consider- 
ing the eyeball as a clock face) so that all the space between 
tympanum and eye is of a dark color and the tympanum falls 
within the dark area, only its lower margin touching the white 
upper lip ; lower section of the loreal region below the nostril and 
to approximately 4:00 o'clock on the right side of the eyeball, 



162 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

white, with a slight infuscation but no markings or reticulations 
of any sort ; the white color continues posteriorly below the eye 
and tympanum to the forearm ; a light colored line above the 
brown lateral band from upper eyelid to lumbar area; a white 
line extending anteriorly within the dark band from groin, to 
about halfway on the flanks ; an oblique, wedge-shaped bar on the 
anterior side of the forearm; limbs crossbanded, the bands dif- 
fusing and forming reticulations on the posterior aspect of the 
thighs. Below, white, immaculate, except for a narrow dark band 
across the anterior part of the chest. 

Measurements. Snout-vent 26.8 ; head length 9.3 ; head breadth 
9.2 ; femur 9.2 ; tibia 12.8. 

Habits. P. neblina is an inhabitant of humid slopes. Its voice 
consists of a series of short trills, while that of P. trinitatis, a frog 
of small streams is a series of clearly defined single notes. Upon 
being approached, P. trinitatis moves hurriedly away, while P. 
neblina permits close approximation and then moves away with 
short hops (Test, op. cit.). 

Range. Definitely known only from the type locality. U.C.V. 
21, 22 and 34, included under P. trinitatis trinitatis (see "Locali- 
ties" under that species) may be referable to this form. Un- 
fortunately, they were available for examination only before 
P. neblina was described. 

Remarks. According to Test, the variation in this species is 
small, although males tend to have narrower chest bands (some- 
times scarcely indicated) and more infuscation of the light areas. 
The dorsal surfaces may be uniform blackish or with faintly in- 
dicated mottles, and the lateral whitish line may be of variable 
forward extent. In some specimens the lateral black area is 
interrupted by whitish mottling and the dark bars of the limbs 
may be equally broken by mottling or freckling. 

Living P. neblina can be easily distinguished from P. trinitatis 
trinitatis, by the yellow ventral surfaces of the legs and to a 
lesser extent of the body, and by the uniformly dark brown back 
margined by the tan (whitish in alcohol) dorsolateral stripes 
(Test, op cit.). Alcoholic specimens are much more difficult to 
distinguish. In P. neblina the tympanum and the space between 
tympanum and eye are enclosed within the black area of the 
lateral band, while in P. t. trinitatis, the lower parts of tympanum 
and orbito-tympanic space are white. P. neblina is also said to 
have a less constant and conspicuous chest bar, more uniformly 



RIVERO : SALIENTIA OF VENEZUELA 163 

colored (white) upper lip, more uniform dorsal color, more dis- 
tinct dorsolateral stripes, a larger size, narrower head and more 
prominently banded limbs. However, many good P. t. trinitatis 
have very distinctly banded limbs. 

It is possible that some of Hellmich's P. trinitatis material 
(see synonymy of that species) may have included P. neblina, 
although the ventral surfaces are described as gray, with throat, 
anterior chest and sides of the belly dark blue gray. Lutz, 1927, 
describes the females of his specimens of P. trinitatis as having 
orange ventral surfaces, an indication that he may have had the 
two species. 

Prostherapis alboguttatus (Boulenger) 

Pliyllobates alboguttatus Boulenger, 1903, Ann. Mag. Nat. Hist., (7) 11: 
482: Merida, 1600 m.; Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 40, 47. 
5 (U.M.M.Z. 58904) Merida. 

1 (U.M.M.Z. 51266) Merida. 

2 (C.N.H.M. 3661-2) Merida. 

3 (A.M.N.H. 10551-53) La Culata, 3,000 m. 

8 (A.M.N.H. 644, 646-8, 649-51, 3137) Merida. 

Description. Snout short, rounded; nostrils more or less equi- 
distant between eye and tip of the snout ; tongue oval, free, entire 
or faintly nicked behind ; interorbital space broader than an 
upper eyelid ; canthus rounded or bluntly angular ; loreal almost 
vertical, not concave ; tympanum indistinct, y 2 to % the eye 
diameter; a large rounded outer and a small oval, inner meta- 
carpal tubercle ; subarticular tubercles rounded, not prominent ; 
fingers free, the first extending to the disk of the second ; larger 
disks smaller than the tympanum ; two small metatarsal tu- 
bercles ; first four toes with a short basal web that extends as 
lateral fringes to the disks, outer toe practically free ; heel of 
the adpressed hind limb extends to the eye. Skin above and 
below, smooth. 

Color. Above, brownish gray with obscure darker markings 
and two broad, dark brown, lateral bands that extend from the 
tip of snout to inguinal region ; above these there is apparently 
a light line from behind the eyelids backwards ; flanks clotted 
with white ; hindlimbs spotted or crossbanded ; throat, breast and 
anterior part of the belly brown, dotted with white ; posterior 
part of the belly and thighs generally white. 

Measurements. Snout-vent $ 25, 9 26.5; head breadth $ 
9.5, 9 10; head length $ 8, 9 9 ; femur $ 11, 9 11.5; tibia 
$ 11, 9 11.5. 



164 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

Range. The subtropical zone and above in the Merida Andes. 

Remarks. In some specimens the brown ventral color is re- 
stricted to the pectoral region and a transverse bar similar to 
the one that occurs in P. trinitatis and P. collaris is formed. 
Apparently this is pure coincidence and does not indicate close 
relationship between these forms. 

On some of the specimens all the ventral surfaces are dark 
brown, distinctly dotted with white. On the dorsal surface, how- 
ever, the spots described by Boulenger were not detected in any 
specimen. 

Prostherapis alboguttatus can be easily distinguished from P. 
trinitatis by its rounded, not truncate snout, nostril equidistant 
between eye and tip of the snout (in some specimens they are 
slightly closer to the eye), and different coloration. 

Key to the Species of Phyllobates Reported from Venezuela 

I. Third and fifth toes equal, no tarsal tubercle or fold ; tympanum dis- 
tinct bromelicola 

II. Third and fifth toes subequal; a tarsal fold; tympanum indistinct, al- 
most hidden; usually with a chain of diamond shaped markings above 
brvnnevs 

Phyllobates bromelicola Test 

Figure 12 

Phyllobates bromelicola Test, 1956, Occ. Pap. Mus. Zool. Univ. Mich., No. 
577: 6: Pico Periquito, Eaneho Grande, 1375 m. 
1 (U.M.M.Z. 113029, paratype) Pico Periquito, Rancho 

Grande, 1310 m. 
Description. Snout short, truncate ; tongue pyriform, half free, 
nicked behind ; eye diameter greater than distance between eye 
and nostril but slightly shorter than snout; interorbital space 
little broader than upper eyelid; canthus angular; loreal verti- 
cal; tympanum distinct, y 2 the eye diameter; two metacarpal 
tubercles, the outer divided and more extensive than the inner ; 
subarticular tubercles indistinct, except in the first finger, where 
the only tubercle is rounded and slightly prominent ; first finger 
considerably shorter than second, which is slightly shorter than 
fourth ; larger disks smaller than the tympanum ; two small meta- 
tarsal tubercles; toes free, their disks as large as those of the 
fingers; last toe feeble, shorter than third, and having the 
smallest disk of all ; heel of the adpressed hind limb extends to 
the anterior corner of the eye. Skin above, smooth. Below, 
feebly granular on the throat, belly and thighs. 



RIVERO : SALIENTIA OF VENEZUELA 



165 



Color. Above, light yellowish brown or tan with considerable 
darker dusting (contracted melanophores) ; a dark brown canthal 
streak beginning at the tip of the snout and continuing pos- 
teriorly to the lumbar region; on the flanks the stripe diffuses 
ventrally and near the groin it narrows to a broad line that 
extends almost to the tip of the urostyle ; a narrow white line 
above the brown band, from tip of snout, through margin of 




Fig. 12. Phyllobates bromelicola Test, Paratype U.M.M.Z. 113029. 



upper eyelid, to end of the brown band ; loreal region below the 
lateral band and upper lip immaculate; two distinct, short, 
oblique bars on the proximal end of the thigh, the distal one 
extending anteriorly to the anterior aspect ; another short bar on 
the middle of the tibia and other dusky, ill-defined ones on the 
ventral aspect of the tarsus and foot. Below, whitish, immacu- 
late. 

Measurements. $ , snout-vent 14.7 ; head length 5 ; head 
breadth 4.2 ; femur 7 ; tibia 7. 

Habits. As its name implies, P. bromelicola lives and ap- 
parently breeds in bromeliads, and was never found by its author 
outside of the "tree pines." 

Range. Only known from the upper slopes of Pico Periquito. 

Remarks. In life, the color of the species is described as fol- 
lows : ' ' ground color of the dorsum is olive, bordered by a distinct 
white hairline; lateral band black; bars on thighs and shank 
black; tips of digits black; underparts bright yellow through- 
out ..." (Test, op. cit.). 



166 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Superficially, P. bromelicola looks a lot like P. nubicola fiotator 
Dunn, but in the latter species the first finger is longer than the 
second; the white dorsolateral line enters into and divides the 
lateral dark band at its posterior end, and the thighs do not have 
oblique bars on the upper surfaces. The enlarged third finger 
of the male P. nubicola has not been described for P. bromelicola. 

Phyllobates brunneus (Cope) 

Prostherapis brunneus Cope, 1887, Proc. Amer. Philos. Soc, 24: 54: Matto 

Grosso. 
Prostherapis trinitatis Boettger, 1896, Ber. Senckenb. Naturf. Ges., 1895- 
6 : LIV. 

3 (U.M.M.Z. 113885 :AB4464, 4497, 4842) nr. La Cumbre, 
Maracay-Turiano Rd. 

1 (U.M.M.Z. 55548) Boqueron, Yaracuy. 

Description. Snout truncate; tongue narrow, free, entire or 
slightly nicked behind; eye diameter slightly greater than dis- 
tance between eye and nostril but shorter than the snout ; inter- 
orbital space slightly greater than an upper eyelid; canthus 
angular; loreal vertical; tympanum indistinct, almost totally 
hidden, only the anteroventral margin showing, about y 2 the eye 
diameter; metacarpal tubercles small but distinct; subarticular 
tubercles rounded, distinct ; first finger longer than second when 
adpressed in the middle, second longer than fourth; a short, 
oblique tarsal fold; metatarsal tubercles small, the inner elon- 
gated, the outer conical ; a short basal web between toes 2 and 3 
and 3 and 4 ; disks of toes larger than those of the fingers ; heel 
of the adpressed hind limb extends to the eye. Skin above, 
smooth, with scattered tubercles at the posterior end, slightly 
rugose on the flanks. Below, smooth. 

Color. Above, light yellowish brown or tan with a central, 
darker longitudinal band that widens and narrows a number 
of times and sometimes breaks into dots or scribblings at the 
sides; a broad, dark brown lateral band from tip of snout to 
ffroin ; above this a whitish narrow line from behind the upper 
eyelid to groin; upper lip white; a dusky wedge-shaped bar at 
the anterior aspect of the forearm ; no distinct white line on the 
flanks, within the brown lateral band ; limbs cross-barred. Below, 
whitish, immaculate. 

Measurements. Snout-vent $ 18, 9 18.5 ; head length <J 6, 
9 6.5 ; head breadth $ 6, 9 6 ; femur $ 8.2, 9 8.2 ; tibia $ 8, 
9 8.5. 



RIVERO : SALIENTIA OP VENEZUELA 167 

Habits. A ground species inhabiting humid slopes (Test, 
1956:6). 

Additional Localities. Mt. Marahuaca, 5000 ft. (U.P.R. 229-31, 
250) ; La Culebra, 1000 ft. (U.P.R. 251) ; Alto Orinoco (Boett- 
ger, 1896) ; Rio Guainia, Alto Orinoco (Senckenb. Mus. 7285 = 
Boettger, 1896). 

Range. The Coastal Range and the Venezuelan Guayana. N. 
Colombia and British Guiana to Rio de Janeiro. 

Remarks. The tympanum in the Venezuelan examples is in- 
distinct and almost hidden, but not totally concealed as described 
for the type specimen. The first finger is also described as equal 
to second and that is the way it looks if it is moved towards the 
second, but if both are adpressed in the middle of the two, the 
first appears longer. It also appears longer when the fingers are 
naturally separated. 

The Coastal Range Phyllobates brunneus agrees with the de- 
scription of Phyllobates marchesianus Melin in practically every 
respect. The dorsal longitudinal band of brunneus is not de- 
scribed for marchesianus, which is said to be brownish above, 
with indistinct black spots, but on the other hand, they both 
agree in having the first finger longer than the second (see 
above), a similar basic color, and in lacking a white line from 
groin to middle of flank. The two forms may prove to be con- 
specific. 

U.P.R. 229-31, 250, 251 were originally referred to P. mar- 
chesianus and then to P. brunneus when no good characters were 
found to differentiate them from this species, except perhaps the 
longer first finger and the presence of an indistinct tympanum. 
As other specimens with typical brunneus coloration have a tym- 
panum and a longer first finger, it was tentatively concluded 
that Cope's statement that the first finger is equal to the second 
in the type of brunneus may have resulted from his adpressing 
the first finger to the second in the position of the latter, instead 
of having them meet in the middle of the two. Regarding the 
tympanum, U.P.R. 251 has it definitely concealed while all the 
other specimens have it somewhat exposed. 

In most of the U.P.R. specimens the "dorsal chain of dia- 
monds" is not evident, and there is no whitish dorsolateral line 
in any. The latter may be the result of having the specimens 
for a few weeks in formalin after they were collected. However, 
they majr not have had the line at any time. Their toe disks are 
also bigger than those from the Coastal Range material. 



168 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

Melin, 1941, expressed doubt in allocating his slightly webbed 
marcliesianus to a genus, and ended up by calling it Phyllobates? 
marcliesianus. Unless different habits and habitats justify the 
separation, the division into the genera Prostherapis and Phyl- 
lobates should be reconsidered in the light of the more recently 
described forms. The division on the basis of presence or absence 
of web seems now untenable. 

Dendrobates leucomelas Steindachner 

Dendrobates leucomelas Steindachner, 1864, Batrachol. Mittheil. : pi. xiii, 

figs, la-d: Colombia. 
Dendrobates tinctorius Boettger, 1896, Ber. Senckenb. Naturf. Ges. : LIV. 
7 (U.P.R. 198-204) La Culebra, 1000 ft,, iv.50. 

1 (U.P.R. 205) Base Mt. Duida, 2000 ft. approx., vi.50. 

2 (U.P.R. 206-7) Tapara, iv.50. 

Description. Snout truncate ; tongue narrow, entire and free ; 
eye diameter equal to distance between eye and nostril; inter- 
orbital space flat, broader than an upper eyelid ; canthus angular 
but not sharp ; loreal slightly inclined inwards, not concave ; 
tympanum moderate, its posterior margin somewhat hidden un- 
der a flat, indistinct fold that runs to the shoulder, about % the 
eye diameter ; a large central metacarpal tubercle ; subarticular 
tubercles round, moderate; fingers free, the first shorter than 
second ; disks large, truncate and with converging margins ; disk 
of the first toe distinctly smaller than the others ; a short, oblique 
tarsal fold may be present or absent; two moderate metatarsal 
tubercles ; toes free, their disks much smaller than those of fingers ; 
heel of the adpressed hind limb extends to the tympanum. Skin 
above, smooth. Beneath, smooth on the throat and breast, slightly 
rugose on the belly and thighs. 

Color. Above, bright yellow (in life), usually with a broad, 
black transverse band across the scapular region and another 
across the sacral region ; the yellow spaces between the bands may 
have narrower bands or rounded or elongated black spots ; limbs 
yellow, spotted with black; a black transverse band across the 
distal end of the tibia seems to be constant; posterior part of 
the thighs black, the color extending over the anus to the distal 
coccygeal region. Ventral surfaces solid black, sometimes with a 
lateral intrusion of yellow from above or with an occasional yel- 
low spot. 

Measurements. Snout -vent $ 31.5, 9 37 ; head breadth $ 10, 

2 11; head length $ 9, 9 10; femur S 14.5, 2 15.5; tibia 

$ 13.5, 9 15. 



RIVERO: SALIENTIA OF VENEZUELA 169 

Habits. All the frogs were collected in rather shady places 
on the forest floor, on moist stones or on the mossy, water-drip- 
ping trunk of forest trees. The species is said to be common in 
all the Territorio Amazonas and someone informed me that he 
once saw a specimen inside a church in Ciudad Bolivar. 

The Maquiritare Indians consider this Dendrobates sacred as 
one of them enters into the history of their good female deity 
Cawishowa. This did not prevent the Indians from capturing 
the frog or from using them to test the effectiveness of curare. 
If the animal dies, the drug is supposed to be sufficiently strong 
to kill warm-blooded prey. 

The local name in Territorio Amazonas is "sapito minero" 
(miner's toad). 

Additional Localities. Auyantepui (A.M.N.H. 46044-7, 46051) ; 
Yapacana (U.S.N.M. 83937-41) ; Upper Orinoco (Boettger, 
1896). 

Range. The Venezuelan Guayana. Southeastern Colombia, 
British Guiana and N. Brasil. 

Remarks. No difference was found between the Venezuelan 
specimens from different localities. 

ATELOPODIDAE 

Key to the Genera of Atelopodidae Reported from Venezuela 

I. Hand and foot adapted for grasping; first toe longer than second 

Oreophrynella 

II. Hand and foot not adapted for grasping; first toe shorter than second 
Atelopus 

Key to the Species of Atelopus Reported from Venezuela 

I. Third linger shorter than distance between eye and tip of the snout ; 

yellow or brownish; Merida Andes oxyrhynchus 

II. Third finger longer than distance between eye and tip of the snout ; 
Coastal Range. 

A. Above, with a labyrinth of brown markings on a yellow or green 
background c. cruciger 

B. Above, uniformly yellow or brownish c vogli 

Atelopus oxyrhynchus Boulenger 

Atelopus oxyrhynchus Boulenger, 1903, Ann. Mag. Nat. Hist., (7) 12: 554: 
Rio Albarregas, Culata, Sierra Nevada de Merida, 10,000-11,000 ft.; 
Nieden, 1926, Das Tierreich. Anura II: 79; Lutz, A., 1927, Mem. Inst. 
Osw. Cruz, 20: 38, 41. 



170 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

2 (M.C.Z. 3804-5) Merida. 

Description. Snout flat, pointed, projecting considerably be- 
yond the mouth ; nostrils situated a little closer to the tip of the 
snout than to the eye ; tongue narrow, entire and free ; eye diame- 
ter slightly greater than distance between eye and nostril ; inter- 
orbital space flat, much broader than an upper eyelid ; canthus 
sharply angular, curved ; loreal vertical, concave ; tympanum 
absent ; a faint, central, metacarpal tubercle ; fingers free or with 
a short rudimentary web when viewed against a light ; first 
finger very short and thick ; subarticular tubercles almost im- 
perceptible ; tips of fingers slightly swollen ; a faint outer meta- 
tarsal tubercle ; toes about V2 webbed ; subarticular tubercles of 
toes absent or very faint ; heel of the adpressed hind limb extends 
to the shoulder or to the position of the tympanum. Skin above 
and on the sides of the body studded with small tubercles and 
large warts, especially on the posterior half of the body and on 
the limbs ; temporal area and base of the forelimbs with small 
spiny tubercles ; a ridge that commences as a bony ridge at the 
posterior corner of the eye extends posteriorly as a lateral row 
of glandules. Belly granular and rugose. Breeding male with 
longer and slightly enlarged arms and a brown rugosity on the 
inner side of the first digit. 

Color. Lemon or brownish yellow with greenish upper eyelids ; 
a brown streak that originates on the nostril extends posteriorly, 
passing the margin of the upper eyelids and marking the glan- 
dules of the lateral row. Small brown spots are indicated on the 
sides of the head of the male specimen. Boulenger describes a 
blotch of vermilion or orange red on the belly of some of his 
specimens, but in the material available the ventral surfaces are 
uniformly yellow. 

Measurements. Snout-vent 6 40, 9 48 ; head breadth $ 11, 
9 12.5 ; head length <$ 13, 9 14; femur $ 17, 9 20; tibia $ 16, 
9 17. 

Additional Localities. La Carbonera, 7000 ft. (U.M.M.Z. 57403 
[22] ; U.S.N.M. 118676) ; La Culata (A.M.N.H. 10593-6) ; Merida 
(U.M.M.Z. 58902 [5], 51268; C.N.H.M. 3625); Rio Mucujun 
(C.N.H.M. 5646) ; Venezuela, 3000-4000 m. (Nieden, 1926 [cited 
from Boulenger, 1903 ? J ) . 

Range. The subtropical zone and above in the Merida Andes. 

Remarks. U.M.M.Z. 58902 is a male with its natural color or 
most of it still existing. The ground color is brownish yellow 
and the larger dorsal tubercles are marked with brown ; the 



RIVERO : SALIENTIA OF VENEZUELA 171 

lateral band is well defined at the canthus, but behind the eye 
it narrows and only borders the lateral row of warts; there is 
also a brownish drab on the temporal area and behind it the 
tubercles are also marked with brown. It is possible that the 
lateral band was broader in this region and faded out, leaving 
only the tubercles marked with brown. There are also some brown 
spots on the throat, chest and limbs. In U.M.M.Z. 58902 the 
brown lateral line is more continuous and better defined than in 
other specimens. 

Irrespective of sex, there seems to be some variation in the 
form of the snout, which is very long and with a pointed tip in 
some specimens while in others the tip is rounder and shorter. 
The skin is sometimes smooth on the anterior half while in some 
individuals distinct tuberculatum extends to the head. 

Atelopus oxyrhynchus agrees with Atelopus c. cruciger in hav- 
ing a lateral row of warts, a brown, lateral band, spinulosity on 
the temporal area, tuberculation on the dorsal surface and a 
somewhat similar aspect. It differs from Atelopus cruciger in its 
much larger size, longer and more pointed snout, much shorter 
fingers and different coloration. 

Atelopus planispina Espada appears to have a snout similar to 
A. oxyrhynchus but its coloration apparently resembles that of 
A. c. cruciger and it differs from A. oxyrhynchus in some im- 
portant characters. 

Atelopus cruciger cruciger (Lichtenstein and Martens) 

Phrynidium cruciger "urn Lichtenstein and Martens, 1856, Nomencl. Rept. 
Amphib. Mus. Berol. : 41 : Veragoa. 

Phryniscus cruciger Giinther, 1858, Cat. Batr. Sal. Brit. Mus.: 44, pi. iii, 
fig. B.; Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 154. 

Phryniscus bibronii Giinther, 1858, Cat. Batr. Sal. Brit. Mus.: 137: Boulen- 
ger, 1882, Cat. Batr. Sal. Brit. Mus., ed. 2: 155: Boettger, 1893, Ber. 
Senckenb. Naturf . Ges. : 39. 

Atelopus spumarius Lutz, A., 1927, Mem. Inst. Osw. Cruz, 20: 38, 41, pi. 
xiii, figs. 27, 28. 

Atelopus cruciger Nieden (part), 1926, Das Tierreich. Anura II: 84; Lutz, 
A., 1927, Mem. Inst. Osw. Cruz, 20: 38; Miiller, L., 1934, Zool. Anz., 
108: 151; Eohl, 1949, Fauna Deser. de Ven., ed. 2: 399, fig. 179. 

Atelopus spumarius Aleman, 1952, Mem. Soc. Cienc. Nat. La Salle, 12: 28, 
fig. 2. 

2 (M.C.Z. 8657-8) Quebrada La Chapa, Nirgua. 
5 (U.M.M.Z. 55556) Quebrada La Chapa, Nirgua. 



172 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

9 (M.C.Z. 8659-61 ; U.M.M.Z. 55557) Sn. Esteban. 

6 (M.C.Z. 17744-9) Cordillera Ocumare. 

1 (U.M.M.Z. 68789) Campo Alto, nr. Maracay. 

Description. Snout flat, subtriangular, obtusely pointed and 
projecting slightly beyond the mouth ; nostrils closer to the tip 
of the snout than to the eye ; tongue narrow, entire and free ; 
eye diameter equal to distance between eye and nostril ; canthus 
straight, sharply angular ; loreal vertical, slightly concave ; in- 
terorbital space flat, broader than an upper eyelid; a faint, 
round palmar tubercle ; fingers practically free, except the first 
which has a short, basal web ; first finger short and stout ; sub- 
articular tubercles of fingers very indistinct, of toes more dis- 
tinct ; fingers and toes with swollen tips ; a flat, rounded, outer 
and a very indistinct inner metatarsal tubercle ; first toe very 
short ; toes V2 to % webbed ; heel of the adpressed hind limb 
extends to the anterior corner of the eye. Skin above, studded 
with minute granules or tubercles and flat warts on the posterior 
third ; a glandular row of warts from eye to groin. Below, 
areolate and rugose. 

Color. Above, yellowish or green with a labyrinth of irregular, 
usually confluent, dark brown reticulations ; one of the markings 
sometimes forms a large X at the nape ; a brown lateral stripe 
from tip of the snout to groin ; ventral surfaces of the limbs and 
flanks, below the lateral stripe, broadly marbled with brown; 
belly and throat yellowish white. 

Measurements. Snout-vent $ 33.5, 9 32 ; head breadth S 9, 
9 8 ; head length $ 8, 9 9 ; femur 6 15, 9 15 ; tibia 6 16, 
9 15.5. 

Habits. This species occurs in great numbers in the proximity 
of streams and sometimes on plants up to a height of 1.5 m. 
(Roze). 

Additional Localities. Curanna (Cumana, Gunther, 1858) ; 
Curupao (U.S.N.M. 117546-50) ; Curucuruma (U.C.V. 41) ; R. 
Chacaito (U.S.N.M. 128877-81) ; El Hatillo (Aleman, 1952) ; 
Guamitas (U.C.V. 9) ; Los Canales (U.S.N.M. 128853-76, 117542- 
5) ; Between Maracay and Ocumare (U.S.N.M. 81137) ; Puerto 
Cabello (Giinther, 1858; Boettger, 1893); Rancho Grande 
(U.S.N.M. 97196-97200; U.C.V. 54, 59-62, 64, 73, 80-1) ; Turgua 
(Aleman, 19.12); Venezuela (Nieden, 1926). 

Range. The Coastal Range from Yaracuy to Sucre. 

Remarks. There seem to be two color varieties of this form, 
one of them as shown by Gunther (1858, pi. Ill) is characterized 



RIVERO : SALIENTIA OP VENEZUELA 173 

by very close and fine reticulations on the dorsum, usually an 
X-shaped marking at the nape and another chevron-shaped mark- 
ing at the sacral region. The other, as shown by Lutz (1927, pi. 
XIII) has a more broken pattern, there is no well defined X- 
shaped marking, and the basic color seems to be darker. Only 
the latter variety is found in Rancho Grande and in Nirgua and 
San Esteban but both are found in the M.C.Z. series from Cor- 
dillera Ocumare. The specimens from Guamita and Curucuruma 
have the fine pattern. The first is a large example, 42 mm. in 
length. 

U.C.V. 64 is peculiar in being very dark, almost black above, 
and in having distinct broad marbling on the lower flanks and 
ventral portion of the limbs, and scattered dark spots on the 
belly and throat. 

Although Phrynidium crucigerum was described from Veragua, 
Panama, the alleged cotypes (Berlin Mus. 3387(3)) represent 
the Venezuelan species, which the original description also fits. 
It thus appears that the A. crucigcr coining from Venezuela suf- 
fered an exchange of locality data with the Eleutherodactylus 
gollmeri coming from Panama, with the result that Eleuthero- 
dactylus gollmeri was for a long time considered to have Caracas 
as its type locality, while A. crucigcr was thought to be Pana- 
manian. In this paper, the range of Atelopus crucigcr is limited 
to the Coastal Range of Venezuela. 



Atelopus cruciger vogli Midler 

Atelopus crucigcr vogli Miillcr, L., 1934, Zool. Anz., 108: 151: Las Pefias, 
nr. Maracay. 

1 (U.M.M.Z. 92431, paratype) Las Penas, nr. Maracay. 
3 (M.C.Z. 20923-5, paratypes) Las Penas, nr. Maracay. 
Description. Atelopus c. vogli differs from the typical form in 
its more slender habit, slightly more projecting snout ; longer and 
more slender inner finger; less webbed toes (% to i/o) and dif- 
ferent coloration. The dorsal and ventral color is yellow, but, 
according to Midler, specimens with brownish dorsums and 
faintly indicated markings on the anterior part of the back are 
occasionally found. There is a lateral stripe along the flanks as in 
A. c. cruciger. 

Measurements. Snout-vent 6 28, 9 36.5 ; head breadth S 7, 
9 9; head length $ 8, 9 10; femur «J 12, 9 16; tibia $ 13, 
9 16.3. 



174 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Additional Localities. Las Pehas, Hacienda La Trinidad 
(U.S.N.M. 113265-6, 113268-74) ; near Maracay (U.S.N.M. 
107328-33 paratypes). 

Remarks. The only information I have been able to obtain 
about the habitat of this form was sent to me by Mr. J. Roze of 
Caracas, who obtained it through the courtesy of Padre Vogl, the 
collector. According to this note, A. c. vogli is found in a very 
inaccessible place to which it is necessary to climb by holding to 
vines and branches. 

My examination of this form includes a series of 46 specimens, 
all of which are uniform yellow above. Atelopns cruciger cruci- 
ger is also of a plain color dorsally up to snout-vent lengths of 
15 mm. (U.S.N.M. 128862-5) ; at 16.5 mm. snout-vent length 
(U.S.N.M. 128861) all the adult markings have appeared. 

Oreophrynella quelchii quelchii (Boulenger) 

Oreophryne Quelchii Boulenger, 1895, Ann. Mag. Nat. Hist., (6) 15: 521: 

Summit Mt. Roraima, bet. Brit. Guiana and Venezuela, 8,500 ft. 
Oreophrynella Quelchii Boulenger, 1895, Ann. Mag. Nat. Hist., (6) 16: 125; 

1900, Trans. Linn. Soc. London, (2) 8: 55, pi. v, fig. 3. 
Oreophrynella quelchii Nieden, 1926, Das Tierreieli. Anura II: 75. 

3 (M.C.Z. 3500-2, ctps.) Mt. Roraima, 8500 ft. 

2 (M.C.Z. 24072-3) Roraima Plateau, 8000 ft. 

1 (C.N.H.M. 43671) Mt. Roraima, 45. 

1 (U.S.N.M. 118230) Roraima. 
Description. Snout short, rounded; tongne narrow, entire and 
free ; eye diameter almost as long as the snout ; interorbital space 
as broad as an upper eyelid; canthus short and curved, well 
defined ; loreal vertical or almost so ; tympanum absent ; two flat, 
rounded, metacarpal tubercles ; palm and fingers with small, flat 
tubercles; fingers very short, distally swollen, the first shorter 
than second, which is slightly shorter than fourth; metatarsal 
tubercles very small; sole with small, flat tubercles; first three 
toes joined at the base and opposed to the other two, first longer 
than second ; heel of the adpressed hind limb extends to the 
shoulder or near the eye. Skin above, including the upper eyelids 
and limbs densely tubercular ; flanks tubercular ; occasionally 
present, a longitudinal fold or ridge along the posterior part of 
the thigh. Belly granular. 

Color. Above, very dark brown, almost black; loreal region 
usually smooth and shiny. Below, spotted or broadly marbled 
with yellow. 



RIVERO : SALIENTIA OF VENEZUELA 175 

Measurements. Snout-vent $ 20, 9 21.5; head breadth $ 7, 
9 7.5; head length $ 5.5; 9 6; femur $ 8.5, 9 8 ; tibia $ 7.5, 
9 8. 

Localities. The Museum of Zoology of the University of Michi- 
gan has 38 specimens from the Roraima Plateau. M.C.Z. bas two 
from Roraima, Brasil. 

Range. The Venezuelan, British Guianan and Brasilian sum- 
mit of Mt. Roraima. 

Oreophrynella quelchii macconnelli Boulenger 

Oreophrynella Macconnelli Boulenger, 1900, Trans. Linn. Soc. London, (2) 

8: 55, pi. v, fig. 1 : Base of Mt. Roraima, 3500 ft. 
Oreophrynella macconnelli Nieclen, 1926, Das Tierreich. Anura II: 76. 

No material examined. 

Original Description. Distinguished from Oreophrynella 
quelchii quelchii by its more prominent snout, projecting much 
beyond the mouth and by the greater expansion of the digits 
which end in distinct truncate disks. Interorbital space broader 
than an upper eyelid ; first toe much longer than second, as long 
as fourth; no distinct metatarsal or subarticular tubercles; first 
and second toes appear to be opposable to the fourtb and fifth, 
both fascicles being bound by the thick integument, the third 
toe, which is the shortest, being free between them ; tarso-meta- 
tarsal articulation of the adpressed hind limb extends to the eye. 
Skin above, covered with small, smooth, feebly prominent warts. 
Lower parts with flat granules. 

Color. Above, brown with lighter marblings, and a series of 
small yellowish spots, forming a line on each side of the back 
from eye to groin, continued obliquely across the upper surface 
of the femur; upper lip yellowish, with two dark brown bars 
below the eye ; lower parts whitish. 

Measurements. Snout-vent 22. 

Range. Only known from the type specimen. 

Remarks. I treat this frog as a race of Oreophrynella quelchii 
on account of its close structural similarity to the nominate form 
(as expressed in the description), and also because one seems to 
represent the other at the higher elevations of Roraima and be- 
cause no other member of the genus is known to occur in South 
America. 

See remarks under Otophryne roousta. 



176 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

KANIDAE 
Rana palmipes Spix 

Rana palmipes Spix, 1824, Spec. Nov. Test. Kan., 29, pi. v, fig. 1: Amazon; 
Boulenger, 1920, Proc. Amer. Acad. Arts Sci. (9)55: 473; Lutz, A., 
1927, Mem. Inst. Osw. Cruz, SO: 40, 47, pi. ix, figs. 7-9; Schmidt, K. P., 
1932, Zool. Ser. Field Mus. Nat. Hist., 18: 161 ; Aleman, 19.12, Mem. Soc. 
Cienc. Nat. LaSalle, 12: 28, fig. 4. 
Ranula Gollmeri Peters, 1859, Monatsb. Akad. Wissensch. Berlin: 402. 
Rana affinis Peters, 1859, Monatsb. Akad. Wissensch. Berlin: 403; Cope, 
1866, Proc. Acad. Nat. Sci. Philadelphia: 130. 

1 (U.P.R. 226) Tapara, iv.50. 

Description. Snout subovoid ; tongue deeply notched behind; 
vomerine odontoids in two short or rounded, oblique and pos- 
teriorly convergent groups between the choanae ; eye diameter 
as long as distance between eye and nostril ; interorbital space 
as broad as an upper eyelid ; canthus distinct ; loreal little slop- 
ing, deeply concave; tympanum large, % the eye diameter; sub- 
articular tubercles round, distinct; first finger slightly longer 
than second; tips of fingers slightly dilated; one elongate, not 
very distinct, inner metatarsal tubercle ; subarticular tubercles 
of toes elongated, distinct ; toes fully webbed with the exception 
of the fourth which has about 2 free phalanges; tips of toes 
dilated into small disks ; heel of the adpressed hind limb extends 
to a little beyond the anterior corner of the eye. Skin above and 
on the hind limbs tubercular; a pair of dorsolateral folds from 
posterior corner of the eye to groins. 

Color. Above, olive, turning to green on the head (in life) ; 
anterior and posterior aspects of the thighs black and light 
marbled ; limbs crossbarred ; below, yellowish and black marbled. 

Measurements. 9 Snout-vent 50; head breadth 18.3; head 
length 21 ; femur 22 : tibia 24. 

Habits. The only specimen was caught very early in the morn- 
ing, half submerged in the waters of the Cunucunuma River. The 
species does not seem to be common in the region. 

Additional Localities. Arabopo (U.M.M.Z. 85207-8) ; Baruta 
(Aleman, 1952) ; Bejuma (U.M.M.Z. 55586) ; Bermudez St. 
(C.N.H.M. 17772-4); Caracas (Peters, 1859); Caserio Silva; 
Oumanacoa (Schmidt, 1932) ; El Hatillo (Aleman, 1952) ; Rio 
Lamona (U.M.M.Z. 55576) ; Maracay (Lutz, 1927) ; Nirgua, La- 
guna de los Coronjos (U.M.M.Z. 55577-8) ; San Feliz, Estacion 
Tachira (U.M.M.Z. 55579) ; Tributary of Rio Uribante (U.M.M.Z. 
85111); Venezuela (U.M.M.Z. 85176; Cope, 1866). 



RIVERO : SALIENTIA OF VENEZUELA 177 

Range. The base of the Merida Andes, the Coastal Range, the 
Venezuelan Guayana and probably all the other physiographical 
provinces of Venezuela. Central America and Trinidad to Peru 
and Matto Grosso. Rana palmipes hoffmanni Miiller, 1924, has 
been described from Costa Rica and Rana palmipes rionapensis 
Andersson, 1945, from Ecuador. Rana brevipalmata rhoadsi 
Fowler, 1913, is considered a synonym. 

Remarks. A female specimen with a snout-vent length of 122 
mm. has been examined. The heel may extend to between eye and 
nostril or to the nostril, while the tympanum may be as much 
as % the eye diameter. A few specimens are salmon-colored 
below. 

Specimen C.N.H.M. 17773 is probably a male, with a tym- 
panum 3 4 the eye diameter and an infuscated throat. It has the 
following measurements : snout-vent 66 ; head breadth 25 ; head 
length 28; femur 31; tibia 36. 

Cope's "Ranula sp. nov., 0. Salvin ; Vera Paz, Venezuela," 
which was listed immediately following his Ranula affinis, was 
presumably Rana palmipes. 

MICROHYLIDAE 

Key to the Genera of Venezuelan Microhylidae 

I. Clavicle and procoracoid reduced, the former not extending to the 

scapula and confined to the mesial portion of the procoracoid 

Elachistocleis 

II. Clavicle and procoracoid extending from midline of girdle to scapula 
Otophryne 

Elachistocleis ovalis (Schneider) 

Bana ovalis Schneider, 1799, Hist. Amphib. : 131 (type loc. not cited). 
7 (U.P.R. 219-225) Puerto Ayacucho, vi.50. 
Description. Head triangular; snout pointed, about 1% longer 
than the eye diameter, projecting considerably beyond the mouth ; 
tongue attached anteriorly and behind but loose in the middle; 
eye diameter shorter than distance between eye and nostril ; 
interorbital space more than twice as broad as an upper eyelid ; 
canthus absent ; loreal slightly oblique ; tympanum hidden ; three 
small, indistinct metacarpal tubercles : fingers short, the first 
shorter than second, which is very little shorter than fourth; 
one inner metatarsal tubercle, no outer ; toes free ; subarticular 
tubercles flattened ; heel of the adpressed hind limb does not 



178 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

reach the shoulder. Skin above, smooth ; a transverse fold behind 
the eyes; a flat oblique fold from posterior corner of eye to 
shoulder. Below, smooth. 

Color. Above, black with minute light points ; flanks and ven- 
tral surfaces gray, marbled or spotted with orange ; a large 
orange spot at the groin and proximal portion of the thigh; 
another broad, orange spot along the posterior aspect of the 
thigh and hidden portion of the tibia; smaller orange spots are 
present on their margins ; throat darkened in the male. 

Measurements. Snout-vent $ 30.5, 5 37 ; head breadth $ 8, 
$ 7.5 ; head length $ 6, 9 7 ; femur $ 11.5, $ 7 ; tibia $ 11, 
9 8. 

Additional Localities. Espino (U.C.V. 47) ; Palenque (U.S. 
N.M. 128847-9) ; Sta. Elena, 3500 ft., Bolivar (U.M.M.Z. 85143). 

Remarks. I find no difference between the Puerto Ayacucho 
specimens and two paratypes of Hypopachus pearsei Ruthven, 
from Fundacion, Colombia (M.C.Z. 3727, 6075). A large, 55 mm. 
specimen collected by Dr. P. J. Darlington in Rio Frio, Colom- 
bia, however, has a longer snout that is slightly greater than twice 
the eye diameter. The spotting on the flanks, groins and thighs 
of all these specimens is very similar to M.C.Z. 2037 from Suri- 
nam. U.M.M.Z. 85143, U.S.N.M. 128847-9 and U.C.V. 47 from 
Venezuela, as well as M.C.Z. 4089 from Trinidad, have a narrow, 
well-defined line on the posterior part of the thighs. The Trini- 
dad specimen appears to have very long legs, probably because 
of its desiccated condition, but groin and ventral spots are still 
obvious. U.C.V. 47 has A^entral spots, small spots under the 
femoral line and groin spots. U.M.M.Z. 85143 has groin spots, 
probably scattered spots underneath (very desiccated) and no 
spot surrounding the femoral line, while the U.S.N.M. specimens 
are very small and dry and the ventral coloration cannot be 
detected. I am inclined to believe that there is some variation in 
the form of the femoral line but that Elarhistocleis pearsei and 
E. ovalis are conspecific. 

The Museum of Comparative Zoology has 8 specimens from 
Buenavista, Bolivia, 6 of which fall into the description of E. 
oicolor while 2 have small yellow spots below, an inguinal spot, 
and on one of them, a broad femoral line. E. oicolor has a brown 
or orange belly that is surrounded by yellow on the throat, sides 
of the belly and lower flanks, thighs and tibiae. The upper lip 
also has a yellow margin that extends posteriorly to the shoulder. 
One of the specimens is 33 mm., another 32, while the others are 



RIVERO: SALIENTIA OF VENEZUELA 179 

smaller. In the same lot, a specimen (M.C.Z. 15686) with uni- 
formly yellow ventral surfaces appears to have small spots on 
the slightly darkened throat. In M.C.Z. 12862, from Bolivia, 
which should be E. ovalis as there are groin and ventral spots 
(small) below, the sides of the darkened throat ( $ ) are yellow 
and the femoral line narrow, reminiscent of the condition in 
E. bicolor. I believe these forms need further study; the fact 
that the two color variations exist in Bolivia should not be defi- 
nitely interpreted to mean that the two are full species as this is 
a region where intergradation of other forms is known to occur. 
Cei (1956) has already used the name E. ovalis bicolor for the 
Argentinian form. 

Measurements. $ Snout-vent 54 ; head breadth 19 ; head length 
18; femur 22.5; tibia 22. 

Additional Localities. Arabopo (U.M.M.Z. 85136, 85138). 

Range. Mt. Roraima region of Venezuela, British Guiana and 
probably Brasil. U.M.M.Z. 85139-40 are from the Kurupung R. 
in British Guiana. 

Remarks. The specimens were dry and stiff so that the limbs 
could not be adpressed to the sides. According to Boulenger the 
tarso-metatarsal articulation extends to the tympanum. 

Phelps (1938b) has demonstrated that, of the several expe- 
ditions that visited Mt. Roraima, only the second expedition by 
McConnell and Quelch could have collected in the small British 
Guianan segment of the summit plateau. Specimens from 3500 
ft. come, however, from Venezuelan territory. 

Otophryne robusta Boulenger 

Otophryne robusta Boulenger, 1900, Trans. Linn. Soc. London, 8: 55, pi. v, 
fig. 5: Mt. Roraima, 3500 ft., "British Guiana" (in error for Vene- 
zuela); Nieden, 1926, Das Tierreich. Anura II: 73; Parker, 1934. 
Monogr. Fa. Microhylidae (London) : 109. 

1 (U.M.M.Z. 85137) Arabopo. 
Description. Snout obliquely truncate ; tongue oval, entire and 
free behind ; two well defined, crenulated ridges in front of the 
pharynx ; eye diameter slightly longer than distance between 
eye and nostril but shorter than the snout; interorbital space 
much broader than an upper eyelid ; canthus angular ; loreal 
vertical, slightly concave ; tympanum disproportionately large, 
much larger than the eye, with which it is in contact ; a fine 
supratympanic fold from posterior corner of eye to shoulder 
and a lateral fold from this to inguinal region ; metacarpal tu- 
bercles indistinct ; subarticular tubercles absent or only faintly 



180 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

indicated ; fingers free, the first shorter than second which is 
shorter than fourth ; outer toe very short ; metatarsal tubercles 
absent; toes distally spatulate, about y 2 webbed, the web ex- 
tending to the tips as conspicuous lateral fringes. Skin above 
and below, smooth. 

Color. Above, very dark brown, obscurely spotted with darker ; 
groins and posterior aspect of the thighs spotted with yellowish 
or orange and black. Ventral surfaces of the belly and limbs 
dirty white, with scattered brown spots; throat and chest dark 
brown. 

LOCALITIES 

The following is a list of Venezuelan localities in which frogs 
have been collected, and a few others mentioned in the text which 
may be difficult to find. The descriptions are taken from the 
literature or from personal observation. The equivalents of 
abbreviations are given below : 

H. Temp. = Highest, average, median temperature (C). 

L. Temp. = Lowest, average, median temperature (C). 

Ann. Temp. = Average, annual, median temperature (C). 

H. Rain. = Highest, average, median rainfall (mm.). 

L. Rain. = Lowest, average, median rainfall (mm.). 

Ann. Rain. = Average, annual, median rainfall (mm.). 

1. Acarigua, Edo. Portuguesa — 9°33'27"N., 69°11'54"W. — 
Elev. 186 m. — II. Temp., Mar. (28.5°), L. Temp., Jul. 
(25.6°), Ann. Temp., 26.8° H. Rain., Jun. (287), L. 
Rain., Feb. (5), Ann. Rain. 1503. 

2. Acosta, Distrito, Edo. Falcon — In the northeastern part of 
Falcon, between Zamora and Silva. 

3. Albarregas, Rio, Edo. Merida — Rio Chama system, with 
headwaters in Sierra La Culata and mouth above Merida. 

4. Altagracia de Orituco, Edo. Guarico — 9°12'30"N., 
66°00'00"W. — Elev. 358 m. — H. Rain., Aug. (168), L. 
Rain., Feb. (1), Ann. Rain. 913. 

5. Amana, Rio, Edo. Monagas — Tributary of Rio Guanipa. 
with headwaters in the Coastal Range and mouth 60 km. 
east of Maturin. 

6. Anaben, Colombia - ■ 4°4'N., 67°44'W. approx. Elev. 
about 100 to 150 m. A small Colombian village at the angle 
formed by the Atabapo with the Inirida. Most frogs col- 
lected in tall grass some 200 yards from the river (Ata- 
bapo) edge or in a cacao planting nearby. 



RIVERO : SAEIENTIA OF VENEZUELA 181 

7. Antimano, D. F. — 10°28'N., 66°58'W. approx. — Elev. 
928 m. 

8. Apure, Eio — Large tributary of the Orinoco on the west- 
ern Llanos. 

9. Agiiita Waterfalls, Terr. Amaz. — Elev. 988 m. A camp 
on a 45° slope of Mt. Duida. Conditions, subtropical. 

10. Arabopo, Edo. Bolivar — 5°2'30"N., 62°42'30"W. approx. 
— Elev. 1,216 m. A village at the base of Cerro Roraima, 
on the valley of Rio Arabapo. The river begins with a 
cascade in the southwestern cliff of Roraima and joins the 
Cuquenan River some 20 miles to the south. The valley is 
broad, chiefly savanna covered and bordered on both sides 
by rolling hills but at about a mile north of the village the 
grassland terminates. The border of the Weitipu forest, 
stretching for some 10 miles to the northern slopes of Cerro 
Weitipu, lies y 2 hour walk to the southeast. The extensive 
forests of Arabopo, starting from the foothills of Roraima, 
curve around the northern end of the valley and around 
the ridge reaching southwest to Weitipu. In addition, forest 
growth occurs in deep valleys and along the margins of 
streams ( Chapman ) . 

11. Aragua de Barcelona, Edo. Anzoategui — 9°27'29"N., 
64°49'33"W. — Elev. 96 m. — H. Temp., Apr. (28°), L. 
Temp., Jan. (24.9°), Ann. Temp. 26.2° — H. Rain., Aug. 
(208), L. Rain., Feb. (2), Ann. Rain. 1,030. 

12. Aroa, Edo. Yaracuy — 10°26'N., 68°54'50"W. approx. — 
Elev. around 370 m. On the valley of Rio Aroa. This is 
dry and the native vegetation has been mostly destroyed. 
Some rocky "quebradas" occur near the town (William- 
son). 

13. Auyantepui, Cerro, Edo. Bolivar — 5°48'N., 62°33'W. — 
Elev. 2,400 m. One of the sandstone mesas of the Guayana 
Highlands. The southern slopes are mostly treeless to 2,000 
m. but otherwise the zones are usually well defined, the 
transitional slopes being narrow or absent (Tate). 

14. Avila, Cerro, D. F. — In the vicinity of Caracas. Frogs 
collected in the forest on the margins of streams between 
1,800-2,200 in. Above 1,500 m. the forest is subtropical 
(Roze). 

15. Barcelona, Edo. Anzoategui — 10°08'06"N., 64°41'05"W. — 
Elev. 10 m. 

16. Barinas, Edo. Barinas — 8°37'49"N., 70°12'10"W. — Elev. 
180 m. — H. Temp., Mar. (27.1°), L. Temp., Sept., Oct. 



182 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



(26.0°), Ann. Temp. 26.6° — II. Rain., Jun. (364), L. 
Rain., Jan. (11), Ann. Rain. 1,863. 




Fig. 13. Faunal regions in Venezuela. 



17. Barquisimeto, Edo. Lara - 10°03'57"N., 69°18'45"W. 
Elev. 566 m. — H. Temp., Nov. (24.9°), L. Temp., Jan., 
Jun. (23.6°), Ann. Temp. 24.2° — H Rain., Feb. (5), Ann. 
Rain. 519 Conditions, xerophilous. 

18. Barrancas, Edo. Monagas - - 8°42'30"N., 62°11'W. approx. 
— Elev. not more than 100 m. 

19. Baruta, Edo. Miranda — Between Caracas and El Hatillo. 
Elev. around 1,100 m. Conditions, subtropical. 

20. Bejuma, Edo. Carabobo — 10°10'00"N., 68°17'00"W. 
Elev. 662 m. — Ann. Temp. 23.9°, H. Rain., May (279), 
L. Rain., Feb. (2), Ann. Rain. 1,208. In a circular plain 
surrounded by high hills. Most of the valley of Rio Bejuma 
and the steep surrounding hills are generally under culti- 
vation. "Quebradas" in the hills are usually surrounded 
by luxuriant growth (Williamson). 



BIVERO: SALIENTIA OF VENEZUELA 183 

21. Berniudez State — Old Venezuelan state that included the 
present Anzoategui, Sucre and Monagas. 

22. Boca de Rio, Edo. Aragua — 10°17'47"N., 67°38'25"W. — 
Elev. 415 m. — H. Temp., Apr. (26.0°), L. Temp., Jan. 
(23.8°), Ann. Temp. 24.9° H. Rain., Aug. (161), L. 
Rain., Jan. (1), Ann. Rain. 821. 

23. Bonicito, Rio, Edo. Zulia — A small river in the Maracaibo 
District. 

24. Caicara, Edo. Bolivar — 7°38'N., 66°14'W. — Elev. around 
60 m. 

25. Caju, Caiio, Terr. Amaz. — - A swift running stream at the 
base of the vertical wall of Marahuaca. Conditions, upper 
tropical but vegetation is not thick and the trees not high. 
Collections from about 1,200 m. 

26. Calabozo, Edo. Guarico — 8°56'06"N., 67°25'00"W. — Elev. 
106 m. — H. Temp., Apr. (29.2°), L. Temp., Jul., Aug. 
(26.6°), Ann. Temp. 27.5° — H. Rain., Jun. (322), L. Rain., 
Jan. (1), Ann. Rain. 1,303. 

27. Campo Alto, Edo. Aragua — Apart from the fact that it is 
near Maracay, little has been learned about this locality. 

28. Caracas, D. F. (center of city) — 10°30'30"N., 66°54'50"W. 
— Elev. 920 m. — H. Temp, (at Observatorio Cagigal) May 
(21.5°), L. Temp., Jan. (18.8°), Ann. Temp. 20.4° — H. 
Rain., Oct. (139), L. Rain., Feb. (7), Ann. Rain. 897. 

29. Carapa, Edo. Anzoategui — 8°21'N., 63°16'W. approx. — 
Elev. not more than 100 m. 

30. Carayaca, D. F. — 10°32'N., 67°07'W. — Elev. 500 m. 
approx. Conditions, tropical to subtropical. Frogs collected 
in the surroundings of a small lake (Roze). 

31. Caripito, Edo. Monagas 10°07'11"N., 53°05'26"W. — 
Elev. 50 m. approx. — Ann. Temp. 26.1° — H. Rain., Jun. 
(305), L. Rain., Feb. (39), Ann. Rain. 2,182. 

32. Carora, Edo. Lara — 10°11'07"N., 70°15'31"W. — Elev. 
420 m. — H. Temp., Jun. (28.7°), L. Temp., Mar. (25.6°), 
Ann. Temp. 27.7° — H. Rain., Oct. (144), L. Rain., Feb. 
(11), Ann. Rain. 632. 

33. Casa de Julian, Terr. Amaz. — Two houses and an Indian 
plantation or "conuco" in the middle of the jungle between 
Tapara and Cano Chana. Frogs collected in the "conuco" 
at 609 m. or in the margins of a nearby stream at approxi- 
mately the same elevation. Conditions, tropical. 



184 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

34. Caserio Silva, Edo. Carabobo — A "posada" between Valen- 
cia and Bejuma. The place is situated on the margin of 
a small, clear stream (Rio La Mona) which is generally in 
the sun but in some places has a little forest remaining 
along it (Williamson). 

35. Casiquiare Canal, Terr. Amaz. — The famous branch of the 
Upper Orinoco that connects with the Amazon through Rio 
Negro. Conditions, mostly humid tropical. 

36. Casurua, Terr. Amaz. — An abandoned hut and "conuco" 
on the northern margin of Rio Cunucunuma, about 5 hours 
by motor boat from the mouth of the river. Soil sandy and 
vegetation not very luxuriant. Estimated elevation, about 
300 m. 

37. Chaeaito, Quebrada, D. F. — A stream at the eastern boun- 
dary of Distrito Federal. Elev. 960-1,000 m. 

38. Ghana or Chanane, Caho, Terr. Amaz. — A stream between 
Casa de Julian and Cerro Marahuaca. Conditions, humid 
tropical. Collections come from the surrounding forest at 
an elevation of 781 m. 

39. Chama, Rio, Edo. Merida — A river that originates in the 
vicinity of Paramo de Macuchies, runs between Sierra La 
Culata and Sierra Nevada de Merida and empties into one 
of the cienagas on the south of Lake Maracaibo. 

40. Chupadero, Chorro, Terr. Amaz. — A small stream that 
empties into the eastern side of Rio Orinoco at about 6 or 7 
hours by motor boat up the river from San Fernando de 
Atabapo. Heavy tropical vegetation on all sides. 

41. Ciudad Bolivar,' Edo. Bolivar — 8°08'52"N., 63°33'06"W. — 
Elev. 54 m. — H. Temp., Sept. (28.3°), L. Temp., Jan. 
(26.5°), Ann. Temp. 27.5° — H. Rain., Jul. (186) L. Rain., 
Mar. (8), Ann. Rain. 1,022. 

42. Cocollar, Edo. Sucre — 10°10'00"N., 63°47'00"W. - - Elev. 
835 m. H. Rain., Oct. (159), L. Rain., Feb. (17), Ann. 
Rain. 1,137. Almost arid in the dry season, the area pro- 
duces a coarse, low grass and usually low trees (Tate). 

43. Cocuhy, Rio — I have been unable to find this locality in 
Venezuela. Cucuy or Cocuy is a place in the southwestern 
corner of Venezuela and perhaps there is a nearby river of 
the same name. 

44. Colonia Tovar, Edo. Aragua — 10°24'38"N., 67°17'03"W. - 
Elev. 1,790 m. — H. Temp., May (16.3°), L. Temp., Jan. 
(14.1°), Ann. Temp. 15.4° H. Rain., Aug. (188), L. 
Rain., Feb. (14), Ann. Rain. 1,300. 



RIVERO : SAL1ENTIA OF VENEZUELA 185 

45. Coro, Edo. Falcon — 11°24'48"N., 69°40'42"W. — Elev. 21 
m. — H. Temp., Sept, (29.9°), L. Temp., Jan. (27.1°), Ann. 
Temp. 28.4° - - H. Rain., Nov. (100), L. Rain., Feb. (14), 
Ann. Rain. 442. 

46. Cosine, Cerro, Edo. Falcon — 10°53'N., 68°41'W. approx. 
— Elev. probably 250-300 m. 

47. Cotiza, Rio, Camino de Galipan, D. F. — Granja Cotiza is 
near Caracas at an elevation of 900 m. Conditions, sub- 
tropical (Chardon). 

48. Cua, Edo. Miranda — 10°09'42"N., 66°53'20"W. — Elev. 
241 m. — H. Rain., Jim. (218), L. Rain., Feb., Mar. (5), 
Ann. Rain. 959. 

49. Culata, Edo. Merida — See La Culata. 

50. Cumana, Edo. Sucre — 10°27'47"N., 64°10'37"W. — Elev. 
3 m. — H. Temp., Oct. (27.6°), L. Temp., Jan. (25.7°), 
Ann. Temp. 26.9° - - H. Rain., Aug. (64), L. Rain., Feb., 
Mar. (4), Ann. Rain. 375. 

51. Cumanacoa, Edo. Sucre 10°15'00"N., 63°55'10"W. — 
Elev. 240 m. — Ann. Temp. 24.7° approx. — H. Rain., Aug. 
(223), L. Rain., Feb. (17), Ann. Rain. 1,336. In the valley 
of Rio Manzanares. The floor of the hill encircled valley 
is level and apparently formed by sedimentation. Most of 
the area is under cultivation (Tate). 

52. Cumarebo, Edo. Falcon - 11°29'25"N., 69°21'26"W. 
Elev. 13 m. 

53. Cunucunuma, Rio, Terr. Amaz. — A tributary of the Ori- 
noco with its mouth at about 250 km. up the river from 
San Fernando de Atabapo. Except for an occasional small 
savanna or ' ' conuco, ' ' the margins of the Cunucunuma are 
heavily forested from its mouth to Tapara and probably 
farther up. 

54. Cuquenan, Cerro, Edo. Bolivar — The second largest of 
Mt. Roraima mountains. Elev. 2,584. 

55. Cuquenan, Valley, Edo. Bolivar — Valley of Rio Cuque- 
nan, an upper tributary of Rio Caroni, with headwaters in 
Mt. Roraima. 

56. Curucuruma, Edo. Aragua — Elev. 1,000 m. approx. Con- 
ditions, subtropical (Roze). 

57. Curupao, Edo. Miranda 10°30'00"N., 63°3818"\V. 
Elev. 420 m. — H. Rain., Jun. (155), L. Rain., Mar. (5), 
Ann. Rain. 1,082. At 1,120 m. H. Rain., Sept. (159), L. 
Rain., Mar. (18), Ann. Rain. 1,305. 



186 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

58. Distrito Federal — District in which the capital city of 
Venezuela is located. 

59. Duida, Cerro, Terr. Aniaz. — A sandstone cerro on the 
southern margin of Rio Cunucunuma. See Esmeralda, 

Agiiita Waterfalls, Carlo Pescado, and La Culebra. 

60. Egido, Edo. Miranda — 8°32'45"N., 71°15'17"W. — Elev. 
1,167 m. — Ann. Temp. 22.6° — H. Rain., Nov. (176), L. 
Rain., Jan. (23), Ann. Rain. 984. 

61. El Baul, Edo. Cojedes — 7°05'57"N., 68°17'01"W. — Elev. 
102 m. — H. Rain., Jun. (402), L. Rain., Feb., Mar. (0), 
Ann. Rain. 1,662. 

62. El Hatillo, Edo. Miranda — 10°27'N., 66°54"W. approx. 
Elev. around 1,100 m. 

63. El Junquito, D. F. — Elev. 1900-2100 m. Conditions, 
subtropical, except in the higher elevations where only grass 
and low shrubs occur. 

64. El Limon, D. F. — 10°28'N., 67°18'W. approx. — Elev. 
577 m. — Ann. Temp. 22.2° — H. Rain., Oct. (204), L. 
Rain., Feb. (33), Ann. Rain. 1,477. 

65. El Mene de Acosta, Edo. Falcon — 11°00'30"N., 68°31'- 
00"W. approx. Elev. not more than 100 m. 

66. El Mene de Buchivacoa, Edo. Falcon — 10°43'N., 71°00'W. 
approx. Elev. not more than 100 m. 

67. El Periquito, Cerro, Edo. Monagas — 10°13'N., 63°41'W. — 
Elev. 500 m. approx. Conditions, tropical to subtropical. 

68. El Sombrero, Edo. Guarico — 9°23'10"N., 67°03'17"W. — 
Elev. 160 m. — H. Rain., Jul. (252), L. Rain., Feb., Mar. 
(0), Ann. Rain. 1,205. 

69. El Valle, D. F. — 10°29'07"N., 66°50'52"W. — Elev. 885 
m. — H. Temp., May (21.7°), L. Temp., Jan. (19°), Ann. 
Temp. 20.7° - - H. Rain., May (117), L. Rain., Feb. (5), 
Ann. Rain. 797. 

70. Encontrados, Edo. Zulia — 9°04'30"N., 72°18'00"W. approx. 
In the humid, tropical plain southwest of Lake Maracaibo. 

71 . Escorial, Edo. Merida — 8°42'N., 72°02'W. approx. — Elev. 
2,500 m. and up. Mammals have been collected above timber 
line at 3,000 m. 

72. Esmeralda, Terr. Amaz. — An abandoned native hacienda 
at about 50 km. up the Orinoco from the mouth of Rio 
Cunucunuma. Located on a mostly flat, grassy savanna 
with occasional low, swampy areas. 

73. Espino, Edo. Guarico — 8°34'N., 66°01'W. — Elev. about 
200 m. — Specimens collected in gallery forest at the mar- 
gin of a stream (Roze). 



RIVERO : SALIENTIA OF VENEZUELA 187 

74. Ge, Rio, Edo. Zulia — Tributary of Rio Palmar, near Villa 
del Rosario, west side of Lake Maracaibo. 

75. Guachi, Rio, Edos. Merida and Zulia — The Rio Guachi 
arises in the northwestern slopes of the Andes and empties 
into the southern end of Lake Maracaibo, some 12 miles 
east of the mouth of Rio Chama (Osgood and Conover). 

76. Guamitas, Parque Naeional, Edo. Aragua — Elev. 700 m. 
approx. Conditions, tropical to subtropical. 

77. Hiquerote, Edo. Miranda — Collections come from the sur- 
roundings of the many rivulets and channels that connect 
with the sea in this area (Roze). 

78. Jacare, Terr. Amaz. — A small Indian village on the north- 
ern margin of Rio Cunucunuma between Casurua and La 
Culebra. Conditions, heavy tropical. 

79. Kukenam — See Cuquenan. 

80. Kunana, Edo. Zulia — Savanna on the southern side of Rio 
Negro, Perija Range. Elev. 1100 m. 

81. La Azulita, Edo. Merida — 8°42'03"N., 71°25'32"W. — 
Elev. 1,135 m. — H. Rain., Jun. (197), L. Rain., Jan. (42), 
Ann. Rain. 1,398. Village on west side of Rio Guachi. Some 
of the surrounding country is cleared for the production 
of sugar and coffee, but most of it is heavily forested. Con- 
ditions, humid tropical to subtropical (Osgood and Con- 
over). 

82. La Carbonera, Edo. Merida - 8°37'30"N., 71°21'00"\V. 
approx. — Elev. 2,128 m. An ''hacienda" at the north- 
eastern slopes of Paramo Tambor, near the headwaters of 
a western branch of Rio Guachi. Temperate forest and open 
cleared meadows prevail (Osgood and Conover). 

83. La Chapa, Quebrada, Edo. Carabobo — A stream near the 
town of Miranda. Collections come from the wooded ridges 
at 1,109 m. 

84. La Cruz Pruviera, Edo. Guarico — I have been unable to 
obtain any information about this locality. Specimens from 
this place were collected by Dr. Henri Pittier. 

85. La Culata, Edo. Merida — A sierra of the Merida Andes 
extending for 80 km. between Paramo de Cheque (3894 

m.) to La Cuchilla (2,475 m.). A paramo (440 m.) in the 
above mentioned sierra. 

86. La Culebra, Edo. Miranda — At about 30 km. in straight 
line from El Junquito but separated from this locality by 
two large and deep valleys. Although the approximate 



188 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

elevation is 1,800 m., the conditions are mostly tropical due 
to its location in the Internal Range where the trade winds 
are little felt ( Roze ) . 

87. La Culebra. Pico. Terr. Amaz. — An eastern peak of Mt. 
Duida rising from Savanna La Culebra at 334 m. and 
reaching an approx. elev. of 1398 m. Talus forest to 1000 
m. where the subtropical forest commences. At about 1340 
m. trees disappear and the terrain is characterized by loose 
rocks, among which a small, spiny pineapple grows 
abundantly. 

88. La Culebra, Savanna, Terr. Amaz. — Savanna at the base 
of Pico La Culebra. Elev. 334 m. 

89. La Fria, Edo. Tachira — 8°13'N., 72°19'W. approx. — 
Elev. 140 m. The forest to the south of the town consists 
of a heavy, mixed growth ; to the north the terrain is nearly 
flat with occasional, muddy and swampy spots, and to the 
east there is a sandy ' ' quebrada ' ' with a good flow of water 
(Williamson). 

90. La Guaira, D. P. — 10°30'48"N., 66°56'02"W. — Elev. 5 m. 
— H. Temp., Sept. (28.1°), L. Temp., Jan., Feb. (25.2°), 
Ann. Temp. 26.8° — H. Rain., Nov. (104), L. Rain., Apr. 
(17), Ann. Rain. 553. Conditions, arid and semiarid but 
mountains rise to 900 m. immediately behind the town and 
constant precipitation in them produces a number of 
streams that are followed by some forest growth along their 
course to the sea. 

91. Lagunillas, Edo. Zulia — 10°07'20"N., 71°15'30"W. — Elev. 
5 m. H. Temp., Oct. (28.8°), L. Temp., Jan. (27.5°), Ann. 
Temp. 28.3° — H. Rain., May (125), L. Rain., Jan. (7), 
Ann. Rain. 900. Village on the eastern shore of Lake Mara- 
caibo. Behind the village there is an extensive, shallow 
lagoon or cienaga filled with water plants and bordered by 
rushes and mangroves. The surrounding country is arid to 
semiarid (Osgood and Conover). 

92. La Mona, Rio, Edo. Carabobo — See Caserio Silva. 

93. La Pascua, Edo. Guarico - - 9°12'30"N., 66°01'00"W. ap- 
prox. — Elev. 195 m. 

94. Las Penas, Ravine, Edo. Aragua — Near Maracay. See re- 
marks under Atelopus cruciger vogli. 

95. Las Quiggas, Edo. Carabobo — A village on Rio San Este- 
ban and a stream that passes by (Williamson). 

96. La Trinidad, Hacienda, Edo. Aragua — 10°21'10"N, 
67°36'07"W. — Elev. 455 m. - - H. Rain., Aug. (185), L. 
Rain., Feb. (1), Ann. Rain. 984. 



RIVERO : SALIENTIA OF VENEZUELA 189 

97. La Trinidad, Edos. Sucre-Monagas — A coffee plantation 
in Mt. Turnmiquire at an elevation of 6000 ft. Conditions, 
subtropical. 

98. Los Canales, Planta Electrica de Naiguata - I have been 
unable to obtain information about this locality. 

99. Los Coronjos, Laguna, Edo. Yaracuy — A lagoon in Nir- 
gua. 

100. Los Teques, Edo. Miranda — 10°24'48"N., 67°02'02"W. — 
Elev. 1169 m.— H. Rain.. Jun. (161), L. Rain., Mar. (10), 
Ann. Rain. 1,126. 

101. Los Venados, D. F. — 10°32'28"N., 66°53'66"W. — Elev. 
1,524 m. — Ann. Temp. 9.2° — Ann. Rain. 716. 

102. Macuto, D. F. — 10°36'50"N., 66°53'00"W. approx. Three 
miles east of LaGuaira. It is said to present a most fertile 
and flourishing character (Robinson and Lyon). 

103. Machiques, Edo. Zulia — 10°03'18"N., 72°23'49"W. - - Elev. 
150 m. - - H. Rain., Oct. (241), L. Rain., Jan. (8), Ann. 
Rain. 1,519. 

104. Manamo, Cano, Delta Amacuro — Channel that forms the 
boundary between Monagas State and the Delta Amacuro 
Territory. 

105. Maracaibo, Edo. Zulia — 10°38'32"N., 71°36'26"W. - - Elev. 
6 m. H. Temp., Aug. (29.4°), L. Temp., Jan., Feb. 
(27.4°), Ann. Temp. 28.3° - - H. Rain., Oct. (149), L. Rain., 
Feb. (0), Ann. Rain. 573. Conditions, arid. 

106. Maracay, Edo. Aragua - - 10°15'17"N., 67°35'57"W. - - Elev. 
445 m. — H. Temp., Apr. (26.2°), L. Temp., Jan. (23.7°), 
Ann. Temp. 24.7° — II. Rain., Aug. (170), L. Rain., Jan. 
(1), Ann. Rain. 858. 

107. Marahuaca, Cerro, Terr. Amaz. --One of the Gnayanan 
Cerros, constituting with the Duida and Huachamacari, a 
distinct triangle at the western end of the Pacaraima Sys- 
tem. Conditions tropical to about 1231 m., where a sub- 
tropical pocket occurs on the northern side. Elev. 2581 m. 

108. Maturin. Edo. Monagas -- 9°44'30"N., 63°10'30"W. - - Elev. 
75 m. — H. Temp., Apr. (26.8°), L. Temp., Jan. (24.5°), 
Ann. Temp. 25.7° — H. Rain., Jun. (208), L. Rain., Mar. 
(18), Ann. Rain. 1,248. 

109. Merida. Edo. Merida — 8°35'56"N., 71°09'24"\V. - - Elev. 
1,623 m. — H. Temp.. Aug. (19.7°), L. Temp.. Jan. (18.2°). 
Ann. Temp. 19.2° — II. Rain.. Sept. (273), L. Rain.. Feb. 
(45), Ann. Rain. 1.816. Situated in a mesa between Rio 
Chama and Rio Albarregas. In the river valleys conditions 



190 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

are tropical or subtropical but mountains rise abruptly on 
either side and temperate forests are accessible within a 
few hours travel (Osgood and Conover). 

110. Money, Edo. Trujillo — 9°24'02"N., 70°36'21"W — Elev. 
340 m. — H. Rain., Aug. (162), L. Rain, Mar. (27), Ann. 
Rain. 970. 

111. Motatan, Edo. Trujillo - - 9°32'00"N, 70°24'00"W. - - Elev. 
296 m. — H. Rain., Oct, (206), L. Rain, Feb. (52), Ann. 
Rain. 1,504. 

112. Minapana, Quebrada, Edo. Yaracuy — A stream near 
Palma Sola. 

113. Mucujun, Rio, Edo. Merida — Rio Chama System, with 
headwaters in Sierra La Culata and mouth above Merida. 

114. Negro, Rio, Terr. Amaz. — River that connects the Amazon 
with the Orinoco through the Casiquiare Canal. 

115. Nirgua, Edo. Yaracuy - - 10°09'06"N, 68°34'13"AV. — Elev. 
798 m. — H. Rain, May (171), L. Rain, Feb. (1), Ann. 
Rain. 923. Conditions, similar to Bejuma but country is 
rougher and adjacent to the town more despoiled. Rio 
Borria runs at the foot of the plateau on which the town 
is built. Above the intake of the city water supply, the river 
flows through brush and small trees. A few miles north of 
town there is a quebrada in pastures, brush, and coffee and 
banana plantings; on top of the hill, a humid forest (Wil- 
liamson). 

116. Ocumare de la Costa, Edo. Aragua 10°27'30"N, 
67°46'07"W. approx. — Elev. 15 m. 

117. Ocumare del Tuy, Edo. Miranda 10°07'03"N, 
66°46'07"\V. — Elev. 210 m. — H. Rain.. Jul. (206), L. 
Rain, Feb. (15), Ann. Rain. 1,125. 

118. Orope, Edo. Tachira - - 8°22'30"N, 72°22'00" W. approx. — 
Elev. not more than 100 m. In the heart of the humid tropi- 
cal forest where its conditions are developed to the highest 
degree (Osgood and Conover). 

119. Oropito, Rio, Edo. Zulia — River that passes near La Fria 
in the southern part of Lahe Maracaibo. 

120. Palenque, Edo. Guarico -- 8°59'30"N, 66°58'00"\V. approx. 
— Elev. between 100 and 200 m. 

121. Palma Sola, Edo. Falcon 10°36'35"N, 68°33'35"W. - 
Elev. 40 m. approx. — Ann. Rain., 1,703. In a nearly flat, 
heavily wooded country. The Aroa River, which passes by, 
is swift and shallow in this region. 



RIVERO: SALIENTIA OF VENEZUELA 191 

122. Pariaguan, Edo. Anzoategui - - 8°06'00"N., 64°44'00"W. — 
Elev. 123 m. — H. Temp., Apr. (27.4°), L. Temp., Jan., 

Jun., Dec. (26.0°). Aim. Temp. (26.3°). H. Rain., Jul. 
(311), L. Rain., Feb. (0), Ann. Rain. 1,428. 

123. Parmana, Edo. Guarico - - 7°50'00"N., 65°50'00"W. approx. 

— Elev. 80 m. approx. Vegetation intermediate between 
that of gallery forests and llanos. During the rainy season, 
the surrounding area is flooded but Parmana remains above 
the waters (Roze). 

124. Paso del Diablo, Terr. Amaz. — 2°16'30"N., 66°33'00"W. 
approx. — Elev. 100-200 m. 

125. Pauji, Edo. Falcon - - 11°01'N., 68°38'W. approx. - - Elev. 
200-500 m. 

126. Paulo, Edo. Bolivar — Elev. 1,216 m. An Arecuna village, 
some seven miles to the southwest of the cliffs of Roraima, 
stands at the tip of a ridge springing from the Cuquenan 
foothills. The Cuquenan River and another stream to the 
west, which unite a quarter of a mile to the south, flank the 
ridge. Excepting a few small portions, damaged but not 
wholly destroyed by a fire, the forest has been replaced by 
deep mat of bracken, with innumerable bare whitening 
tree trunks thrusting through. Pokeweed usually mixed 
with the all-pervading bracken, has grown in as a replace- 
ment plant very abundantly at altitudes of 4,000-6,000 ft. 
The steep country south and east of Paulo is mostly devoid 
of forest; only in sheltered ravines and re-entrants small 
pockets of forest exist, invariably much damaged by fire 
(Chapman). 

127. Pescado, Caho, Terr. Amaz. — A stream between Esmeralda 
and Cerro Duida. The ground in this region is level, trees 
tall and straight and many palms are distributed through 
it (Tate). 

128. Petare, Edo. Miranda — 10°28'40"N; 66°48'32"W. -- Elev. 
844 m. — Ann. Temp. 22.0° — H. Rain., May (145), L. 
Rain., Mar. (5), Ann. Rain. 784. 

129. Pie del Avila, D. F. — See Avila. 

130. Pie del Cerro, Edo. Aragua 10°19'30"N., 67°18'50"W. 
approx. — Elev. 810-1,350 m. 

131. Puerto Ayacucho, Terr. Amaz. - - 5°53'40"N., 67°41'49"W. 

— Elev. 110 m. — II. Temp., Mar. (30.1°), L. Temp., Jul. 
(26.4°), Ann. Temp. 27.9° H. Rain., Jun. (551), L. 
Rain., Feb. (3), Ann. Rain. 2,455. Conditions, mostly semi- 
arid, large extensions of savanna occur to the south and 



192 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

east of the town but a strip of forest margins the Orinoco 
up to Sanariapo and high forest is also found on the left 
(east) of the road between Puerto Ayacucho and Sanariapo. 

132. Puerto Cabello. Edo. Carabobo - - 10°29'42"N., 68°00'31"W. 

— Elev. 5 m. — H. Temp., Oct. (27.3°), L. Temp., Feb. 
(25.2°), Ann. Temp. 26.2° - - H. Rain., Jul. (10), L. Rain., 
Feb., Mar. (3), Ann. Rain. 82. 

133. Puerto La Cruz, D. F. — 10°32'00"N., 67°21'00"W. approx. 

— Elev. 10 m. approx. 

134. Rancho Grande, Edo. Aragua - 10°21'30"N., 67°41'00"W. 

— Elev. 1,130 m. - - Ann. Temp. 18.9° approx. — H. Rain., 
Aug. (317), L. Rain., Jan. (19), Ann. Rain., 1,747. 

135. Raudal de Dios, Terr. Amaz. - - A rapid in the Cunucunuma 
River between La Culebra and Tapara. Elev. about 1,200 
ft. Conditions, heavy tropical. 

136. Riecito, Edo. Falcon — A feAV km. east of Cerro Cosme. 

137. Roraima, Cerro, Edo. Bolivar -- 5°12'00"N., 60°44'00"W. - 
Elev. 2,614 m. A flat-topped, quartzite cerro with vertical 
sides and a summit area of 25 sq. km. The Summit Camp 
of Tate and previous visitors was located in a shallow, 
rocky basin only a few hundred yards in extent, close to 
the summit of the ledge. See also Arabopo and Cuquenan. 

138. Rosario, Edo. Zulia - - 9°13'N., 72°39'W. approx. — Elev. 
not more than 100 m. A small village at the end of the 
extensive "cienagas" of the Santa Ana-Catatumbo. 

1:59. Sabana de Mendoza, Edo. Trujillo - 9°27'27"N., 
70°46'11"W. — Elev. 118 m. - H. Rain., Oct. (150), L. 
Rain., Jan. (31), Ann. Rain. 1,122. 

140. Sanariapo, Terr. Amaz. — 5°15'00"N., 67°49'00"W. - - Elev. 
150 m. approx. Conditions, semiarid. East of the village 
there is a small spring that is surrounded by a patch of 
humid forest. 

141. San Antonio, Terr. Amaz. - - 3°30'30"N., 66°44'30"W. ap- 
prox. — Elev. between 100-200 m. An abandoned planta- 
tion at the northeastern margin of the Orinoco. Conditions, 
heavy tropical. 

142. San Carlos, Terr. Amaz. - - 1°55'50"N., 67°03'50"AV. approx. 

— Elev. not more than 100 m. 

143. San Cristobal, Edo. Tachira — 7°46'11"N., 72°14'20"W. 
Elev. 830 m. — H. Temp.. Mar. (22.8°), L. Temp., Jan. 
(21.4°), Ann. Temp. (22°). II. Rain.. Jun. (218), L. Rain., 
Feb. (22), Ann. Rain. 1,463. 



RIVERO : SALIENTIA OF VENEZUELA 193 

144. San Esteban, Edo. Carabobo - - 10°25'50"N., 68°01'00"W. 
approx. — Elev. between 100-200 m. A town on Rio San 
Esteban. The river is a swift stream except below the town, 
where it flows for several miles through nearly level sand 
plain. Above San Esteban the entire valley is wooded with 
much coffee and cacao on the main stream and more native 
forest on the higher "quebradas"; below the town, the 
stream is largely in the sun (Williamson). 

145. San Felipe, Edo. Yaracuy — 10°20'38"N., 68°44'04"W. — 
Elev. 255 m. H. Temp., Oct. (26.6°), L. Temp., Jan. 
(25.4°), Ann. Temp. 26.1° — H. Rain., May (209), L. 
Rain., Mar. (20), Ann. Rain. 1,496. In the alluvial plain 
of the Yaracuy River. 

146. San Fernando de Apure, Edo. Apure — 7°53'40"N., 
67°28'11"W. — Elev. 73 m. — H. Temp., Apr. (28.7°), L. 
Temp., Jun. (26.3°), Ann. Temp. 27.4° — H. Rain., (287), 
L. Rain., Jan. (1), Ann. Rain. 1,415. 

147. San Fernando de Atabapo, Terr. Amaz. — 4°02'20"N., 
60°42'30"W. approx. — Elev. between 100-200 m. San Fer- 
nando marks the place where the savanna and semiarid 
country ends and the dense jungle begins. Collections come 
mostly from the savannas surrounding the town and from 
a fairly extensive forest patch at the edge of the Atabapo 
River. 

148. San Juan de los Morros, Edo. Guarico — 9°53'00"N., 
67°20'42"W. — Elev. 430 m. H. Temp., Apr. (25.9°), L. 
Temp., Jan. (23.2°), Ann. Temp. 24.5° — H. Rain., Jul. 
(301), L. Rain., Feb. (5), Ann. Rain. 1,342. 

149. San Juan, Rio, Edo. Zulia — Tributary of Rio Motatan, in 
the southeastern part of Lake Maracaibo. 

150. San Julian, D. F. — 10°36'N., 66°51'W. approx. — Elev. 
near sea level. The hamlet, consisting of a few scattered 
huts, is located in an irrigated valley above the village of 
Caraballeda. The mountains behind San Julian are well 
forested (Robinson and Lyon). 

151. Santa Ana, Rio, Edo. Zulia — Large river on the western 
side of Lake Maracaibo. 

152. Santa Catalina — Apart from the fact that it is in Rio 
Orinoco, I have been unable to obtain any other information 
about this locality. 

153. Santa Elena, Edo. Bolivar — 4°36'05"N., 61°06'52"W. — 
Elev. 907 m. — H. Temp., Mar. (22.4°), L. Temp., Jul. 
(21.1°), Ann. Temp. 21.8° — H. Rain., Jun. (251), L. 



194 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Rain., Feb. (62), Ann. Rain. 1,628. In the Gran Sabana, 
in the angle formed in the southeast by Venezuela, British 
Guiana and Brasil. 

154. Santa Elena, Edo. Merida — 8°50'30"N., 71°35'30"W. ap- 
prox. — Elev. not more than 100 m. In the alluvial plain 
characterized by heavy, humid tropical forest (Osgood and 
Conover). 

155. Santa Lucia, Edo. Miranda — 10°18'N., 66°39'W. approx. 
— Collections come from elevations between 300 and 810 m. 

156. Sierra Maestra, Edo. Aragua — I have been unable to ob- 
tain any information about this locality. 

157. Sosa, Edo. Guarico — Elev. 200 m. approx. On Rio 
Guarico. Frogs collected in the grass at the margin of the 
river. Gallery forest follows the course of Rio Guarico but 
the rest of the country is savanna covered (Roze). 

158. Tachira, Edo. Tachira 8°07'N., 72°20' W.approx. — 
Elev. about 365 m. Heavily wooded region of steep or pre- 
cipitous mountain sides and many streams (Williamson). 

159. Tanaguarena, D. F. — In the vicinity of La Guaira and 
Macuto. Elev. 10 m. Conditions, arid to semiarid (Roze). 

160. Tapara, Terr. Amaz. — The last navigable (canoe) point 
of Rio Cunucunuma. Conditions, humid tropical except at 
the edge of the river where the trees are sparse and small. 

161. Temiche, Terr. Amaz. — A camp on the southern slopes of 
Cerro Marahuaca, at an elevation of 4,050 ft. Conditions, 
humid tropical or lower subtropical. A great abundance of 
an enormously leaved palm and tall trees that make the 
place very umbrageous. A stream at about 50 ft. from 
camp. 

162. Trujillo, Edo. Trujillo - - 9°22'24"N., 70°26'08"W. - - Elev. 
790 m. — H. Rain., Nov. (137), L. Rain., Jan. (41), Ann. 
Rain. 936. 

163. Tucacas, Edo. Falcon — 10°15'30"N., 68°19'08"W. — Elev. 
4 m. H. Rain., Nov. (212), L. Rain., Feb. (37), Ann. Rain. 
1,053. Dry woods and dry Heliconia patches adjacent to 
the town; back of it, a small boggy stream. 

164. Tunapucito, Edo. Sucre - 10°36'00"N., 63°09'30"W. ap- 
prox. — Elev. about 300 m. 

165. Turgua, Edo. Miranda — At about 20 km. from Caracas, 
between Petare and Baruta. Elev. 1,100 m. approx. Con- 
ditions, subtropical (Roze). 



RIVERO : SALIENTIA OF VENEZUELA 195 

166. Turumiquire, Cerro, Edos. Sucre-Monagas — 10°7'N., 
63°52'W. approx. Elev. 2,630 m. A mountain on the eastern 
part of the Coastal Range. 

167. Unare, Edo. Anzoategui - - 10°05'3O"N., 65°11'00"W.— 
Elev. sea level approx. 

168. Upata, Edo. Bolivar — 8°01'07"N., 62°24'33"W. — Elev. 
340 m. H. Rain., Mav (138), L. Rain., Feb. (0), Ann. Rain. 
868. 

169. Uribante, Rio, Edos. Tachira and Barinas — Tributary of 
Rio Apure, with headwaters in the Andes of Tachira and 
mouth at 7°18'N., 70°46'W. approx. 

170. Valencia, Edo. Carabobo — 10°11'10"N., 67°59'58"W. — 
Elev. 478 m. — H. Temp., Oct. (24.9°), L. Temp., Jan. 
(24.0°), Ann. Temp. (24.6°). H. Rain., Aug. (187), L. 
Rain., Mar. (3), Ann. Rain. 1,120. 

171. Villa del Rosario, Edo. Zulia — 10°19'54"N., 72°19'30"W. 

— Elev. 80 m. approx. 

172. Wanadi, Caiio, Terr. Amaz. — A swift running stream on 
the slopes of Mt. Marahuaca. Collections from the stream 
or from the surrounding heavy jungle at an elevation of 
1,225 m. 

173. Waika-piapu Mt, (Ualacatipu), Edo. Bolivar — A moun- 
tain on the west north-west of Roraima and Cuquenan. 
Together with the Urutipu and Charantipu it forms a ridge 
that cuts off the Mazaruni and Cuyuni Rivers. 

174. Yapacana, Cerro, Terr. Amaz. - - 3°42'N., 62°45'W. approx. 

— Elev. 1,250 m. 

175. Yumarito, Rio, Edo. Yaracuy — A stream in Boqueron. 

176. Zaraza, Edo. Guarico - - 9°20'09"N., 65°19'03"W. — Elev. 
60 m. — H. Rain., Jul. (241), L. Rain., Feb. (6), Ann. 
Rain. 1,144. 

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Sammlung des bayr. Staates, Zool. Anz., 57 (%): 38-42. 

1927. Amphibien und Reptilien der Ausbeute Prof. Bresslau's in Bra- 
silien 1913-14. Abh. Senckenb. Naturf. Ges., 40 (3): 259-304. 

1934. Uber eine neue Rasse von Atelopus cruciger (Licht. u. Marts.) 
von Venezuela. Zool. Anz., 108 (7/8) : 145-155. 

1938. Batrachologische Mitteilungen. Zool. Anz., 121 (9/10) : 284- 
288. 



RIVERO : SALIENTIA OF VENEZUELA 203 

MULLER, LORENZ AND WALTER HELLMICH 

1936. Wissenschaftliche Ergebnisse der deutschen Gran Chaco-Expe- 

dition; Aniphibien und Reptilien. Stuttgart. Pp. i-x 4- 1-120, 

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Netting, M. Graham 

1930a. A new Toad of the Genus Eupemphix. Ann. Carnegie Mus., 19 

(3) : 167-168, pi. VII, figs. 1-2. 
1930b. The Systematic Status and Breeding Habits of Eupemphix 

trinitatis Boulenger. Ann. Carnegie Mus., 19 (4) : 249-254. 
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1923. Anura I. Das Tierreich, 46: i-xxxii 4- 1-584 pp., figs. 1-380. 

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1926. Anura II. Das Tierreich, 49: i-xvi 4- 1-110 pp., figs. 1-55. 
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Noble, G. K. 

1917. The Systematic Status of Some Batrachians from South Amer- 
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1918. The Amphibians Collected by the American Museum Expedition 
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1920. The New Batrachians from Colombia. Bull. Amer. Mus. Nat. 

Hist., 42 : 441-446. 
1923. New Batrachians from the Tropical Research Station, British 

Guiana. Zoologica (N.Y.), 3 (14): 289-305, 3 text maps. 
1925. A new Genus of Surinam Toads (Pipidae). Amer. Mus. Novit., 

No. 164: 1-3. 

1931. The Biology of the Amphibia. New York. Pp. i-xiii + 1-577, 
figs. 1-174. 

Osgood, Wilfred H. and Boardman Conover 

1922. Game Birds from Northwestern Venezuela. Zool. Ser. Field. 
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Parker, H. W. 

1927. A Revision of the Frogs of the Genera Pseudopaludicola, Physa- 
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478, figs. 1-5. 

1928. Notes on Reptiles and Batrachians from Matto Grosso and E. 
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1932. The Systematic Status of Some Frogs in the Vienna Museum. 
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1933. A List of the Frogs and Toads from Trinidad. Trop. Agr. 
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1934. A Monograph of the Frogs of the Family Microhylidae. Lon- 
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1935. The Frogs, Lizards and Snakes of British Guiana. Proc. Zool. 
Soc. London, 1935: 505-530. 

1936. A Collection of Reptiles and Amphibians from the Upper Ori- 
noco. Bull. Mus. Roy. Hist. Nat. Belgique, 12 (26) : 1-4. 



204 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

1938. The Vertical Distribution of Some Reptiles and Amphibians in 

Southern Ecuador. Ann. Mag. Nat. Hist,, (11) 2: 438-450. 
1940. Undescribed Anatomical Structures and New Species of Reptiles 

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Peracca, Mario Giacinto 

1904a. Viaggio del Dr. A. Borelli nel Matto Grosso brasiliano e nel 

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1914. Reptiles et Batrachiens de Colombie. Mem. Soc. Sci. Neuchatel, 

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Peters, Wilhelm 

1859. Hr. Peters legte eine neue Gattung und eine neue Art von 

Frbschen aus Caracas vor. Monatsb. Akad. Wissensch. Berlin : 

402-403. 
1863. Hr. W. Peters machte eine Mittheilung fiber eine neue Schlangen- 

gattung, Styporliynchus, und verschiedene andere Amphibien 

des zoologischen Museums. Monatsb. Akad. Wissensch. Berlin: 

399-413. 
1868. Hr. W. Peters machte fernere Mittheilungen fiber neue Batra- 

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1872. Hr. W. Peters machte eine Mittheilung fiber eine Sammlung von 

Batrachiern aus Neu-Freiburg in Brasilien. Monatsb. Akad. 

Wissensch. Berlin: 680-684. 
1877. Sammlung des Hr. Dr. Carl Sachs in Venezuela. Monatsb. 

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Phelps, William 

1938a. La Expedicion del American Museum of Natural History al 

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1938b. La Proeedencia Geografica de las Aves Coleccionadas en el Cerro 

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1943. Las Aves de Perija. Bol. Soc. Ven. Cienc. Nat., 8 (56) : 265- 

338, 2 pis., 2 maps. 
Phelps, William and William H. Phelps Jr. 

1948. Lista de las Aves de Venezuela con su Distribucion, Parte 2, 

Passeriformes. Bol. Soc. Ven. Cienc. Nat., 12 (75): 1-427. 
Piatt, Jean 

1934. The Systematic Status of Eleutherodactylus latrans (Cope). 

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Procter, Joan B. 

1921. On a Small Collection of Reptiles and Batrachians made by Mr. 

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7: 189-192. 
Reinhardt, J. and C. Lutken 

1861. Bidrag til Kundskab om Brasiliens Padder og Krybdyr. Vid- 

densk. Medd. Nat. Forening, no. 1 : 143-203, pis. I-IV. 



RIVERO : SALIENTIA OF VENEZUELA 205 

Robinson, Wirt and Marcus Ward Lyon Jr. 

1902. An Annotated List of Mammals Collected in the Vicinity of La 
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1949. Fauna Descriptiva de Venezuela, Vertebrados, ed. 2 (Caracas). 
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Roux, Jean 

1927. Contribution a 1 'erpetologie du Venezuela. Verb. Naturf. Ges. 
Basel, 38 : 252-261. 
Ruthven, Alexander 

1914. Description of a new Engystomatid Frog of the Genus Hypo- 

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1916. A new Species of Paludicola from Colombia. Occ. Pap. Mus. 
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1917. Two new Species of Eleutherodactylus from Colombia. Occ. Pap. 
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1919. The Amphibians of the University of Michigan-Walker Expedi- 
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Ruthven, Alexander and Helen T. Gaige 

1923. Description of a new Species of Pipa from Venezuela. Occ. 
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1953. The Family Position of Neotropical Frogs Currently Referred 
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1735. Loeupletissimi Rerum Xaturalium Thesauri Aecurata Descrrptio, 
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1932. Reptiles and Amphibians of the Mandel Venezuelan Expedition. 
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Schneider, Johann Gottlob 

1799. Historiae Amphibiorum Naturalis et Literarie. Jena, 1: 1-264, 
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1947. On Venezuelan Reptiles and Amphibians Collected by Dr. II. G. 
Kugler. Bull. Mus. Comp. Zool., 99 (5): 517-537. 



20(i BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Spix, Johann Baptist von 

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1891. Description of a New Genus and Species of Tailless Batrachian 
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1930. Notes on Mount Roraima Region. Geogr. Rev., 20: 31-52, 9 

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1884. Note sur un Batracien d 'espece nouvelle provenant de Panama. 
Bull. Soc. Philom. Paris, (7) 8: 151-152. 



RIVERO: SALIENTIA OF VENEZUELA 207 

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1854. Uber den Beutelfrosch. Archiv. Anat. Physiol. 1854: 449-477, 
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Hyla platydactyla Boulenger, type specimen. Courtesy of British Mu 
seum (Natural History). 



Bulletin of the Museum of Comparative Zoology 

AT HAEVAED COLLEGE 

Vol. 126, No. 2 



A REVISION OF THE GENUS PSEUDISOBRACHIUM IN 
NORTH AND CENTRAL AMERICA 
(HYMENOPTERA, BETHYLIDAE) 



By Howard E. Evans 



CAMBEIDGE, MASS., U.S.A. 
PEINTED FOE THE MUSEUM 

December, 1961 



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Bulletin of the Museum of Comparative Zoology 

AT HARVARD COLLEGE 

Vol. 126, No. 2 



A KEVISION OF THE CrENUS PSEUDISOBRACHIUM IN 
NORTH AND CENTRAL AMERICA 
(HYMENOPTERA, BETHYLIDAE) 



By Howard E. Evans 



CAMBRIDGE, MASS., U.S.A. 
PRINTED FOE THE MUSEUM 

December, 1961 



No. 2 - A Revision of the Genus Pseudisobrachium in North 
and Central America (Hymenoptera, Bethylidae) 

By Howard E. Evans 

INTRODUCTION 

The bethylid genus Pseudisobrachium contains approximately 
100 known species, about four-fifths of which occur in the West- 
ern Hemisphere and about two-fifths of which are described for 
the first time in the present revision. There are many unde- 
scribed Neotropical species, but I have had to omit the West 
Indies and South America from present treatment because of 
inadequate material. The genus is well represented in Europe 
and in Africa, and the species from these continents are closely 
similar to the American forms. Two species from the Philippines 
described by Kieffer (1922) represent the only known Oriental 
species. Although Kieffer (1914) states that the genus occurs in 
Australia, I have seen no specimens from that continent and the 
only species described from there, P. australiensis Kieffer, was 
correctly removed to Propristocera by Kieffer himself. 

The North American species were first treated by Ashmead 
(1893) under the name Isobrachium, a name now properly re- 
stricted to an unrelated Palaearctic genus. Ashmead described 
and presented a key to six species; he apparently had before him 
about 15-20 specimens. The characters he used now seem un- 
reliable and in some cases are actually incorrect ; his sex associa- 
tions also tended to be capricious. Kieffer (1906) described one 
species from Nicaragua and a few years later (1914) presented 
a key to the species of the world, though there is no evidence that 
he studied much if any North American material. Fouts (1928) 
described five new species and presented a revised key to species ; 
he apparently had about 15 specimens before him in addition to 
Ashmead 's material. 

I first became interested in the genus about ten years ago and 
began slowly to gather material. It quickly became apparent that 
the genus was a large and difficult one and that the existing 
literature was completely inadequate. The present revision, al- 
though based on about 1400 specimens, pretends to be no more 
than a very preliminary treatment of the genus. Reliable species 
characters have been more difficult to find than, for example, 
in the related genera Dissomphahis and Propristocera, and much 
difficulty has been encountered in associating the sexes properly. 



212 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Nevertheless, I do feel that the revision represents a sufficient 
advance over earlier work to justify my publishing it at this 
time. 

There are those who question the value of taxonomic studies 
on a strictly "alpha" level, as this one admittedly is. I can 
appreciate their arguments and I readily admit that the species 
of the alpha taxonomist are no more than hypothetical entities. 
Only more sophisticated studies, involving not only careful, sta- 
tistically analyzed inventories of the structure of properly 
sampled populations, but also comparative studies of physiology 
and behavior, can establish with certainty how a complex of 
organisms ought properly to be classified. But such studies must 
have a beginning. Pseudisobrachium has been, until recently, like 
a vaguely plotted continent on an explorer's map. The present 
study is no more than a beach-head on that continent, but it may 
perform the important function of serving as a base for further 
operations. 

ACKNOWLEDGMENTS AND SOURCES OF MATERIAL 

Pseudisobrachium is poorly represented in the collections of 
most museums, the sole exception being the U. S. National 
Museum. However, by gathering material from many sources I 
have been able to amass enough material for this preliminary 
study. I am particularly indebted to Karl V. Krombein and 
Henry K. Townes for making available the valuable material in 
their personal collections. A. T. McClay, at Davis, California, 
M. Wasbauer, at Sacramento, California, and Mrs. L. K. Gloyd, 
at Urbana, Illinois, have provided me with excellent series of 
certain species taken from light trap collections. My own col- 
lecting has enabled me to fill in many gaps. A trip to Mexico 
during the summer of 1951 was supported by the American 
Philosophical Society, and a second trip to Mexico and the south- 
western United States during 1959 was made possible by a 
fellowship from the John Simon Guggenheim Memorial Founda- 
tion. Additional specimens were collected during several trips 
to the southern states supported by the National Science Founda- 
tion for studies on the behavior of digger wasps. 

The following list of institutions and individuals is meant 
to serve as an acknowledgment to each, as well as an indication 
of the abbreviation by which each is designated in the text : 

Academy of Natural Sciences of Philadelphia (ANSP) 

American Museum of Natural History, New York (AMNH) 



EVANS: REVISION OF PSEUDISOBRACHIUM 213 

California Academy of Sciences, San Francisco (CAS) 

California Dept. of Agriculture, Sacramento (CDAS) 

California Insect Survey, Berkeley (CIS) 

Canadian National Collections, Ottawa (CNC) 

Carnegie Museum, Pittsburgh (CM) 

Cornell University, Ithaca, N. Y. (CU) 

Florida State Plant Board, Gainesville (FSPB) 

H. K. Townes Collection, Ann Arbor, Mich. (HKT) 

Illinois Natural History Survey, Urbana (INHS) 

Kansas University, Lawrence (KU) 

Kansas State University, Manhattan (KSU) 

K. V. Krombein Collection, Arlington, Va. (KVK) 

Museum of Comparative Zoology, Cambridge, Mass. (MCZ) 

Oficina de Estudios Especiales, Secretaria Agricultura y 

Ganaderia, Mexico, D. F. (OEE) 
R. R. Dreisbach Collection, Midland, Mich. (RRD) 
University of Arizona, Tucson (UA) 
University of California, Davis (UCD) 
United States National Museum, Washington, D.C. (USNM) 

BIOLOGY OF THE GENUS 

Pseudisobrachium is properly placed as one of the more spe- 
cialized genera of the tribe Pristocerini. The most closely related 
genus in the Americas is probably Propristocera, a genus in which 
the males lack a transverse carina margining the propodeal disc 
behind, as in Pseudisobrachium. It is somewhat more distantly 
related to Pristocera, Cleistepyris, and Dissomphalus. Pluto- 
Parisobrachium Kieffer I regard as synonyms of Pseudiso- 
brachium (see further discussions under generic diagnosis 
below). 

It must be admitted regrettably that Ashmead's 1893 state- 
ment regarding the biology of the genus cannot yet be im- 
proved upon. In Ashmead's words, "the genus is found associ- 
ated with various ants; it may be parasitic upon the ants, or 
upon the myrmecophilous Coleoptera so frequently found in their 
nests. ' ' There are now many more records of females being taken 
with ants than there were in Ashmead's time (see list below), 
but the precise relationship remains to be determined. It should 
be noted that the species of Pristocera attack coleopterous larvae, 
and there is some evidence that the species of Dissomphalus do 
also. Unfortunately, nothing is known of the biology of other 
genera of Pristocerini. It is, of course, entirely possible that 



214 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Pseudisobrachium has switched from coleopterous to formicid 
larvae, possibly by way of myrmecophilous forms. There is now 
strong circumstantial evidence that members of this genus do in 
fact attack ant larvae (see especially remarks under P. arenarium 
below). This may account for the great success of the genus 
as compared to most other genera of the tribe. Females are also 
sometimes taken in Berlese samples of soil or leaf litter, pre- 
sumably because the sample includes a portion of an ant nest. I 
have also seen an occasional female from light trap collections, 
suggesting that females may sometimes leave the soil during the 
night. 

A perusal of the list presented below shows that female 
Pseudisobrachium have been taken with ants of several subfami- 
lies (Formicinae, Dolichoderinae, Myrmicinae, Dorylinae, Poner- 
inae). Where there are several records for one species, in most 
cases that species appears to be associated with several genera, 
sometimes of more than one subfamily. For example, prolonga- 
tum has been taken with the formicine genera Camponotus, 
Acanthomyops, and Formica, as well as the myrmicine genus 
Aphaenog aster. The closely related arenarium is recorded with 
the ponerine genera Proceratium and Stigmatomma. P. ashmeadi 
has been taken with the genera Formica and Acanthomyops (For- 
micinae) and Tapinoma (Dolichoderinae). While Formicinae 
have been taken with three of six Nearctic species, none of the 
exotic species has yet been taken with ants of this subfamily. 

Since the females are never taken by routine collecting meth- 
ods, they are rare in collections. Males are much more frequently 
encountered. Males of some species are diurnal and often taken 
by sweeping low vegetation either in forests or in open country. 
I have taken them on honeydew only rarely, and know of only 
one or two records of them having been taken on flowers. Males 
of certain other species appear to fly either by day or in the 
evening, while many species, especially those characteristic of 
arid country, are fully nocturnal. Nocturnal species tend to have 
larger eyes and ocelli and are often paler in color (especially 
the wings). However, some forms with small eyes and ocelli 
have been taken only at night. Many species exhibit considerable 
variation, in color, eye size, and ocellar size, and it is possible that 
some of this variation is associated with ecological conditions 
and the time of flight of local populations. The majority of males 
of this genus in collections have been taken at light. 



EVANS : REVISION OF PSEUDISOBRACHIUM 215 

Most of the South American and Palaearctic males I have seen 
are dark in color and have relatively small ocelli, and I am not 
aware of reports that these species are nocturnal. The evolution 
of numerous nocturnal species seems to be especially character- 
istic of desert regions of North America, somewhat paralleling 
the development of the brachycistidine Tiphiidae. 

Further comment is perhaps in order on the difficulty in as- 
sociating sexes in this genus. Of the North American species, 
the sexes of only two (prolong atum and rufiventre) have been 
associated with any degree of certainty. Theoretically, one 
should be able to collect males in numbers in certain situations, 
then search the ant nests of that area diligently for females ; 
having found a female, one ought to be able to place her in 
an open container in the field and capture the males as they 
are attracted to her. I have actually given a good deal of thought 
and time to this scheme, but with completely negative results. 
On one occasion, I collected 67 male prolong atum on a small 
wooded hillock within a few hours (Plummer's Island, Mary- 
land, Sept. 23, 1960). However, a very careful search of all the 
ant nests I could find in the area revealed no females. Of course, 
in the late summer (when male Pseudisobrachium are on the 
wing in the Northeast), most ant nests contain little brood. It is 
entirely possible that both sexes emerge in the fall and mate 
and that the females overwinter but do not enter ant nests until 
spring, when ant colonies are growing and contain much brood. 
The majority of females which have been taken in ant nests have 
been taken in the spring or early summer. 

RECORDS OF PSEUDISOBRACHIUM TAKEN WITH 

ANTS 1 

Nearctic Species 

P. arenarium Evans. 2 Specimen from St. Charles, Mo., 1949 (M. 
Talbot), labeled "in vial with Stig[matomma] pallipes" (Pon- 
erinae) [USNM]. Specimen from Philadelphia, Pa., 20 May 
1939 (W. L. Brown), labeled "from nest of Proceratium sp." 
(Ponerinae) [USNM]. Dr. Brown has written me that this 
specimen was taken in Wissahickon Park with Proceratium 
silaceum (Roger). The nest was under a rock in deciduous 



1 All records Involve female Pseudisobrachium except for one Oriental record 
which is noted otherwise. 

2 These species are described in the text from the male sex, and in each case 
the sex association is tentative. 



216 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

woods; there were 13 adult ants in the nest, 11 workers and 2 
ergatoid females. The Pseudisobrachium was in the middle of 
the nest with brood in a space less than one inch in diameter. 
There were no myrmecophilous beetles in this nest and in fact 
both this ant and Stigmatomma pallipes are not known to har- 
bor such beetles. 

P. ashmeadi Evans. 2 Specimen from Arlington, Mass., 24 May 
1953 (W. L. Brown), labeled ''Formica fusca nest in woods" 
[MCZ]. Specimen from Forest Hills, Mass., 4 May 1915 (W. 
M. Mann), with two Tapinoma sessile (Say) [det. W. L. 
Brown] on card point on same pin [USNM]. Specimen from 
Forest Hills, Mass. (no date) (W. M. Mann) with Acantho- 
myops claviger (Roger) on card point on same pin [USNM]. 
Tapinoma belongs to the Dolichoderinae, the other two genera 
to the Formicinae. 

P. occidentale Evans. 2 Specimen from Stanford Univ., Calif., 
13 Feb. 1910, with a worker and dealate queen Tapinoma sessde 
(Say) [det. W. L. BroAvn] on card point on same pin (Dolicho- 
derinae) [USNM]. 

P. prolongation (Provancher) ( = mandibulare Aslimead, mon- 
tanum Ashmead, myrmecophilum Aslimead). Ashmead 
(1893) records mandibulare from a nest of Camponotus 
pennsylvanicus, 27 May 1883 (T. Pergande) ; this specimen is 
in the USNM, from Washington, D. C, bearing a note to that 
effect. A second specimen in the USNM, from Washington, 
D. C, 19 Aug., is pinned with two specimens of Camponotus 
nearcticus Emery [det. W. L. Brown] . The type of mandibulare, 
from Retreat, Haywood Co., N. C. [USNM] is pinned with a 
worker Aphacnogaster fulva Roger (Myrmicinae). Ashmead 
(1893) also recorded myrmecophilum from nests of Formica 
rufibarbis at Helena, Mont., and montanum from nests of this 
ant at Assiniboine, Mont, (both II. G. Hubbard) ; the ant 
associated with the specimen from the latter locality has been 
identified as Formica cinerea Mayr by W. L. Brow T n. Ash- 
mead's record of rufiventre from Helena, Mont., from a nest 
of Formica obscuripcs Forel, also doubtless applies to prolon- 
gatum. There are two additional specimens in the USNM, both 
from Forest Hills, Mass., May 1915 (W. M. Mann) ; one is 
pinned with a Formica of the pallide-fulva group, the other 
with Acant]iomyops sp. ( Iclaviger Roger) [det. W. L. Brown]. 
There are two specimens in the MCZ from Lexington, Mass., 



EVANS : REVISION OF PSEUDISOBRACHIUM 217 

5 June 1955 (W. L. Brown), one labeled "in nest of Aca/ntho- 
myops claviger," the other labeled "in nest of Acanthomyops 
sp. indet." A specimen from Walsh Co., N. D., 23 June 1950 
(W. E. LaBerge) bears the notation "with Formica sp." 

P. flaviventre (Kieffer). Dr. W. L. Brown has sent me a specimen 
probably belonging to this species which he took at Little 
Grassy Lake, Williamson Co., 111., 10 Aug., 1958, in oak-hickory 
woods, in leaf litter in or near a nest of Solenopsis of group 
molest a Say (Myrmicinae). A number of specimens of this 
species have been taken in soil (see text for data). 

P. rufiventre (Ashmead). Ashmead's (1893) record of this spe- 
cies from Montana probably applies to prolongatum, as noted 
under that species. A specimen from Blue Hills, Canton, Mass., 
20 July 1956 (W. L. Brown) is labeled "Formica ouscuri- 
ventris Mayr nest under rock" [MCZ]. 

Neotropical Species 

P. merklei Bruch. Bruch (1917b) described this species along 
with several other myrmeeophiles taken near La Plata, Ar- 
gentina. In his introduction he states : ' ' mencionare por ahora 
los insectos de nuestras capturas y que podemos considerar en 
cierto modo como huespedes de Solenopsis saevissima" (Myr- 
micinae). 

P. solenopsiclicola Bruch. Bruch (1917a) described this species 
"de un nido de Solenopsis saevissima Sm. var. tricuspis Forel, 
en la Sierra de la Ventana [Argentina] ' (Myrmicinae). 

P. terresi Mann. "Described from a single specimen taken in a 
nest of Aphaenogaster relicta at Diquini [Haiti]" (Myrmi- 
cinae) (Mann, 1915). 

Palaearctic Species 

P. leptanillae Duchaussoy. Duchaussoy (1916) described this 
species from "Tunisie, Kairouan, Sousse ; trouve par M. le 
Dr. Santschi, dans les nids des fourmis du genre Leptanilla" 
(Dorylinae). 

P. suocyaneum Haliday (= cantianum Chitty). Chitty (1906) 
took the type of cantianum "in nest and runs of Ponera con- 
tracta," Charing, Kent, England, 3 Aug. 1903. He surmises 
that subcyancum may attack Myrmecina latreillei, (= M. 
graminicola Latr.) but gives no reason for this belief. 



218 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Oriental Species 

P. sp. A specimen of an apparently undescribed male Pseudiso- 
brachium from San Francisco del Monte, Manila, P.I. (R. C. 
(McGregor) [MCZ] is pinned with two worker Odontomachus 
simillimus (F. Smith) (Ponerinae) [det. E. 0. Wilson]. There 
are no accompanying data, so it can only be tentatively as- 
sumed that this male was taken from a nest of the ant. 

STRUCTURE AND TERMINOLOGY 

Sexual dimorphism is so pronounced in this genus that it is 
necessary to treat the two sexes separately both here and in 
the keys and descriptions. Following a discussion of the useful 
characters of each sex I have provided a summary of the various 
abbreviations employed in the text. 

Males. — Males are fully winged, usually dark in color, and 
vary in size from 2 to 6 mm. The mandibles (Figs. 7-32) have 
from 3 to 5 teeth ; these teeth are simply numbered one to 
five, starting with the apical (outermost) tooth. Mandibular 
dentition is extremely important, and it is desirable to spread 
the mandibles of freshly caught specimens. The clypeus has 
a median elevation and is extended apically as a more or less 
truncate lobe (Figs. 1-6, 43-50). The head (Figs. 1-6) is sub- 
lenticular or somewhat elongate and has prominent, hairy eyes 
and three ocelli. Length of the head (LH) is measured from 
the vertex crest to the median apical margin of the clypeus, width 
of the head (WH) at the maximum point, including the eyes; 
both of these measurements are made at full frontal view. AVidth 
of the front (WF) is measured at its minimum point, usually 
about the middle of the eyes. The height of the eye (HE) is 
measured at its maximum in lateral aspect. As an indication of 
ocellar size, I have measured the transverse diameter of the 
anterior ocellus (DAO) and compared this with the width of 
the front; in species with the smallest ocelli DAO is about .10 X 
WF, while in species with the largest ocelli DAO is nearly 
.40 X WF. The ocello-ocular line (OOL) is the shortest distance 
from the eye tops to the lateral ocellus, measured by looking 
directly down upon it with the specimen appropriately tilted. This 
measurement is frequently compared to the width of the ocellar 
triangle (WOT), which is simply the distance across (and in- 
cluding) the posterior ocelli. The front angle of the ocellar 
triangle varies interspecifically and provides an important means 



EVANS: REVISION OF PSEUDISOBRACHIUM 219 

of separating certain species. I have measured this by super- 
imposing the right angle of a grid micrometer over the outer 
sides of the anterior and one lateral ocellus and determining 
whether the angle of the line extending to the other lateral ocellus 
is greater or less than a right angle. The occipital carina is 
obsolete dorsally except as otherwise noted. The antennae have 
13 segments, of which the scape is much the largest. The relative 
length of the flagellum is of much importance, and in order to 
have an easily comparable measurement I have selected segment 
eleven (the antepenultimate) for measurement. In species with 
especially long antennae, this segment measures two or more 
times as long as wide, while in specimens with very short an- 
tennae it may actually be wider than long. This measurement is 
especially subject to error because of the tendency of the anten- 
nae to coil and because the segments are capable of some ex- 
tension (specimens preserved in alcohol tend to give higher 
measurements). 

The term thorax is here employed to mean the functional 
thorax or alitrunk. The pronotum is gradually expanded behind 
and shows little variation in the genus. On the mesonotum, the 
mesoscutum possesses well developed parapsidal furrows, while 
the more median notauli show much variation in degree of de- 
velopment ; the scutellum has a basal transverse groove, a median 
elevated disc, and large lateral foveae. The propodeum shows 
much variation in length and measurements have been found 
useful. Its total length, from margin of metanotum to posterior 
rim (articulation with gaster) is compared with its maximum 
width, both these measurements being made in full dorsal view. 
In lateral view, the height of the propodeum is measured from 
the carina running forward from above the hind coxae. A single 
median carina and lateral carinae margining the disc are pres- 
ent in all species except as otherwise noted. The mesopleurum 
is oblique and elongate and is the source of useful taxonomic 
characters, though these are often subtle and difficult to express. 
At the top, just below the base of the hind wings, is a more or 
less well defined swelling, here called the mesopleural callus ; this 
is subtended by a usually elongate impression ; the remainder of 
the mesopleurum tends to be sculptured in various ways. The 
legs are slender and short-haired, and aside from slight differ- 
ences in dimensions of the femora show no important specific 
differences. The tarsal claws normally have a single erect tooth, 
but this tooth may be extremely weak or (in several South Ameri- 
can species) entirely absent. The fore wings (Figs. 51-62) have a 



220 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

characteristic venation which shows little variation within the 
genus. The vein which extends beyond the origin of the basal and 
transverse median veins and is more or less in a straight line 
with the median vein is here termed the discoidal vein, the cell 
below it the discoidal cell. This vein may be well defined, weak, 
faintly indicated but unpigmented, or entirely absent. On the 
outer part of the wing there are a number of hyaline streaks 
which presumably mark the course of former veins ; these are 
omitted from the figures, and I have found them of no particular 
value in classification. The hind wing has a single strong but 
short vein at the base and a strong jugal lobe. The length of the 
fore wing (LFW) is a useful measurement for size of the wasp, 
as total body length is difficult to measure accurately. 

The term abdomen is here used to mean the gaster (true abdo- 
men minus the propodeum) in the common manner of hymenop- 
terists. The apical abdominal sternite or subgenital plate is of 
simple structure and usually somewhat truncate apically. The 
genitalia are highly characteristic of the genus and show very 
little variation within the genus (including also Old World 
species). Their typical form is shown in Figure 63. The para- 
meres are deeply divided into a relatively broad outer arm and 
a more slender inner arm, the latter having much fewer setae. 
The volsellae are complex, with digitus and cuspus about as 
figured in all species, but looking very different from different 
points of view ; at the base of the volsellae, medially, is a portion 
containing several strong radiating sulci, resembling a fan and 
therefore here termed the vannus (absent in one species). The 
aedoeagus is normally flat, somewhat bottle-shaped, and of simple 
structure, though in one species it is compressed and of more 
complex structure (Fig. 67). 

Female. — Females are completely apterous and without tegu- 
lae, and show corresponding reductions in the thorax (Reid, 1941, 
figured and discussed the thorax of the related genus Mangesia, 
from Africa). They are nearly sightless, usually pale in color, 
and clearly modified for a hypogaeic existence. They vary in 
length from 1.8 to 6.5 mm., the females of a given species being 
(on the average) considerably smaller than the males. The mand- 
ibles have three or four teeth, sometimes only two teeth which are 
prominent, the third being very small (Figs. 33-42). The clypeus 
is very broad, truncate or somewhat emarginate, and has a 
median ridge. The scape is large, the flagellum short and some- 
what thickened from the base, the flagellar segments (except the 



EVANS : REVISION OF PSEUDISOBRACHIUM 221 

last) broader than long; the antennae are much alike in all 
species. The eyes consist of a single facet on each side ; in some of 
the smaller species the eyes cannot be seen at all, but it is difficult 
to be sure that the facet is not there but indistinguishable from 
the surface sculpturing. The head itself is subquadrate, slightly 
longer than wide. Length of the head (LH) is measured from 
median apex of clypeus to median crest of vertex in full frontal 
view. Width of the head (WII) is measured at the mid-point of 
LH ; usually the head is not wider elsewhere than at this point, 
but in a few species it is slightly wider anteriorly than here. The 
occipital carina is obsolete dorsally except in one species as noted. 
Ocelli are absent. 

The length of the thorax (LT), again including the propo- 
deum, is measured from the anterior end of the pronotal disc 
(omitting the depressed collar) to the posterior end of the pro- 
podeum. The pronotum is large and is longer than broad ; its 
median length is also measured without consideration of the 
collar, while its width is measured posteriorly, across the rather 
prominent shoulder-like corners. The mesonotum is strongly 
depressed anteriorly, fitting against the pronotum ; this is the 
mesoscutum according to Reid (1941). The remainder of the 
mesonotum, although called the mesonotum in the text, is said 
by Reid to be the scutellum ; it is triangular, somewhat pointed 
posteriorly, and is measured at its maximum points (anteriorly 
for width, medially for length, omitting the depressed portion). 
The mesopleura are large and have dorsal portions which flank 
the mesonotum ; these ronud off laterally and form much the 
widest part of the thorax. The propodeum is strongly narrowed 
anteriorly to a pair of pointed processes which embrace the 
posterior point of the mesonotum ; the disc is subcarinate lat- 
erally, the carinae being interrupted at one point by the spi- 
racles ; the disc is without strong sculpturing of any kind. The 
legs are short-haired and without spines except for the middle 
tibiae and tarsi, which are strongly spinose. The abdomen ( = 
gaster) is sessile, sub fusiform, and terminates in a short sting 
which is embraced by stout sting-sheaths. 

ALPHABETICAL LIST OP ABBREVIATIONS 
USED IN TEXT 

Ant. 11 : antennal segment eleven (antepenultimate) 
DAO : diameter of anterior ocellus (transversely) 
HE : height of eye (maximum, lateral view) 



222 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

L : length 

LP W : length of fore wing 

LH : length of head (full front view, including clypeus but not 

mouthparts) 
LT: length of thorax (excluding collar but including propo- 

deum ) 
OOL : ocello-ocular line 
W: width 

WF : width of front (at minimum point) 
WII: width of head (full frontal view; see preceding section 

for details) 
WOT: width of ocellar triangle (including lateral ocelli) 
X : times 

Genus PSEUDISOBEACHIUM Kieffer 

Isobrachium Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 35-40. [Kec Forster, 

1856; misidentifieation.] 
Pseudisobrachium Kieffer, 1904, Ami. Mus. Civ. Stor. Nat. Genova, (3a) 1: 

368. [Type species P. laiiceps Kieffer ($, Bolivia); designated by 

Kieffer, 1906, in Andre, Spec. Hym. Eur., 9: 297.] 
Monepyris Kieffer, 1905, Ann. Sci. Roe. Bruxelles, 29: 101, 124. [Type 

species Epyris lialidayi Westwood (—subcyaneum Haliday) ($, 

Europe) ; monobasic.] Placed in synonymy by Kieffer, 1906, op. cit., 

p. 297. 
Plutobethylus Kieffer, 1910, Ann. Soc. Ent, France, 79: 51. [Type species 

P. distans Kieffer ($, Peru); original designation.] New synonymy. 
Lyssepyris, Kieffer, 1913, Boll. Lab. Zool. Portici, 7: 108. [Type species 

Holcpyris flavicornis Kieffer ( S , Nicaragua); monobasic] New 

synonymy. 
Xuntepyris Kieffer, 1913, Boll. Lab. Zool. Portici, 7: 108. [Type species 

Epyris flaviventris Kieffer (S, Texas); monobasic] New synonymy. 
Lissepyris Kieffer, 1914, Das Tierreich, 41: 236. Error for Lyssepyris. 
Xanthepyris Kieffer, 1914, Das Tierreich, 41 : 417. Correction of typographi- 
cal error. 
Parisobra chiu m Kieffer, 1914, Das Tierreich, 41: 424. [Type species 

Rhabdepyris (?) albipes Kieffer {$, Paraguay); monobasic] New 

synonymy. 
Generic characters. — Male. Mandibles with four or five 
(rarely three) teeth; clypeus with a median carina, usually trun- 
cate but occasionally dentate or emarginate apieally; eyes with 
abundant short hair ; antennae simple, 13-segmented, the flagellar 
pubescence appressed or suberect, never erect and bristling. Pro- 
notum without transverse grooves or carina; scutellum with a 
basal transverse groove and with lateral fovae ; propodeum with 



EVANS : REVISION OF PSEUDISOBRACHIUM 223 

a single median carina (rarely obsolete) and without a carina 
margining the dorsal surface posteriorly. Tarsal claws with a 
single tooth which may be very weak or occasionally absent. 
Wings with basal vein reaching subcosta well basad of stigma; 
discoidal vein present or absent, discoidal cell occasionally fully 
outlined by Aveak veins. Genitalia with parameres deeply divided 
into two separate arms ; basis volsellaris with vannus present (ex- 
cept rarely) ; aedoeagus simple and depressed (with rare ex- 
ceptions). Female. Mandibles with three or four teeth; clypeus 
with a median carina, truncate or emarginate apically ; eyes 
each consisting of a single facet, sometimes indistinct; ocelli 
absent ; head longer than broad ; antennae short, 13-segmented, 
flagellum somewhat thickened. Wings and tegulae absent; pro- 
notum longer than broad ; mesonotum subtriangulate, subacute 
behind ; propodeum gradually narrowed anteriorly to a pair of 
points which flank the posterior point of the mesonotum, the 
thorax much constricted laterally at the junction of the propo- 
deum and mesonotum ; mesopleura large, bulging laterally ; mid- 
dle tibiae stout and strongly spined, middle tarsi also somewhat 
spinose, legs otherwise short-haired but not spinose. 

Remarks. I have seen no specimens of the type species of 
Pseudisobrachium , but have no reason to question that it belongs 
to the genus as here understood. Through the courtesy of Dr. 
G. Steinbach of the Zoologisches Museum der Humboldt-Uni- 
versitat zu Berlin, I have had an opportunity to study the type 
specimen of the type species of Pluiobethylus Kieffer. This 
species is a typical Pseudisobrachium in every respect other than 
the tarsal claws, which are simple as indicated by Kieffer. Since 
the tooth on the tarsal claws is very minute and difficult to see 
in some Nearctic species, it seems to me a mistake to emphasize 
this character to that extent. 

With the aid of Dr. R. L. Doutt, of the University of Cali- 
fornia at Albany, T have been able to borrow for study the 
type specimen of flavicornis Kieffer, type of the genus Lyssepyris 
Kieffer. This species is a perfectly typical Pseudisobrachium 
and in fact will run to that genus in Kieffer 's key to genera. In 
his original description of flavicornis, Kieffer states clearly that 
the scutellum has a transverse furrow at the base. But in order 
to properly run out Lyssepyris in his key, one has to assume the 
scutellum has two pits at the base ! 

Xanthepyris Kieffer, based on flaviventris Kieffer, from Texas, 
is also a synonym of Pseudisobrachium. In this instance I am 



224 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

indebted to Mr. Karl-Johan Heqvist of the Naturhistoriska Riks- 
museet of Stockholm for sending me the type of flaviventris. 
Xanthepyris was described as having two unconnected pits at 
the base of the scutellum. The type of flaviventris is pinned 
through the mesonotum in such a way that the structure of the 
base of the scutellum is difficult to observe. However, the species 
is a not uncommon one and in every way a typical Pseudiso- 
brachium; in other specimens the scutellum can be seen to have a 
transverse groove at the base. 

Parisobrachium Kieffer, 1914, based on a single species from 
Paraguay, is said to differ from Pseudisobrachium in lacking a 
discoidal vein and in having three propodeal carinae (although 
Kieffer for some reason places the genus in the Epyrini rather 
than the Pristocerini) . Dr. Delf a Guiglia, of the Museo Civico di 
Storia Naturale, Genoa, Italy, has been kind enough to send me 
a cotype of the type species, albipes, and I find it to be a rather 
typical Pseudisobrachium. Study of the North American species 
shows that the presence or absence of a discoidal vein is of little 
significance in this complex, and the propodeum of albipes differs 
scarcely at all from that of several other species. 

Protisobrachium Benoit, 1957, based on males of two species 
from the Congo, is said to differ from Pseudisobrachium in lack- 
ing a discoidal vein, while Afrisobrachiuin Benoit, 1957, also 
based on males of two Congo species, is said to differ in having the 
basal vein meeting the tip of the pterostigma. I am inclined to 
question the generic value of these characters, but in the absence 
of material I prefer to make no definite disposition of these names. 

KEY TO SPECIES OF PSEUDISOBRACHIUM 

Males 

1. Mandibles with five teeth, the basal three teeth similar in size and 

shape, or sometimes the basal tooth slightly thicker and more rounded 

than the third and fourth teeth (Figs. 7-18) 2 

Mandibles with four teeth, rarely with three, occasionally with five, 
in the latter case the basal tooth broad and curving gradually into the 
inner mandibular margin and the third and fourth teeth very small 
(Figs. 19-32) 24 

2. Abdomen with a distinct, slender petiole (Fig. 68); discoidal vein 

arising far down on the transverse median vein (Fig. 52); vannus 
of genitalia wholly wanting (Fig. 65) (Panama) 

43. petiolatuvi n. sp. 

Abdomen sessile; discoidal vein arising near top of transverse median 
vein, or absent; vannus present, with radiating ridges (Figs. 63-64) 

3 



EVANS: REVISION OF PSELTDISOBRACIIIirM 225 

3. Relatively robust species, propodeum short, measuring less than 1.5 

times as long as broad, in lateral view less than 2.5 times as long as 

high 4 

Very slender species, propodeum in particular very elongate, measuring 
at least 1.5 times as long as broad, in lateral view at least 2.5 times 
as long as high (nccidentale group) 14 

4. Mesopleural callus convex, strongly polished, non-alutaceous or nearly 

so, contrasting to the remainder of the mesopleurum, which is alu- 

taceous and/or punctate 5 

Mesopleural callus alutaeeous, not especially prominent or strongly 
contrasting to the remainder of the mesopleurum, which is also alu- 
taeeous (in one species the whole mesopleurum is convex and shining, 
the callus not well differentiated) (obscurum group) 18 

5. Aedoeagus compressed, of complex structure (Fig. 67) ; mesonotum, 

anteriorly between the notauli, very strongly polished, with only a 

few punctures; propodeum short, rugose (eastern U. S.) 

44. anomalum n. sp. 

Aedoeagus depressed, simple (Fig. 63); other characters not entirely 
as above (crassum group) 6 

6. Front strongly shining, barely if at all alutaeeous, with very strong 

punctures 7 

Front weakly to moderately shining, alutaeeous, punctures not es- 
pecially strong 11 

7. Ocelli very large, DAO at least .25 X WF, OOL not much if any more 

than half WOT; wings very pale; abdomen rufous (Texas) 

5. crassum n. sp. 

Ocelli smaller, DAO less than .2 X WF, OOL subequal to or greater 
than WOT ; wing veins and abdomen dark 8 

8. Antennae medium to dark brown; notauli strongly impressed for most 

of length of mesoscutum 9 

Antennae light reddish-brown; notauli impressed on anterior half of 
mesoscutum only 10 

9. Mandibles broad apically and with the teeth large and subequal (Fig. 

9) ; eyes large, bulging laterally, removed from vertex crest by a 
distance slightly less than their own height; WF 1.27 X HE (Central 
Mexico) 1. michencri n. sp. 

Mandibles with the basal teeth smaller and more rounded than the 
apical tooth (Fig. 10) ; eyes small, not bulging, removed from vertex 
crest by a distance greater than their own height; WF 1.6 X HE 

(central and southern Mexico) 2. blomi n. sp. 

10. Ocelli of moderate size, in a fairly broad triangle (OOL about .9 X 
WOT) ; legs (except front coxae) bright straw-yellow (southern 
Mexico) 3. perpunctatum n. sp. 

Ocelli small, in a compact triangle far removed from the eyes (OOL 
1.3-1.45 X WOT) ; legs brown, paler apically (Mexico and Guate- 
mala) 4. dalmati n. sp. 



226 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

11. Clypeus unusually broad, its lateral angles slightly produced (Fig. 43) ; 

propodeal disc shining, with rather strong transverse striae (Panama) 

9. clypeatum n. sp. 

Clypeus with its lateral angles obtuse, not produced; propodeal disc 
less strongly shining and with less definite striae, if any 12 

12. Clypeus with a median tooth (Fig. 47) ; basal three teeth of mandibles 

small, forming a unit which is somewhat set off from the other 
teeth (Fig. 12) ; propodeum with transverse striae (Costa Rica) .... 

8. cooperi n. sp. 

Clypeus without a median tooth ; mandibular dentition not as above ; 
sculpturing of propodeum weaker and with little tendency to form 
striae 13 

13. Clypeus wide at the base, its sides strongly converging (Fig. 44) ; ocelli 

far removed from eyes (OOL 1.56-1.75 X WOT) ; antennae shorter 
(segment eleven less than twice as long as thick) (Panama and Costa 

Rica) 7. rettenmeyeri n. sp. 

Clypeus narrower basally, its sides less strongly convergent (Fig. 48) ; 
ocelli less far removed from eyes (OOL 1.0-1.3 X WOT) ; antennae 
longer (segment eleven at least twice as long as thick) (Texas) . 
6. texanum n. sp. 

14. Front very broad in relation to eye height (WF 1.4-1.56 X HE) ; ocelli 

small, far removed from eyes (OOL 1.1-1.4 X WOT) ; antennae 
rather short, segment eleven less than 1.4 X as long as thick 

(central Mexico) 14. brunneum n. sp. 

Front narrower (WF .90-1.5 X HE) ; ocelli not so far removed from 
eyes (OOL .43-1.12 X WOT) ; in specimens having the front rather 
broad and the ocelli fairly far removed from the eyes (some speci- 
mens of ooeidentale) the antennae are much longer, segment eleven 
at least 1.5 X as long as thick 15 

15. Front angle of ocellar triangle slightly greater than a right angle ; 

antennae short, segment eleven not more than 1.5 X as long as thick ; 

very small, maximum wing length 2.1 mm. (Texas) 

12. mattheiosi n. sp. 

Front angle of ocellar triangle slightly less than a right angle ; antennae 
longer, segment eleven from 1.4-2.6 X as long as thick ; length of 
fore wing 2.1 to 5.0 mm 16 

16. Front very narrow, WF less than HE (.90-.95 X HE) ; body castane- 

ous ; distance from eye tops to vertex crest equal to less than half HE 

(southern California) 11. castaneum n. sp. 

Front somewhat broader, WF subequal to or greater than HE ; body 
dark brownish-fuscous; distance from eye tops to vertex crest equal 
to more than half HE 17 

17. Larger, minimum wing length 2.8 mm.; front of moderate breadth 

(WF 1.13-1.53 X HE) (California and Arizona) 

10. occidentale n. sp. 



EVANS : REVISION OF PSEUDISOBRACHIUM 227 

Smaller, maximum wing length 2.7 mm.; front narrow (WF .97-1.42 
X HE) (central Mexico) 13. nigriculum n. sp. 

18. Anterior part of mesopleurum with many strong punctures; wing veins 

and stigma brown, discoidal vein present and weakly pigmented; fore 

wing at least 2.9 mm. long 19 

Anterior part of mesopleurum without strong punctures; wing veins 
light brown to colorless; minute species, LFW 1.6-3.1 mm 20 

19. Antennae and legs brown; eyes small, removed from vertex crest by a 

distance about equal to their own height ; OOL much exceeding WOT 

(Mexico) 15. michoacanum n. sp. 

Antennae ferruginous, legs bright castaneous; eyes larger, removed 
from vertex by a distance much less than their own height; OOL 
slightly if at all exceeding WOT (South Carolina and Florida) .... 
16. carolinianum n. sp. 

20. Front broad and eyes small (WF at least 1.4 X HE); ocelli very 

small (DAO about .15 X WF) 21 

Front narrow and eyes large (WF not over 1.3 X HE) ; ocelli enlarged 
at least slightly (DAO .18-.26 X WF) 23 

21. Thoracic dorsum and pleura strongly shining, very weakly alutaceous; 

legs and antennae dark brown (New Mexico) 

17. gibbosum n. sp. 

Thoracic dorsum and pleura alutaceous, weakly to moderately shining; 
legs and antennae yellowish or pale castaneous 22 

22. OOL and WOT subequal; discoidal vein of fore wing unpigmented 

(Arizona) 18. otiosum n. sp. 

OOL much greater than WOT; discoidal vein of fore wing present as 
a pigmented streak (Nicaragua) 19. testaceipes Kieffer 

23. Ocelli in a close triangle, front angle less than a right angle; front 

virtually impunctate (Arizona) 20. pallidum n. sp. 

Ocelli in a broad triangle, front angle greater than a right angle ; 
front with small though usually quite distinct punctures (W. Texas 
and Arizona to Chihuahua and Baja California) 

21. obscurum n. sp. 

24. Mandibles with five teeth, though the third and fourth teeth are very 

small and may be partially fused (Figs. 19-21) 25 

Mandibles with only three or four teeth 28 

25. Discoidal vein of fore wing present, well pigmented; anterior part of 

mesopleurum strongly punctate ; large species with relatively long 
antennae (segment eleven at least 1.5 X as long as thick {prolonga- 

tum group ) 26 

Discoidal vein of fore wing absent or at least unpigmented ; mesopleu- 
rum obscurely punctate; minute, deserticolous species with shorter 
antennae than above 20 



228 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

26. Front relatively broad and eyes relatively small (WF 1.5-1.8 X HE, 

.66-.73 X WH) ; ocelli small (DAO not over .15 X WF) (southern 
Canada and northern United States, south in Appalachians to Caro- 

linas) 24. prolongatum (Provaneher) 

Front relatively narrow and eyes relatively large (WF .90-1.25 X HE, 
.52-.62 X WH) ; ocelli somewhat enlarged (DAO at least .18 X WF) 
27 

27. Mesoscutum strongly shining, non-alutaceous, punctures small but very 

distinct, more crowded laterally; ocelli very large, anterior ocellus 

measuring nearly .2 mm. in diameter (central Mexico) 

22. aztccum n. sp. 

Mesoscutum moderately shining, alutaceous at least on the sides, punc- 
tures present but less pronounced than above ; ocelli of moderate size, 
anterior ocellus less than .15 mm. in diameter (eastern United States) 
23. arenarhim n. sp. 

28. Mandibles slender and with only three teeth, the basal tooth large, 

somewhat rounded (Fig. 32) ; propodeum with some weak longi- 
tudinal ridges medio-basally, but without a single distinct median 

carina (Panama) 46. svperbum n. sp. 

Mandibles with four teeth, third tooth often rather small, very rarely 
absent, but when so basal tooth not as above (Figs. 22-31); pro- 
podeum with a distinct median carina 29 

29. Species with small eyes and ocelli (WF more than 1.4 X HE; ocelli 

not or only slightly enlarged) (Figs. 1-4) ; vertex elevated far above 
eye tops, distance from eye tops to vertex crest commonly subequal 

to or greater than eye height {carbonarium group) 30 3 

Species with relatively large eyes, ocelli also usually enlarged (WF less 
than 1.4 X HE) (Figs. 5-6) ; vertex elevated above eye tops a distance 
usually considerably less than eye height (rufiventre group) . . . 41 s 

30. Front and thoracic dorsum and pleura (including the weakly defined 

mesopleural callus) so strongly and regularly alutaceous as to appear 
beaded, punctures barely if at all evident; OOL. at least nearly 

1.5 WOT (eastern United States) 25. carbonarium (Ashmead) 

Front and thoracic dorsum and pleura not beaded in appearance, 
merely somewhat alutaceous and often punctate 31 

31. Elongate species, antennae quite long and slender (segment eleven at 

least 1.4 X as long as thick) ; either the propodeum is unusually 
elongate (over 1.5 X as long as wide) or the mesopleurum has large 

punctures anteriorly 32 

More compact species, antennae only occasionally with segment eleven 
as much as 1.4 X as long as thick, if so other characters not as 
above 33 

3 In doubtful cases rely upon a oareful determination of the WF/HE ratio ; species 
in which the individual variation embraces the ratio 1.4 will key out on both 
sides of this couplet. 



EVANS : REVISION OF PSEUDISOBRACHIUM 229 

32. Eyes bulging, head narrowed behind eyes and vertex rather narrowly 

rounded (Fig. 2); OOL and WOT subequal; mesopleurum without 

well-defined punctures (California) 31. persimile n. sp. 

Eyes not bulging, head wide behind eyes and vertex very broadly 
rounded; OOL much exceeding WOT; mesopleurum with some coarse 
punctures anteriorly (Washington and British Columbia east to 

Maritimes and south to North Carolina) 

24. prolongatum (Provaneher) 

33. All four mandibular teeth projecting and tooth-like, third and fourth 

teeth subequal or fourth somewhat the larger (Figs. 24-26); small 
species occurring in southwestern United States, Mexico, and Central 

America 34 

Basal tooth of mandibles very different from the small (rarely absent) 
third tooth, broad and with its inner margin arching into the inner 
mandibular margin (Figs. 23, 27, 28) 36 

34. Front obscurely if at all punctate; eyes bulging, vertex narrowly 

rounded off far above eye tops (Fig. 4) ; LFW 1.3-2.0 mm 

28. minutissimum n. sp. 

Front with shallow but rather distinct punctures; eyes less bulging 
and vertex much more broadly rounded; LFW 2.3-3.0 mm 35 

35. Legs (except front coxae) light yellowish-brown; wing veins light 

brown to almost colorless; OOL subequal to WOT (Arizona) 

30. navajo n. sp. 

Legs brown, wing veins brown; OOL much exceeding WOT (Arizona 
and New Mexico) 27. minimum, n. sp. 

36. Antennae relatively smooth (as, for example, in flavinervis), setulae 

on fiagellum pale, minute, subappressed, erect setae on flagellum short 

and inconspicuous; oeellar triangle rather broad 37 

Antennae roughly pubescent (as, for example, in prolongation) , flagel- 
lar setulae suberect, some of them .3 X as long as width of fiagellum, 
erect setae numerous and some of them usually half as long as width 
of flagellum 39 

37. Ocelli small (DAO .12-.15 X WF) ; front very broad (WF at least about 

1.5 X HE) (Fig. 3); clypeus rather broadly truncate apieally (Fig. 

50) ; OOL at least equal to WOT 38 

Ocelli somewhat enlarged (DAO at least .19 X WF) ; front generally 
narrower and clypeus more narrowly truncate than above; OOL much 
less than WOT 48 

38. WF at least 1.75 X HE; vertex elevated above eye tops a distance 

about equal to eye height (Fig. 3) (Durango) 32. hurdi n. sp. 

WF not more than about 1.5 X HE; vertex elevated above eye tops 

a distance equal to about .8 X HE (New Mexico) 

33. Icrombeini n. sp. 

39. Ocelli small (DAO not more than .16 X WF), anterior ocellus situated 

above an imaginary line drawn between eye tops (Fig. 1); discoidal 
vein of fore wing present, pigmented for a short distance (Fig. 58) 
40 



230 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Ocelli somewhat enlarged and anterior ocellus situated on or touching 
an imaginary line drawn between eye tops (Fig. 5), or if not, then 
the discoidal vein is absent (Fig. 60) 43 

40. Mesopleural callus shining, barely alutaceous, anterior part of meso- 

pleurum with coarse punctures (Nicaragua and Honduras) 

29. flavicornis (Kieffer) 

Mesopleural callus alutaceous, slightly convex but not otherwise notably 
contrasting to remainder of mesopleurum, which is weakly to mod- 
erately punctate (eastern United States) 26. ashmeadi n. sp. 

41. Front angle of ocellar triangle (as measured from outer sides of 

ocelli) less than a right angle (as in Figs. 4, 5) 42 

Ocelli in a broad triangle, front angle greater than a right angle 
(as in Fig. 6) 48 

42. Thoracic dorsum strongly polished, mostly if not entirely non-alutaceous, 

contrasting with the front, which is distinctly alutaceous; mesopleu- 
rum also strongly shining; mesoscutum with small, sharply defined, 

rather evenly spaced punctures (Arizona) 38. apache n. sp. 

Thoracic dorsum not wholly strongly polished, always in large part 
alutaceous, not strongly contrasting to front 43 

43. Fore wing with discoidal vein present as a distinctly pigmented streak 

(Fig. 59) ; wings with setulae dark, veins brown 44 

Fore wing with discoidal vein absent or merely indicated by an un- 
pigmented line (Figs. 60-61) ; wings pale, veins light brown to color- 
less 46 

44. A minute species, LFW 1.6-2.5 mm.; legs and antennae brownish; an- 

tennae of moderate length, segment eleven about 1.2 X as long as 
thick; pubescence of flagellum unusually coarse and suberect (Louisi- 
ana and Arkansas) 36. pusilhim n. sp. 

Larger, LFW 2.1-3.7 mm. ; legs and antennae light yellowish or reddish 
brown, or if darker brown (some specimens of comanche) antennae 
longer than above and flagellar pubescence less coarse 45 

45. Vertex narrowly rounded, eyes strongly bulging laterally; an elongate 

species, propodeum, in lateral view, much more than twice as long as 
high, thorax, in lateral view, more than 3 X as long as its maximum 

height (Arizona) 37. comanche n. sp. 

Vertex more broadly rounded, eyes less strongly bulging (Fig. 5) ; body 
compact, propodeum about twice as long as high, thorax less than 

3 X as long as its maximum height (eastern United States) 

35. rufiventre (Ashmead) 

46. Mesopleural callus well differentiated, convex and polished, remainder 

of mesopleurum somewhat shining and/or punctate; antennae short 
(segment eleven 0.9-1.2 X as long as thick) ; vertex broad, almost 
straight across (eastern United States) ... .34. flaviventre (Kieffer) 
Mesopleural callus weakly differentiated, not strongly convex and 
polished; antennae of variable length, often longer than above . . .47 



EVANS : REVISION OF PSEUDISOBRACHIUM 231 

47. Clypeus emarginate apieally (Fig. 49) ; ocelli not much enlarged (DAO 

about .17 X WF) ; body dark brown to black (Texas) 

40. cmarginatum n. sp. 

Clypeus truncate apieally (Fig. 45); ocelli usually larger than above 
(Texas to California) 39. foutsi n. sp. 

48. Front with small but usually very clearly defined punctures; eyes rela- 

tively smooth, the setae being shorter and sparser than is usual in 
this genus; wing veins nearly colorless; antennae various shades of 
yellow or light castaneous, very smooth (western Texas to California 

and Baja California) 42. flavinervis Fouts 

Front obscurely if at all punctate; eyes about as hairy as usual in the 
genus; wing veins light brown (Nebraska to eastern Texas and 
Mexico ) 49 

49. Ocelli of moderate size (DAO .19-.24 X WF) ; abdomen brown like 

head and thorax (Nebraska to Nuevo Leon) 41. rectangulatum n. sp. 
Ocelli extremely large (DAO .33-.37 X WF) ; abdomen rufous, con- 
trasting to head and thorax (lower Rio Grande Valley of Texas) 
43. macrops n. sp. 

Females 

1. Mandibles with four well defined teeth (Figs. 33-36) 2 

Mandibles with three teeth, sometimes with only two large, well-defined 

teeth, sometimes with a rudimentary fourth tooth indicated by an 
undulation on the inner mandibular margin (Figs. 37-42) 8 

2. Basal tooth of mandibles large and projecting (Figs. 33-34) ; entire 

body dark brown to almost black 3 

Basal two teeth of mandibles small and drawn back along inner margin 
of mandibles (Figs. 35-36) ; abdomen, at least, pale 4 

3. Very large, length of head much over 1 mm.; median carina of clypeus 

continued as a short apical tooth; head 1.15 X as long as wide, 

occipital carina obsolete dorsally (Panama) 47. gigas n. sp. 

Smaller, head length well under 1 mm.; clypeus without a median 
apical tooth; head 1.3 X as long as wide, occipital carina complete 
dorsally (Panama) 48. zeteM n. sp. 

4. Head and thorax nearly black, eye spots contrastingly pale, abdomen 

contrastingly rufous; entire head with strong, rather evenly spaced 
punctures, between which it is strongly polished (Guatemala) .... 

49. manni n. sp. 

Head and thorax pale to medium brown, only slightly if at all darker 
than abdomen, eyes less strongly contrasting to head color 5 

5. Anterior part of front and sides of head alutaceous and/or striato- 

punctate ; generally larger than below, head length .6 mm. or more . . 6 

Head not at all striato-punctate and not or barely alutaceous, strongly 

shining ; minute species, head length .4-.63 mm 7 



232 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

G. Head slightly, gradually narrowed behind, anterior part of front 
alutaceous but not or only slightly striato-punctate ; pronotal disc 
elongate, about 1.6 X as long as its posterior width (Utah, Califor- 
nia) 50. paucipunctatum Fonts 

Head parallel-sided or very slightly bulged near the middle, anterior 
part distinctly striato-punctate laterally and dorsally ; pronotal disc 
about 1.4 X as long as its posterior width (California, Arizona.) .... 
10. ? occidentale n. sp. 

7. An unusually slender species, head 1.42 X as long as wide, mesonotuni 

about twice as long as wide; head punctures separated, for the most 

part, by about twice their own diameters (West Texas) 

12. ? matthewsi n. sp. 

A somewhat stockier species, head 1.35 X as long as wide, mesonotuni 
1.4 X as long as wide; head punctures separated, for the most part, 

by about their own diameters (southwestern United States) 

21. ? obscurum n. sp. 

8. Basal tooth of mandibles large, strongly projecting (Fig. 42); clypeal 

carina slightly produced beyond apical margin, which is concave; 
head and thorax deep brown, eyes white (Costa Eica) . 

51. costaricanum n. sp. 

Basal tooth of mandibles small, drawn back along inner margin of 
mandible ; other characters not as above 9 

9. Head strongly striato-punctate, at least antero-laterally ; head parallel- 

sided or sides weakly convex, LH 1.1-1.3 X WH; generally larger 
and darker species, LH usually over .8 mm. (rarely down to .68 mm.) 

10 

Head not or very weakly striato-punctate (if rather evidently so, LH 
more than 1.3 X WH and sides of head weakly, gradually convergent 
behind) ; generally smaller species of paler coloration, LH not over 
.8 mm 11 

10. Disc of propodeum strongly shining, barely if at all alutaceous, punc- 

tures small and confined to extreme sides; head 1.2-1.3 X as long 
as wide (southern Canada and northern United States, south to 

North Carolina) 24. prolongation (Provancher) 

Disc of propodeum moderately shining, evidently alutaceous, punctures 
on sides fairly strong, punctate parts of disc equal in area to impunc- 
tate median band; head about 1.15 X as long as wide (eastern United 
States 23. ? arenarium n. sp. 

11. Punctures of front very numerous, separated at least in part by no 

more than their own diameters (except medially), anteriorly de- 
cidedly crowded; front usually somewhat alutaceous, at least in part, 

under side of head often rather strongly alutaceous 12 

Punctures of front sparse, separated for the most part by more than 
their own diameters; front strongly shining, not at all alutaceous, 
under side of head somewhat alutaceous or not 16 



EVANS : REVISION OF PSEUDISOBRACHIUM 233 

12. Length of head not over .(> nun.; front with punctures somewhat less 

crowded than below 13 

Length of head generally over .0 mm. ; front usually somewhat darker 
and with punctures relatively more crowded than above 14 

13. Front shining, weakly alutaceous, with punctures somewhat more evenly 

distributed than below; LII 1.22-1.35 X WH (eastern United States) 

34. ? flaviventre (Kieffer ) 

Front conspicuously alutaceous, in fact obscurely striate anteriorly, 
punctures rather sparse posteriorly and much more crowded anter- 
iorly; LH 1.4 X WH (southwestern United States) 

42. ? flavinervis Fouts 

14. Head rather elongate, LH 1.3-1.4 X WH, sides of head weakly, gradu- 

ally convergent behind (eastern United States) 

25. ? carbonarium n. sp. 

LH 1.1-1.25 X WH 15 

15. Head yellowish-brown; front unusually flat, punctures relatively weak 

(Pacific states) 31. ? persimile n. sp. 

Head ferrugino-castaneous; front not especially flat, punctures strong 
(eastern United States) 35. rufiventre (Ashmead) 

10. Punctures of front large and distinct; LH .47-. 53 mm. (eastern United 

States) 26. ? ashmeadi n. sp. 

Punctures of head very small and shallow; LH about .4 nun. (Arkansas) 
36. '? pusillum n. sp. 

Crassum Species-group 

The males of this group have a relatively compact body form, 
the propodeum being distinctly shorter and higher than in the 
occidentale group. The most distinctive feature of the group 
is the convex, prominent, shining and non-alutaceous mesopleu- 
ral callus, which contrasts strongly with the remainder of this 
sclerite. The mandibles are five-toothed ; in some species the basal 
tooth is somewhat thicker and more rounded than the two fol- 
lowing teeth, approaching the condition in the prolongatum 
group. The group falls readily into two subgroups: in one the 
head and thoracic dorsum are shining and strongly punctate ; in 
the other these areas are dull, alutaceous, and weakly if at all 
punctate. Each of these subgroups has several species in Central 
America and Mexico and a single species in southern Texas. The 
compact form and relatively large size of species of this group, 
the full mandibular dentition, the variation in the shape of 
the clypeus, the complete notauli of some species, and other char- 
acters suggest that this may be the most primitive of the several 
species groups (i.e., closest to related genera such as Pristoccra, 
Propristocera, and deist epyris). 



234 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



No females can presently be assigned to this group with any 
real certainty. I would expect the females to have relatively 
broad mandibles, probably with four strong teeth. Two species 
described later in the paper, gigas and zeteki, have mandibles of 
this type and may represent females of this group. Another 
species, manni, may or may not go with this group. 



Species and locality 



No. 



TABLE 
WF/HE 



00L/W0T 



DAO/WF 



Ant. 11 L/W 



Brownsville, Texas 
San Antonio, Texas 


3.9(3 
3.3 


7-4.2) 


l.H (1 
1.32 


0-1 


2) 


.45 (. 
.62 


42- 


50) 


.29 (.27- 
.25 


.31) 


1.5(1.4-1 

1.4 


.7) 


michenerl 




























Teziutlan, Puebla 


4.9 




1.27 






1.32 






.17 






2.0 




perpunctatum 




























Tehuantepec, Oaxaca 


3.1 




1.23 






.92 






.18 






1.6 




blomi 




























San Cristobal, Chiapas 
Cuernavaca, Morelos 


5.3 
! 4.3(3 


7-4.9) 


1.6 

1.57(1 


54- 


.6) 


1.70 

1.81 (1.62-2.0) 


.13 
.11 






2.2 
2.1 




dalmatl 




























Yepocapa, Guatemala 
"Mexico" 


4.0 
4.9 




1.5 
1.54 






1.45 
1.32 






.12 
.14 






1.6 
2.1 




cooperl 




























Turlalba, Costa Rica 


2.7 




1.52 






1.83 






.12 






1.9 




clypeatum 




























Canal Zone 


3.1 




1.34 






1.60 






.13 






2.0 




rettenmeyeri 




























Canal Zone 
Costa Rica 


! 2.6(2.2-2.9) 
2.5 


1.47(1 
1.46 


.43- 


.5) 


1.66(1 
1.70 


.56- 


1.75) 


.13( 
.11 


11- 


.15) 


1.7(1.6- 
1.7 


.8) 


texanum 





























Brownsville, Texas 
Harlinqen, Texas 
San Antonio, Texas 
Wharton, Texas 



2.9 1.40 1.00 .16 2.0 

3.4(2.5-3.9) 1.41(1.35-1.44) 1.22(1.18-1.27) .15(.15-.16) 2.3(2.2-2.5) 

3.3(3.2-3.3) 1.41(1.38-1.44) 1.07(1.05-1.11) .16(.15-.17) 2.2(2.0-2.5) 

3.6 1.32 1.00 .16 2.3 



1. Pseudisobrachium micheneri new species 

Holotype. — $ , 5 mi. NE of Teziutlan, Puebla, Mexico, 4700 
feet, 27 June 1953 (Univ. Kansas Mex. Exped.) [KU]. 

Description. — Length 5.5 mm.; LEW 4.9 mm. Head and 
thorax black, abdomen dark brown, slightly paler on the sides 
of the first two segments, apical third of mandibles light brown, 
teeth rufous ; antennae medium brown ; legs brown, becoming 
paler apically ; wings lightly infuscated, stigma dark brown, 
veins brown. Mandibles broad apically, with five strong, sharp 
teeth (Fig. 9). Clypeus truncate apically, its median carina 
weakly arched in profile. Antennae elongate, first four segments 
in a ratio of about 26:8:18:15, segment eleven twice as long as 
thick ; pubescence of flagellum coarse, setulae almost half as long 
as thickness of segments, erect setae nearly as long as width of 
flagellum. WF .61 X WH, 1.27 X HE; ocelli of moderate size, 



EVANS : REVISION OP PSEUDISOBRACHIUM 235 

in a small, compact triangle far removed from eyes; DAO .17 
X WP; OOL 1.32 X AVOT. Eyes large, prominently bulging 
from sides of head, removed from vertex crest by a distance 
nearly equal to their height. Front shining, toward vertex a 
little less strongly shining and more evidently alutaceous, over 
whole surface with large punctures which for the most part are 
separated by less than their own diameters. Pronotum with 
rough, reticulate sculpturing except along the posterior margin, 
which is smooth and shining. Mesoscutum shining, with strong, 
close punctures ; notauli deeply impressed for full length of meso- 
scutum; scutellum punctate except in extreme center, basal 
groove broad and shallow, lateral foveae small, shallow. Pro- 
podeum 1.45 X as long as broad, in lateral view about 2.3 X as 
long as high ; spiracles elongate-elliptical, directed dorsad ; entire 
surface, including sides, with strong reticulate sculpturing ; 
median carina strong, extending full length of dorsal surface. 
Mesopleural callus strongly convex, smooth and shining, with a 
few punctures anteriorly; remainder of mesopleurum with 
strong, close-set punctures. Discoidal vein of fore wing strong, in 
fact the entire discoidal cell outlined by faintly pigmented lines. 

Remarks. — Although this species is known from but one 
specimen, this specimen is distinctive in so many ways that there 
is little question that it is specifically distinct from its relatives 
crassum and blomi. The latter species is apparently diurnal, the 
former nocturnal. The ocelli of micheneri are slightly larger than 
those of blomi but much smaller than those of crassum; the wing 
veins and membrane are pigmented as in blomi. In general, I 
would expect micheneri to be a diurnal species, but it may fly in 
shaded situations or in periods of reduced sunlight. The broad 
mandibles, complete notauli, and fully outlined discoidal cell 
suggest the genus Pristocera. 

2. Pseudisobrachium blomi new species 

Holotype. — $ , San Cristobal las Casas, Chiapas, Mexico, 
7500 feet, 28 April 1959 (H. B. Evans) [MCZ, No. 30265]. 

Description. — Length 6.1 mm. ; LFW 5.3 mm. Black, sides of 
base of abdomen suffused with dark reddish-brown ; antennae 
dark brown ; coxae dark brown, legs otherwise medium broAvn ; 
apical two-thirds of mandibles dark ferruginous ; wings lightly 
infuscated, stigma dark brown, veins brown. Mandibles with 
five teeth, basal tooth somewhat rounded (Fig. 10). Clypeus 
truncate apically. Antennae elongate, first four segments in a 



236 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

ratio of about 32 :5 :22:20, segment eleven 2.2 X as long as thick; 
pubescence of flagellum short and dark, erect setae short and 
inconspicuous. WF .71 X WH, 1.6 X HE; ocelli small, in a 
compact triangle far removed from eyes, DAO .14 X WF, OOL 
1.7 X WOT. Eyes relatively small, not notably bulging; distance 
from tops of eyes to vertex crest actually slightly greater than 
eye height. Front strongly shining, barely alutaceous, with 
many small but strong punctures which are separated from one 
another by from 1-2 times their own diameters. Anterior half of 
pronotum somewhat roughened, weakly shining, posterior half 
smoother, punctate, shining. Mesoscutum shining, barely alu- 
taceous, with close-set punctures except along the median line ; 
notauli strong on the anterior two-thirds, reaching posterior 
margin as weak impressions only. Scutellum shining, punctate, 
the basal groove moderately wide, shallow. Propodeum measur- 
ing 1.4 X as long as broad, in lateral view 2.3 X as long as 
high; spiracles elongate-elliptical, directed dorso-laterad ; disc 
with reticulate sculpturing which is strongest anteriorly. Meso- 
pleural callus prominent, shining and impunctate, remainder of 
mesopleurum with large, coarse punctures, the punctures smaller 
posteriorly and nearly absent along posterior margin below 
callus. Fore wing with discoidal vein long and strong, the dis- 
coidal cell in fact faintly outlined by pigmented lines. 

Paratypes. — MEXICO : 1 $ , 4 mi. NW Cuernavaca, Morelos, 
7500 feet, 28 June 1959 (H. E. and M. A. Evans) [CU] ; 1 $ , 
6 mi. N. Cuernavaca, Mor., 7500 feet, 15 Aug. 1954 (J. G. 
Chillcott) [CNC]. 

Variation. — The paratypes are smaller than the type but dif- 
fer only slightly in body measurements (Table I). The mandibles 
are yellowish apically and the legs bright castaneous beyond the 
coxae rather than dull brown as in the type. Since the type and 
paratypes are from widely separated mountain systems, it is not 
surprising that there are differences between them. However, I 
question that any of these differences are of specific value. 

Remarks. — This large and distinctive species is named for 
Franz Blom of San Cristobal las Casas, Chiapas, my host during 
a most profitable stay in that city in April 1959. The type was 
taken on low broad-leaf foliage in a pine forest just west of town 
at about 11 in the morning. The first paratype listed was taken 
at about the same altitude and in precisely the same ecological 
situation ; it was taken at about 10 in the morning. 



EVANS : REVISION OF PSEUDISOBRACHIUM 237 

3. Pseudisobrachium perpunctatum new species 

Holotype. — $ , Tehuantepec, Oaxaca, Mexico, 9 Aug. 1958, 
200 feet (E. G. Matthews) [MCZ, No. 30266]. 

Description. — Length 4.5 mm.; LFW 3.1 mm. Head and 
thorax black, abdomen reddish-brown, somewhat infuscated to- 
ward apex ; antennae castaneous ; legs wholly and uniformly light 
brown, except front coxae piceous ; mandibles castaneous, teeth 
rufous; wings subhyaline, stigma brown, veins light brown. 
Mandibles with five teeth, about as figured for the preceding 
species. Clypeus truncate apically, its median elevation not 
arched in profile. Antennae rather long, first four segments in 
a ratio of about 21 :5 :13 :12, segment eleven 1.6 X as long as 
thick; pubescence of flagellum pale, subappressed, erect setae 
pale, numerous, the longest ones slightly less than half as long 
as width of flagellum. WF .65 X WH, 1.23 X HE ; ocelli some- 
what enlarged, DAO .18 X WF, OOL .9 X WOT. Eyes rather 
large, removed from vertex crest by only .6 their height. Front 
shining, weakly alutaceous, with strong punctures which are 
separated by about or slightly less than their own diameters. 
Pronotum shining, strongly punctate, slightly roughened an- 
teriorly. Mesoscutum strongly punctate except along midline ; 
notauli strong on anterior .6, absent behind ; scutellar disc 
sparsely punctate, basal groove straight, shallow. Propodeum 
measuring 1.35 X as long as broad, in lateral aspect measuring 
2.4 X as long as high ; spiracles elliptical, opening dorso-laterally ; 
median carina short, extending only about half length of disc, 
which is covered with fine, somewhat irregular transverse striae. 
Mesopleural callus convex, non-alutaceous, strongly shining ; re- 
mainder of mesopleurum alutaceous and punctate. Discoidal 
vein of fore wing weak, barely pigmented. 

Remarks. — The one known specimen of this species was col- 
lected in a bare, arid field surrounded by irrigated land in the 
city of Tehuantepec ; it came to the light of a Coleman lantern 
in the early evening. 

4. Pseudisobrachium dalmati new species 

Holotype. — $ [San Pedro] Yepocapa, Guatemala, [Dept. 
Chimaltenango, 4850 feet], May 1948 (H. T. Dalmat) [USNM, 
No. 65151]. 

Description. — Length 5.5 mm.; LFW 4.0 mm. Head and 
thorax black, abdomen dark reddish-brown suffused with lighter 



238 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

brown basally and apically; scape brown, flagellum dull rufo- 
castaneous; front coxae black, legs otherwise medium brown 
except beyond apical two-thirds of tibiae, where they are pale 
yellow -brown ; wings weakly infuscatecl, veins and stigma dark 
brown. Mandibles with five teeth, about as in the preceding 
two species. Clypeus truncate apically, its median elevation 
not arched in profile. Antennae rather short, scape especially 
short, with a ventral excavation which permits a strong elbowing 
of the antennae; first four segments in a ratio of about 
22:6:15:15, segment three only 1.4 X as long as thick, segment 
eleven 1.6 X as long as thick; pubescence of flagellum light 
brown, subappressed, setae numerous, generally slightly less than 
half as long as width of flagellum. WF .68 X WH, 1.5 X HE ; 
ocelli small, in a compact triangle far removed from eyes, DAO 
.12 X WF, OOL 1.45 X WOT. Eyes rather large, removed 
from vertex crest by a distance equal to about .9 their height. 
Front shining, non-alutaceous, with strong punctures which are 
separated from one another by less than their own diameters. 
Pronotum shining, strongly punctate, slightly roughened in 
front. Mesoscutum with the notauli strong on the anterior .6, 
absent behind ; disc strongly punctate except on the median line ; 
scutellar disc sparsely punctate. Propodeum short, in dorsal 
view measuring 1.25 X as long as broad, in lateral view 2.1 X 
as long as high ; spiracles elliptical, directed dorsad ; disc with 
strong reticulate sculpturing anteriorly and laterally, posteriorly 
with finer sculpturing which tends to form weak transverse 
ridges ; median carina short. Mesopleural callus convex, strongly 
shining, non-alutaceous ; remainder of mesopleurum also non- 
alutaceous, but wholly covered with very strong punctures. Fore 
wing with the discoidal vein longer than the basal vein, in fact 
the entire discoidal cell outlined by weakly pigmented veins. 

Paratype. — l i , labeled simply "Mex." [ANSP]. 

Variation. — The single paratype is larger than the type, meas- 
uring about 6.5 mm. long, fore wing 4.9 mm. The abdomen is 
missing beyond the fourth segment. The legs are wholly dark 
except for a light spot at the inner apex of the middle and hind 
tibiae. The antennae are considerably longer than in the type, 
segment eleven measuring 2.1 X as long as broad. WF measures 
1.54 X HE, OOL 1.32 X WOT. In other details the resemblance 
to the type is close. 

Remarks. — I would judge this to be a diurnal species, as 
the ocelli are small and the colors dark. Its relationship to the 
preceding two species is close, but there are enough differences 
so that I feel confident that it is specifically distinct. 



EVANS : REVISION OF PSEUDISOBRACHIUM 239 

5. Pseudisobrachium crassum new species 

Holotype. — S , Brownsville, Texas, Oct. 1942 (at light, E. S. 
Ross) [CAS]. 

Description. — Length 5.2 mm.; LFW 3.7 mm. Head and 
thorax dark brownish-fuscous, nearly black ; abdomen bright red- 
dish-brown ; antennae ferruginous ; coxae brownish, remainder 
of legs straw-yellow ; wings subhyaline, stigma brown, veins 
amber. Mandibles with five teeth, the basal tooth somewhat 
thicker than the two rather small teeth next to it (Fig. 13). 
Apical margin of clypeus truncate ; median carina not arched 
in profile. Antennae of moderate length ; first four segments in 
a ratio of about 20:5:11:10, segment eleven 1.45 X as long as 
thick; pubescence of flagellum fine, pale, subappressed, erect 
setae short and inconspicuous. WF .6 X WH, 1.13 X HE ; ocelli 
very large, DAO .27 X WF ; OOL .5 X WOT ; front angle of 
ocellar triangle slightly less than a right angle. Eyes removed 
from vertex crest bj r slightly more than half HE. Front strongly 
shining, only very weakly alutaceous, with strong punctures 
which are separated from one another by only slightly more than 
their own diameters. Pro- and mesonota also strongly shining 
and barely alutaceous, with strong punctures which are more 
widely separated between the notauli and absent from the center 
of the scutellum ; notauli strongly impressed on anterior .7 of 
mesoscutum, absent behind. Propodeum 1.45 X as long as broad, 
in lateral view about twice as long as high ; disc shining, aluta- 
ceous, median carina extending for only about half length of 
dorsal surface ; spiracles elongate-elliptical, directed dorsad. 
Mesopleural callus strongly shining, non-alutaceous, but with a 
few punctures ; remainder of mesopleurum weakly alutaceous, 
with small but distinct punctures. Discoidal vein of fore wing 
barely pigmented, about as long as basal vein. 

Paratypes. — TEXAS : 1 $ , same data as type [MCZ] ; 1 $ , 
Esperanza Ranch, Brownsville [USNM] ; 1 $ , San Antonio, 
Oct. 1942 (E. S. Ross) [CAS]. 

Variation. — The Brownsville paratypes approximate the type 
in size, but the San Antonio specimen is smaller (length of body 
4.6 mm., of fore wing 3.3 mm). The latter specimen is a little 
less strongly punctate and has a relatively wider front and 
smaller ocelli (Table 1). The specimen from the Esperanza 
Ranch is the most strongly punctate of the lot, and in this speci- 
men the notauli extend for only about half the length of the 
mesoscutum. 



240 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

6. Pseudisobrachium texanum new species 

Holotypc— 6 , Harlingen, Texas, 12 March 1945 (D. E. 
Hardy) [USNM, No. 65150]. 

Description. — ■ Length 5.1 mm. ; LFW 3.6 mm. Head and 
thorax black, abdomen dark brown, paler basally; mandibles 
light brown, teeth rufous; antennae dark brown, coxae dark 
brown, trochanters and femora medium brown, tibiae and tarsi 
light brown ; wings subhyaline, stigma brown, veins light brown. 
Mandibles with five teeth, the three inner teeth subequal. Cly- 
peus narrow, apical margin very slightly concave (Fig. 48). 
Antennae elongate, first four segments in a ratio of about 
23:5:14:13, segment eleven 2.2 X as long as thick; pubescence 
of flagellum pale and subappressed, erect setae short, sparse. 
WF .68 X WH, 1.44 X HE ; ocelli rather small, DAO .15 X WF, 
OOL 1.22 X WOT. Eye tops removed from vertex crest by a 
distance equal to about .6 HE. Front alutaceous, moderately 
shining, with shallow punctures which are separated from one 
another by slightly more than their own diameters. Pro- and 
mesonota alutaceous, moderately shining, punctures numerous 
but small and shallow ; notauli strong on anterior half of meso- 
scutum; basal furrow and lateral foveae of scutellum shallow. 
Propodeum short, only about 1.35 X as long as broad, in lateral 
view about twice as long as high ; spiracles elongate-elliptical ; 
disc with a strong median carina, with reticulate sculpturing 
basally and laterally. Mesopleurum with the callus shining, very 
weakly alutaceous, remainder of mesopleurum with large punc- 
tures. Discoidal vein of fore wing not deeply pigmented, arising 
a short way down on the transverse median vein (Fig. 53). 

Paratypcs. — TEXAS :4 n, same data as type, 12-15 Marcl 
1945 [USNM. MCZ] ; 3 $ S , San Antonio, Aug. 1942 (at light, 
E. S. Ross) [CAS, MCZ] ; 1 $ , Brownsville, Sept. 1942 (E. S. 
Ross) [CAS] ; 1 $ , Wharton, 24 June 1917 (J. C. Bradley) 
[CU]. 

Variation. — Size range is from 3.3 to 5.1 mm., LFW 2.5 to 
3.9 mm. There is little variation in head measurements in the 
type series (Table 1). The Wharton specimen has distinctly 
paler antennae, the flagellum being chestnut-brown. 

7. Pseudisobrachium rettenmeyeri new species 

Holotype. — $ , Barro Colorado Island, Canal Zone, 28 March 
1955 (Carl Rettenmeyer) [KU]. 



EVANS : REVISION OF PSEUDISOBRACHIUM 241 

Description. — Length 4.2 mm. ; fore wing 2.7 mm. Head 
black, thorax dark reddish-brown, abdomen slightly paler, es- 
pecially the basal segments; mandibles yellowish, teeth rufous; 
clypeus light brown ; antennae medium brown, including scape ; 
legs, including front coxae, bright yellowish-brown ; wings lightly 
infuscated, veins and stigma dark brown. Mandibles with five 
teeth, basal tooth slightly stronger than third and fourth teeth. 
Clypeus rather broad basally, sides tapering to a truncate apex 
of moderate width (Fig. 44). Antennae with first four segments 
in a ratio of about 19:4:13:11, segment eleven about 1.6 X as 
long as thick; pubescence of flagellum light brown, coarse, erect 
setae numerous, many of them over half as long as thickness of 
flagellum. WF .7 X WH, 1.48 X HE; ocelli small, DAO 
.13 X WF ; OOL 1.67 X AVOT. Distance from eye tops to vertex 
crest equal to about .8 HE. Front strongly alutaceous, weakly 
shining, punctures small and inconspicuous. Pro- and mesonota 
strongly alutaceous, weakly punctate ; notauli strong on anterior 
.6 of mesoscutum. Propodeum 1.4 X as long as wide, in lateral 
view about twice as long as high; spiracles elliptical, directed 
dorso-laterad ; disc weakly sculptured, the sculpturing showing 
a weak tendency to form transverse striations. Characters of the 
mesopleurum and of the fore wing as described for the following 
species. 

Paratypes. — CANAL ZONE :U, same data as type [KU] ; 
1 $ , Pacora, 13 May 1953 (F. 8. Blanton) [USNM]. COSTA 
RICA: 1 $, Hamburg Farm, April (C. W. Dodge) [MCZ]. 

Variation. — The Pacora specimen is slightly larger than the 
type (LFW 2.9 mm.), the Barro Colorado paratype very small 
(LFW 2.2 mm.). In the Pacora specimen the antennae are light 
brown, the scape yellow-brown. In all other respects the para- 
types closely resemble the type. 
8. Pseudisobrachium cooperi new species 

Holotypc. — $ , Turrialba, Costa Rica, 22 June 1949 (K. W. 
Cooper) [USNM, No. 65152]. 

Description. — Length 3.7 mm.; LFW 2.7 mm. Head and 
thorax black, abdomen dark brown, suffused with lighter brown 
on sides basally ; mandibles yellow, teeth rufous ; antennae brown 
beyond the scape, which is black; front coxae black, legs other- 
wise light brown; wings lightly infuscated, veins and stigma 
dark brown. Mandibles with five teeth, basal three teeth small, 
subequal, forming a unit which is well set off from the apical 
two teeth (Fig. 12). Clypeus with a rounded, polished, median 



242 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

apical prominence (Fig. 47). Antennae slender, first four seg- 
ments in a ratio of about 20:4:11:10, segment eleven 1.9 X as 
long as thick, pubescence of flagellum coarse, semierect, erect 
setae unusually numerous. WF .69 X WH, 1.52 X HE ; ocelli 
small, in a compact triangle far removed from eyes ; DAO .12 X 
WF, OOL 1.83 X WOT. Vertex broadly rounded off far above 
tops of eyes, distance from eye tops to vertex crest nearly equal 
to HE. Front strongly alutaceous, weakly shining, with abund- 
ant shallow and rather inconspicuous punctures. Pro- and 
mesonota also strongly alutaceous and with abundant but shal- 
low punctures ; notauli present as thin lines on anterior .4 of 
mesoscutum. Propodeum about 1.4 X as long as wide, in lateral 
view about twice as long as high ; spiracles elliptical, directed 
dorsad; median carina strong, dorsal surface with ridges which 
are reticulate basally, transverse posteriorly. Mesopleurum with 
the callus shining, barely alutaceous, remainder of this sclerite 
strongly alutaceous and with strong sculpturing. Discoidal vein 
of fore wing darkly pigmented, longer than basal vein, forming 
a straight line with median vein. 

9. PSEUDISOBRACHIUM CLYPEATUM new Species 

Holotype. — S , Barro Colorado Island, Canal Zone, 4 Jan. 
1929 (C. H. Curran) [AMNH]. 

Description. — Length 4.3 mm.; LFW 3.1 mm. Head black, 
thorax piceous, abdomen reddish-brown, sides of the basal seg- 
ments suffused with light yellowish-brown ; mandibles yellow, 
apices rufous ; antennae brown ; coxae, trochanters, and basal 
parts of femora bright straw-yellow ; apices of femora and all 
of tibiae and tarsi yellow-brown ; wings lightly infuscated, veins 
and stigma brown. Mandibles with five teeth, as shown in 
Figure 11. Clypeus broad apically, with a median tooth, the 
lateral angles prominent and slightly reflexed (Fig. 43). An- 
tennae slender, first four segments in a ratio of about 24:5 :15:14, 
segment eleven about twice as long as thick ; pubescence of flagel- 
lum coarse, semierect, erect setae numerous, the longest ones over 
half as long as width of flagellum. WF .65 X WH, 1.34 X HE ; 
ocelli small, far removed from eyes ; DAO .13 X WF ; OOL 1.6 X 
WOT. Vertex broadly rounded off far above the eye tops ; 
distance from eye tops to vertex crest equal to about .8 HE. 
Front moderately shining, strongly alutaceous, with shallow 
punctures which are separated from one another by slightly more 
than their own diameters. Pro- and mesonota strongly alu- 
taceous, weakly shining, with abundant shallow punctures; 



EVANS : REVISION OP PSEUDISOBRACHIUM 



243 



notauli distinct on anterior .6 of mesoscutum; basal groove of 
scutellnm rather broad and short, lateral foveae large, deep. 
Propodenm about 1.4 X as long as wide, in lateral view 2.2 X 
as long as high ; spiracles elliptical, directed dorso-laterad ; 
median carina rather long, dorsum of propodeum otherwise 
covered with delicate transverse ridges, shining. Mesopleural 
callus rather short and broad, remainder of mesopleurum strongly 
alutaceous and with strong punctures. Discoidal vein of fore 
wing interstitial with median vein, long, well-pigmented, in fact 
the discoidal cell completely outlined by weakly pigmented lines. 
Genitalia much as in other species and as shown in Figures 63, 
but both divisions of the parameres unusually slender. 

Remarks. — There is a possibility that zeteki, described on a 
later page from a single female from Barro Colorado Island, is 
the female of clypeatum. 

Occidentale Species-group 

In this group are placed unusually elongate males possessing 
five-toothed mandibles. The mesopleural callus is often polished, 
but in general the callus is less well differentiated than in the 
crassnm group. The species together range from central Cali- 
fornia to central Mexico and east to western Texas. There is 
very little structural variation within the group, and with one 









TABLE II 








Species and locality 


No. 


LFW 


WF/HE 


00L/W0T 


DA0/WF 


Ant. 11 L/W 


occidentale 














Tehama Co., Calif. 


6 


3.2(2.4-3.7) 


1.29(1.23-1.36) 


.71 (.54-. 88) 


.22 (.20-. 24) 


1.6 (1.4-1.8) 


Mendocino Co., Calif. 


2 


3.6(3.1-4.1) 


1.26(1.24-1.28) 


.80 (.66-. 94) 


.19 (.18-. 20) 


2.0 


Lake Co., Calif. 


101 


4.2(3.4-4.9) 


1.40(1.29-1.50) 


.94 (.86-1.0) 


.19 (.18-. 21) 


2.1 (1.9-2.4) 


Colusa Co., Calif. 


4 


4.0(3.1-4.4) 


1.20(1.06-1.31) 


.69 (.66-. 70) 


.25 (.20-. 29) 


1.9 (1.8-2.0) 


Yolo Co., Calif. 


64 


3.4(2.6-4.7) 


1.28(1.13-1.50) 


.70 (.52-. 88) 


.22 (.17-. 30) 


1.8 (1.4-2.1) 


Sacramento Co., Calif. 


36 


3.5(2.8-4.2) 


1.33(1.05-1.45) 


.73 (.43-. 96) 


.21 (.17-. 31) 


1.8 (1.5-2.0) 


Solano Co. , Calif. 


2 


4.1 (4.0-4.2) 


1.14(1.13-1.15) 


.54 


.28 (.26-. 30) 


2.2 


Marin Co., Calif. 


22 


4.0(3.1-4.3) 


1.34(1.21-1.47) 


.96 (.85-1.1) 


.19 (.17-. 21) 


2.2 (2.0-2.4) 


Contra Costa Co., Calif. 


I 


2.9 


1.53 


.90 


.17 


1.6 


Santa "Clara Co., Calif. 


3 


4.6(4.3-5.0) 


1.28(1.20-1.33) 


.91 (.82-1.05) 


.20 (.18-. 23) 


2.3 (2.1-2.6) 


Los Angeles Co., Calif. 


3 


4.2(4.1-4.3) 


1.41(1.37-1.45) 


1.10 (1.06-1.12) 


.19 (.18-. 19) 


2.1 (1.9-2.3) 


Southeastern Arizona 


3 


3.7(3.6-3.9) 


1.25(1.18-1.33) 


.71 (.67-. 75) 


.22 (.21-. 23) 


1.7(1.6-1.8) 


castaneum 














San Diego, Calif. 


7 


2.5 (2.1-2.7) 


.93 (.90-. 95) 


.63 (.57-. 69) 


.27 (.21-. 32) 


1.7 (1.4-1.9) 


nlgrlculum 














Pachuca, Hidalgo 


4 


2.4 (2.2-2.5) 


1.36 (1.30-1.42) 


.90 (.77-1.07) 


.20 (.18-. 22) 


1.6 (1.4-1.7) 


Zimapan, Hidalgo 


13 


2.4 (2.1-2.7) 


1.10 (.97-1.20) 


.71 (.60-. 82) 


.25 (.22-. 28) 


1.7 (1.4-1.8) 


Sombrerete, Zacatecas 


2 


2.7 


1.20 (1.16-1.23) 


.68 (.63-. 73) 


.26 (.24-. 27) 


1.7 (1.4-1.9) 


matthewsl 














Van Horn, Texas 


10 


1.9 (1.7-2.1) 


1.08 (1.00-1.17) 


.53 (.46-. 61) 


.27 (.23-. 29) 


1.3 (1.1-1.5) 


brunneum 














Pachuca, Hidalgo 


1 


2.7 


1.40 


1.33 


.15 


1.2 


Zimapan, Hidalgo 


24 


2.3(2.1-2.7) 


1.48 (1.40-1.56) 


1.20 (1.1O-1.40) 


.16 (.14-. 17) 


1.2 (1.1-1.4) 



244 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

exception the several forms seem to be allopatrie (at least so far 
as the present very limited records indicate). This one exception 
is brunneum, the most distinctive member of the group, which 
occurs sympatrically with nigricnlum in central Mexico. The 
latter species and all the other members of this group may be 
thought of as comprising a single superspecies which ranges 
throughout the arid regions of the southwestern United States 
and northern Mexico. 

Females assigned tentatively to this group have four-toothed 
mandibles with the basal two teeth small and recessed along 
the inner margin. 

10. Pseudisobrachium occidentale new species 

Jlolotype. — 2 , Novato, Marin Co., Calif., 25 Aug. 1953 (H. L. 
Mathis) [CAS]. 

Description. — Length 5 mm. ; LFW 4 mm. Head and thorax 
dark brownish-fuscous, almost black, abdomen bright chestnut 
brown, with weakly developed annulations of paler brown ; coxae 
dark brown, femora and tibiae medium brown, tarsi light brown ; 
antennal scape nearly black, flagellum medium brown ; wings 
very weakly infuscated, veins and stigma brown. Mandibles with 
five teeth in an oblique series (Fig. 14). Clypeus tectiform 
medially, truncate apically. Antennae elongate, flagellum pubes- 
cent and with a few erect setae which on the basal segments are 
about half as long as width of flagellum, first four antennal seg- 
ments in a ratio of about 11 :3 :5 :5 ; segment eleven twice as long 
as thick. Front of moderate breadth, WF .65 X WH, 1.35 X HE ; 
OOL subequal to WOT ; ocelli in a small triangle, front angle less 
than a right angle, of moderate size, DAO .19 X WF. Vertex, 
in anterior view, rounded off well above eyes, distance from eye 
tops to vertex crest equal to about two-thirds the eye height. 
Front moderately shining, alutaceous, with small, shallow punc- 
tures. Pronotum and mesoscutum moderately shining, the latter 
with the punctures somewhat more sharply defined, although 
small ; notauli very strong on anterior two-thirds of mesoscutum, 
absent behind ; scutellum shining, disc nearly impunctate, basal 
and lateral foveae shallow. Propodeum very long, in dorsal view 
measuring 1.7 X as long as wide, in lateral view measuring 2.6 X 
as long as high ; propodeal spiracles elongate, directed dorsad ; 
disc with fine, reticulate ridges except nearly smooth caudad of 
end of median carina. Mesopleural callus strongly shining, non- 
alutaceous ; remainder of mesopleura alutaceous, anteriorly 



EVANS : REVISION OF PSEUDISOBRACHIUM 245 

rather roughly punctate. Fore wing with discoidal vein rather 
strong, longer than transverse median vein, arising a short dis- 
tance from the top of the latter vein (Fig. 55). Abdomen slender 
and elongate. 

Paratypes. — CALIFORNIA : 6 S $ , Red Bluff, Tehama Co., 
20 July 1956 (light trap, E. Yeomann) [UCD] ; 1 $ , Ukiah, 
Mendocino Co., 12 July 1935 (M. L. Jones) [CDAS] ; 1 $ Hop- 
land Grade, Mendocino Co., 5 May 1959 (S. M. Fidel) [UCD] ; 
100 $ $ , Upper Lake, Lake Co., Aug. 1958 (light trap, R. E. Dol- 
phin) [UCD, CAS, MCZ, CU, CM, INHS] ; 1 $ , Soda Bay, Lake 
Co., 25 July 1958 (light trap, R. E. Dolphin) [UCD] ; 1 $ , 
Arbuckle, Colusa Co., 14 July 1959 (light trap, J. Fowler) 
[UCD] ; 3 $ $ , College City, Colusa Co., June 1959 (light trap, 
J. Fowler) [UCD] ; 25 $ $ , 4.5-9 mi. W. Zamora, Yolo Co., 
June-July 1959 (light trap, J. Fowler) [UCD] ; 20 $ S , 4 mi. 
SW. Dunnigan, Yolo Co., July 1959 (light trap, J. Fowler) 
[UCD] ;8 $ S , Winters & vie, Yolo Co., June-Aug. 1959 (light 
trap, J. Fowler) ; 3 $ $ ,3 mi. NW. Yolo, June, July 1959 (light 
trap, J. Fowler) [UCD] ; 4 $ $ , Davis, July-Sept. [UCD] ; 
3 $ $ , Woodland, 17 Aug. 1959 (light trap, J. Fowler) [UCD] ; 
1 $ , Esparto, Yolo Co., 29 June 1959 (light trap, J. Fowler) 
[UCD] ; 34 $ $ , Fairoaks, Sacramento Co., 12 Aug. 1933 (A. 
Bellue) [USNM, CDAS, CU] ; 1 $ , Sacramento, 14 July 1933 
(H. H. Keifer) [CDAS] ; 1 $ , Rio Linda, 6 July 1958 (light 
trap, J. Fowler) [UCD] ; 1 $ , Rio Vista, Solano Co., 19 July 
1959 (light trap, E. Mezger) [UCD] ; 1 $ , Dixon, 13 July 1956 
(light trap, E. Mezger) [UCD] ; 19 $ $ , Novato, Marin Co., 
Aug.-Sept. (light trap, H. L. Mathis) [CAS, UCD, MCZ] ; 
1 6, Mill Valley, Marin Co., 28 Sept. 1953 (H. L. Mathis) 
[UCD]; 1 $, San Venetia, Marin Co., 10 Oct. 1953 (H. L. 
Mathis) [UCD] ; 1 $ , Danville, Contra Costa Co., 18 Aug. 
1949 (F. X. Williams) [CAS] ; 2 $ S , Los Gatos, Santa Clara 
Co., 1 Aug. 1933 (J. A. Kusche) [CAS] ; 1 $ , Alma, Santa 
Clara Co., 30 Aug. 1933 (11.11. Keifer) [CDAS] ; 3 S S , Glen- 
dale, Los Angeles Co., Sept.-Oct. 1951 (W. M. Schlinger) [UCD]. 
ARIZONA :1$, Cave Creek Canyon, Chiricahua mts., 25 Aug. 
1927 5-6000 feet (J. A. Kusche) [CAS] ; 1 $ , Cochise Strong- 
hold, Cochise Co., 2 Oct. 1954 (at light, Butler and Werner) 
[UA] ; 1 $ , Madera Canyon, Santa Rita mts., 2-4 Aug. 1959 
(K. V. Krombein) [KVK]. 

Variation in males. — The 247 paratypes range in size from 
2.9 to 6.0 mm., fore wing from 2.4 to 5.0 mm. There is consider- 
able variation in width of the front, with specimens from one 



246 



BULLETIN : MUSEUM OF COMPAEATIVE ZOOLOGY 



locality often being fairly consistent in this character, but the 
over-all variation not strongly correlated with geography. Ocel- 
lar size and antennal width appear to vary clinically from west 




Central 
California 




Southeastern 
Arizona 



♦ 




* 



Los Angeles 
Co., Calif. 

Map 1. — Some aspects of geographic variation in males of P. ocoidentale. 
Horizontal bars represent the mean L/W for the eleventh antennal segment ; 
see Table II for actual values and for range of variation. Mean ocellar size 
is superimposed upon this as a circle by taking the DAO/WF ratio, multi- 
plying it by ten, and using the same scale as for antennal length. Vertical 
bars indicate whether the front is weakly (thin line), moderately (thick 
line), or strongly alutaceous (thick bar). It will be seen that in coastal 
localities the ocelli are small as compared to the antennal length and the 
front is weakly alutaceous; as one proceeds eastward, regardless of the 
latitude, there is a tendency for larger ocelli, shorter antennae, and a more 
alutaceous front. 



EVANS: REVISION OF PSEUDISOBRACHIUM 247 

to east, with specimens from coastal California having generally 
longer antennae and smaller ocelli than specimens farther inland 
(Table II and Map 1). Although most specimens have the front, 
thoracic dorsum, and mesoplenral callus rather shining, as in 
the type, in many specimens from Yolo and Sacramento counties, 
California, these parts are less shining and more strongly alu- 
taceous. The three specimens from southeastern Arizona are 
distinctly more heavily alutaceous than any others, and in these 
the mesopleural callus is weakly differentiated. Again, there 
may be a west-to-east cline in this character. Although the 
range of this species covers a considerable distance north-to- 
south, there are no discernible clines in this direction, specimens 
from Sacramento County, California, being more like those from 
southern Arizona than they are like those from Marin County, 
California, a relatively short distance west. This species is 
being collected in great numbers in light traps, and may provide 
an interesting study in intraspecific variation, particularly when 
it is possible to fill in some of the wide gaps in its known range. 

Female (assigned here tentatively). — Stanford Univ., Calif., 
22 Dec. 1909 [USNM]. 

Description of female. — Length of body 3.1 mm., of head 
.6 mm., of thorax 1.1 mm. Head castaneous, thorax light cas- 
taneous, abdomen yellowish-brown; mandibles light castaneous, 
teeth rufous; clypeus and scape light castaneous, flagellum dull 
yellowish-brown; legs wholly yellowish-brown. Mandibles slen- 
der, with four distinct teeth but third and fourth teeth drawn 
well back along inner margin, as figured for obscnrum (Fig. 
36). Clypeus broadly truncate (actually very slightly concave) 
apically; median carina strong, not quite reaching margin. 
Head 1.35 X as long as wide ; sides parallel, arcuately contracted 
near posterior margin to a broad, straight vertex ; occipital carina 
obsolete dorsally. Eyes small, amber-colored and therefore not 
contrasting strongly to the brownish head. Anterior part of 
front and sides of head striato-punctate and also obscurely alu- 
taceous ; posterior half of front smooth and shining between the 
punctures, which are elongate except more circular toward 
vertex; under surface of head alutaceous, punctures rather 
evenly spaced. Pronotal disc 1.4 X as long as its posterior 
width, about as long as maximum width of thorax ; mesonotum 
1.5 X as long as wide, .82 X as long as maximum width of thorax ; 
propodeum 1.6 X as long as wide. Pronotal disc weakly alu- 
taceous behind and on sides, otherwise smooth and shining; 
punctures widely spaced, absent only from midline. Mesonotum 



248 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

obscurely alutaeeous on sides, polished medially, with some weak 
punctures on sides. Propodeum polished dorsally, obscurely 
punctate, spiracles subcircular, directed dorso-laterally. Meso- 
pleurum strongly alutaeeous, punctures rather weak. Body, in- 
cluding legs, with abundant short, pale setae. 

Other females. --CALIFORNIA: 1, same data as preceding 
[USNM] ; 1, Crow Canyon, Alameda Co., 19 Feb. 1939 (K. S. 
Ilagen) [CIS]. 

Variation in females. — The two specimens from Stanford 
University are very similar in size, color, and structure. The 
specimen from Alameda County is considerably larger, the head 
being .94 mm. long, the thorax 1.65 mm. long (the abdomen 
is missing). The thorax is rich castaneous like the head. The 
head is only 1.25 X as long as wide; antero-dorsally it is dis- 
tinctly striato-punctate but hardly alutaeeous, and ventrally 
it is barely alutaeeous but with stronger punctures than in the 
Stanford University specimens. In thoracic structure the three 
specimens are nearly identical. 

Remarks. — The only male Pseudisobrachium known to occur 
in the Sau Francisco Bay area is occidentale, which is not un- 
common and rather variable in size. Therefore, it is most logical 
that these females belong with occidentale. While there is noth- 
ing about their structure which contraindicates this assignment, 
there is still much to be learned about the distribution of western 
Pseudisobrachium, so this association must be considered tenta- 
tive. 

11. Pseudisobrachium castaneum new species 

Holotype.— $, San Diego, Calif. (Ricksecker) [TJSNM, No. 
65153]. 

Description. — Length 3 mm; LFW 2.7 mm. Thorax and 
abdomen chestnut-brown, head also of this color but top and 
front somewhat suffused with fuscous; legs, including coxae, 
uniformly light brown; antennae brown, somewhat darker than 
legs but lighter than body; wings hyaline, veins and stigma 
pale brown. Mandibles with five teeth. Anterior margin of 
clypeus very weakly concave. Antennae elongate, with pubes- 
cence and erect setae as in preceding species ; first four segments 
in a ratio of about 15 :6 :9 :8 ; segment eleven 1.6 X as long as 
thick. Front very narrow, WF only .55 X Wli, .95 X HE; 
ocelli moderately large, OOL only .6 X WOT, DAO .22 X WF ; 
front angle of ocellar triangle slightly less than a right angle. 



EVANS: REVISION OF PSEUDISOBRACHIUM 249 

Vertex rounded off a short distance above eye tops, distance from 
eye tops to vertex crest about .4 X HE. Front shining, alu- 
taceous, punctures shallow, rather numerous. Pro- and mesonota 
also shining and with numerous but rather shallow and incon- 
spicuous punctures; notauli weakly impressed on anterior half 
of mesoscutum; scutellum with basal groove narrow, lateral 
foveae well defined. Propodeum measuring 1.65 X as long as 
wide, in lateral aspect 2.5 X as long as high; spiracles, small, 
elliptical, directed laterad ; most of dorsum of propodeum smooth 
and shining, median carina sharply defined. Mesopleural callus 
strongly shining, barely alutaceous; remainder of mesopleura 
shining, very weakly punctate and without other strong sculptur- 
ing. Fore wing with the discoidal vein longer than the transverse 
median vein, but only very weakly pigmented, arising a short 
distance down on transverse median vein (Fig. 56). 

Paratypes. — ti S $ , all same data as type [USNM, MCZ]. 

Variation. — Body length varies from 2.6 to 3.0 mm., wing 
length from 2.1 to 2.7 mm. Coloration is quite uniform through- 
out the series. Means and range of variation for several charac- 
ters are presented in Table II. 

12. Pseudisobraciiium matthewsi new species 

Holotype. — ■ S , Van Horn, Culberson Co., Texas, 22 July 
1956 (at light, E. G. Matthews) [MCZ, No. 30272]. 

Description. — Length 2.2 mm.; LFW 1.8 mm. Head dark 
brownish-fuscous; thorax and abdomen dark chestnut brown 
except sides of basal abdominal segments paler ; legs light brown 
except coxae somewhat darker ; antennae brown, darker than legs 
but lighter than body; wings hyaline, veins and stigma light 
brown. Mandibles with five teeth, basal three teeth small, sub- 
equal in size and shape. Anterior margin of clypeus truncate. 
Antennae of moderate length, with conspicuous pubescence and 
a few very short erect setae ; first four segments in a ratio of 
about 26:9:11:10; segment eleven 1.2 X as long as thick. WF 
.58 X WH, 1.1 X HE ; ocelli rather large, in a broad triangle, 
front angle slightly greater than a right angle ; OOL .5 X WOT ; 
DAO .27 X WF. Distance from tops of eyes to vertex crest about 
.6 X HE ; top of vertex only weakly rounded, almost straight 
across. Front alutaceous, shining, witli shallow punctures. Pro- 
notum and mesoscutum also shining but strongly alutaceous, the 
scutellum however nearly smooth ; punctures of thoracic dorsum 



250 BULLETIN : MUSEUM OP 1 COMPARATIVE ZOOLOGY 

shallow and inconspicuous; notauli strongly impressed on an- 
terior third of mesoscutum. Propodeum 1.6 X as long as broad, 
in lateral aspect 3 X as long as high ; spiracles small, subcircular, 
directed laterad; surface of propodeum mostly smooth and 
shining, median carina long and sharply defined. Mesopleural 
callus shining, barely alutaceous ; remainder of mesopleura alu- 
taceous but without strong sculpturing or punctures. Fore wing 
discoidal vein very weakly pigmented, about as long as trans- 
verse median vein and arising a short distance from top of that 
vein. 

Paratypes. — 9 $ $ , all same data as type [CU, MCZ, TJSNM]. 

Variation. — The paratypes range in size from 2.0 to 2.4 mm. 
and show no noteworthy differences in color from the type. Mean 
and range of variation for certain characters are presented in 
Table II. There appears to be some variation in the shape of 
the ocellar triangle, but without exception the front angle is at 
least as great as a right angle. 

Female (assigned here tentatively). — El Paso Co., Texas, 22 
July 1937 (W. F. Turner and W. H. Anderson, from soil in peach 
orchard) [USNM]. 

Description of female. — Length of body 1.8 mm., of head 
.43 mm., of thorax .8 mm. Head and thorax medium brown, 
abdomen light yellowish-brown ; antennae and legs straw-colored. 
Mandibles with four well-defined teeth, about as figured for 
obscurum (Fig. 36). Clypeus slightly emarginate apically. Head 
rather slender, 1.42 X as long as wide ; sides subparallel, con- 
verging behind to a straight vertex. Eye a single facet which is 
only slightly paler than the head. Front strongly polished, with 
small punctures which are separated, on the average, by about 
twice their own diameters, surface between punctures without 
any sculpturing whatever ; under side of head also strongly 
polished and with small, rather evenly spaced punctures. Pro- 
notal disc elongate, 1.6 X as long as its posterior width; meso- 
notum also elongate, about twice as long as wide, nearly as long 
as maximum width of thorax; propodeum subovoid, 1.65 X as 
long as wide. Pronotum strongly polished, with weak, widely 
spaced punctures, weakly alutaceous along posterior margin; 
mesonotum strongly polished in middle, otherwise weakly alu- 
taceous ; propodeum also strongly polished, weakly alutaceous 
posteriorly, mesonotum and propodeum both with only a few 
weak punctures on the sides. Sides of mesopleurum weakly alu- 
taceous, weakly punctate. Body and legs with abundant short, 
pale setae. 



EVANS : REVISION OP PSEUDISOBRACHIUM 251 

13. Pseudisobrachium nigriculum new species 

Holotype. — $ , Zhnapan, Hidalgo, Mexico, 11-14 June 1951 
(at light, H. E. Evans) [MCZ, No. 30273]. 

Description. -- Length 2.8 mm.; LFW 2.6 mm. Head dark 
brownish-fuscous, nearly black, thorax and abdomen dark brown, 
abdomen with lighter brown markings latero-posteriorly on basal 
segments ; legs and antennae medium brown ; wings hyaline, veins 
and stigma light brown. Mandibles with five teeth, the basal three 
teeth small, subequal. Anterior margin of clypeus truncate. 
Antennae of moderate length, rather coarsely pubescent and with 
numerous erect setae which toward the base are half as long as 
the thickness of the antennae ; first four segments in a ratio of 
about 18:5 :7 :7, segment eleven 1.5 X as long as thick. WF .58 X 
WII, 1.1 X HE ; ocelli rather large, OOL .6 WOT ; DAO .28 X 
WF; front angle of ocellar triangle slightly less than a right 
angle. Vertex extended well above eye tops, where it is arched 
only weakly ; distance from tops of eyes to vertex crest .67 X HE. 
Front weakly shining, strongly alutaceous, with moderately 
strong punctures which are separated from one another by from 
two to three X their own diameters. Pronotum and mesoscutum 
shining, moderately alutaceous, rather weakly punctate ; notauli 
strong on the anterior fourth of the mesoscutum, absent behind ; 
scutellum polished, basal groove and lateral foveae well defined 
but shallow. Propodeum 1.6 X as long as broad, in lateral view 
3 X as long as high ; spiracles small, elliptical, directed dorso- 
laterad; dorsal surface of propodeum shining, weakly alutaceous 
in front, smooth behind ; median carina strong. Mesoplural cal- 
lus strongly polished, smooth ; remainder of mesopleura also 
shining, but weakly alutaceous and weakly punctate. Fore wing 
with discoidal vein slightly longer than the transverse median 
vein, arising near the top of this vein, moderately well pigmented. 

Paratypes. — HIDALGO : 12 $ $ , same data as type (H. E. 
Evans, P. D. Hurd) [MCZ, USNM, CU, CIS] ; 4 $ 8 , Pachuca, 
29 July 1954 (J. G. Chillcott) [CNC]. ZACATECAS: 2 $ $ , 
15 km. E. of Sombrerete, 28-31 July 1951 (H. E. Evans) [MCZ]. 

Variation. — The paratypes vary in size from 2.2 to 3.0 mm. 
Generally they agree well with the type in coloration, but sev- 
eral specimens, including both of those from Zacatecas, have 
the thorax and abdomen nearly black like the head, and the 
legs and antennae are also darker in these specimens. There are 
no marked structural differences between the Zacatecas speci- 
mens and those from Hidalgo (see Table II). Throughout both 



252 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

series some variation can be noted in the strength of the punc- 
tures and microscopic sculpturing. 

Remarks. — All specimens in the type series were taken at 
Coleman lanterns during the evening hours. The type locality 
was in desert at about 6000 feet, the Zacatecas location in arid 
grassland at about 7000 feet. 

14. PSEUDISOBRACHIUM BRUNNEUM new species 

Holotype. — 8 , Zimapan, Hidalgo, Mexico, 11-14 June 1951 
(at light, H. E. Evans) [MCZ, No. 30274]. 

Description. — Length 2.6 mm. ; LFAV 2.4 mm. Dark brown, 
except base and apex of abdomen suffused with lighter brown; 
antennae dark brown ; legs medium brown, a little paler apically ; 
wings hyaline, stigma brown, veins light brown. Mandibles with 
five teeth, the basal three teeth small, subequal (Fig. 15). Cly- 
peus truncate apically, its median carina arched in profile. 
Antennae of moderate length, first four segments in a ratio 
of about 32:9:12:10, segment eleven 1.3 X as long as thick; 
flagellum with coarse, subappressed pubescence and a few erect 
setae which toward the base are about half as long as thickness of 
flagellum. WF .66 X WH, 1.5 X HE ; ocelli small, OOL 1.2 X 
WOT, DAO .17 X WF ; front angle of ocellar triangle less than 
a right angle. Vertex extended far above eye tops, where it is 
more or less squared off; distance from eye tops to vertex crest 
about equal to eye height. Front shining, moderately alutaceous, 
with shallow punctures. Pronotum and mesoscutum of much the 
same character, latter with notauli impressed on anterior third ; 
scutellum strongly polished medially, basal groove and lateral 
foveae strong. Propodeum 1.8 X as long as broad, in lateral 
view 2.7 X as long as high; spiracles small, elliptical, directed 
dorsad ; median carina strong on anterior two-thirds of disc, disc 
wholly alutaceous but more shining behind. Mesopleura shining, 
weakly alutaceous and weakly punctate, callus somewhat ele- 
vated but not differing otherwise from remainder of pleurum. 
Fore wing with discoidal vein weakly pigmented, about as long 
as transverse median vein and arising just below top of latter 
vein (about as in Fig. 56). 

Paratypes. — HIDALGO: 23 6 8, same data as type (H. E. 
Evans, P. D. Hurd) [MCZ, USNM, CU, CIS, CAS, OEE, 
INHS] ; 1 $ , Pachuca, 29 July 1954 (J. G. Chillcott) [CNC]. 

Variation. ■ — The paratypes vary in size from 2.2 to 2.9 mm. 
Color of the body and legs varies from rich chestnut-brown to 
dark brownish-fuscous with the head and thorax almost black. 



EVANS : REVISION OF PSEUDISOBRACHIUM 



253 



In some specimens the punctures of the front are deep and well- 
defined (although not large), while in others they are scarcely 
evident. The distance from eye tops to the vertex crest varies 
from .7o to 1.0 X HE. Variation in other measurements is not 
great (Table II). 

Remarks. — All specimens were collected at light along with 
specimens of nigricalum. Although the two species are very simi- 
lar in size and color, there are several differences, brunneum 
having shorter antennae, less protruding eyes, and smaller ocelli. 

Obscurum Species-group 

The males of this group have a relatively compact body form 
as in the crassum group, but the mesopleural callus is weakly 
differentiated and alutaceous. Two large species with dark 
wings and punctate mesopleura may represent a link with the 
crassum group. The remaining live species are minute, desertico- 
lous species with pale wings and non-punctate pleura. Tn some 
males of this group the basal mandibular tooth is somewhat 
thicker than the adjacent teeth, approaching the condition in 
the prolongation group. The females of only one species, ob- 
scurum, are known, and these can be assigned here only tenta- 
tively. These females differ in no outstanding features from 
those of the preceding species-group. 

TABLE III 



Species and locality 


No. 


LFW 


WF/HE 




00L/W0T 


DA0/WF 


Ant. 11 L/W 


obscurum 


















Culberson Co. , Texas 
Chihuahua, Chlh. 


11 


2.8 (2.3-3.1) 
2.5 


1.21 (1.16-1 
1.00 


.28) 


.60 (.54-. 65) 
.55 


.22 (.20- 
.26 


24) 


1.3(1.2-1.5) 
1.2 


Cochise Co. , Ariz. 
Graham Co. , Ariz. 


2 


2.6 
2.6 


1.11 
1.18 




.62 

.47 


.22(.21- 
.25 


23) 


1.3 
1.4 


Tucson, Ariz. 
Superior, Ariz. 
Oracle, Ariz. 
Santa Rosalia, Baja Cal. 
Comondu, Baja Cal. 
San Miguel, Baja Cal. 
Venancio, Baja Cal. 
La Paz, Baja Cal. 
Santiago, Baja Cal. 
Mlraflores, Baja Cal. 


101 
3 

6 
2 
4 
1 


2.0 

2.2 (1.6-2.9) 

2.8 (2.3-3.0) 

2.3 

2.4 

2.6 

2.5 (2.1-2.7) 

2.2 (2.0-2.4) 

2.2 (1.9-2.5) 

2.3 


1.07 

1.10 (.98-1 . 

1.11 (1.07-1 

1.03 

1.15 

1.30 

1.22(1.16-1 

1 18(1.15-1 

1.10(1.04-1 

1.08 


20) 
.14) 

.28) 
.21) 
.15) 


.50 

.57 (.42-. 65) 

.54 (.50-. 58) 

.58 

.61 

.65 

.62 (.61-. 65) 

.65 (.53-. 72) 

.64 (.60-. 69) 

.64 


.25 

.23 (.21- 

.23 (.22- 

.22 

.22 

.18 

.20(.18- 

.20(.19- 

.2! (.20- 

.22 


.26) 
.24) 

.23) 
.21) 
.22) 


1.2 

1.2(1.1-1.4) 

1.3(1.2-1.4) 

1.4 

1.3 

1.3 

1.2(1.1-1.4) 

1.1 

1.2(1.0-1.3) 

1.3 


otlosum 


















Superior, Ariz. 


3 


2.0 (1.9-2.2) 


1.48 (1.44- 


.52) 


.95 (90-1.00) 


.15 




1.3 


mlchoacanum 


















Tuxpan, Mlchoacan 


1 


3.0 


1.60 




1.70 


.13 




1.5 


carollilanum 


















Grahamville, S. C. 


1 


3.3 


1.30 




1.12 


.16 




1.4 


Miami, Fla. 


3 


3.2 (2.9-3.4) 


1.12 (1.05- 


.23) 


.76 (.70-. 32) 


.20 (.18- 


.23) 


1.4 (1.3-1.4) 


pallidum 


















Yuma Co. , Ariz. 


2 


1.8 


1.08 (1.06- 


.10) 


.74 


.20 




1.0 


gibbosum 


















Hidalgo Co., N. Mex. 


1 


1.6 


1.68 




1.10 


.15 




1.1 


testacelpes 


















San Marcos, Nicaragua 


1 


1.8 


1.43 




1.40 


.14 




1.1 



254 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

15. Pseudisobrachium michoacanum new species 

Holotype. — $ , Tuxpan, Michoacan, Mexico, 6000 feet eleva- 
tion, 6 July 1959 (H. E. Evans) [MCZ, No. 30275]. 

Description. — Length 3.6 mm.; LFW 3.0 mm. Head and 
thorax black; abdomen shining black, except first tergite mar- 
gined with brown; scape black, flagellum dark brown; apical 
half of mandibles rufous; legs dark brown, beyond the middle 
of the tibiae medium brown ; wings lightly infuscated, covered 
with brown setulae, veins and stigma dark brown. Mandibles 
with five teeth, basal three teeth about equally developed (Fig. 
18). Clypeus truncate apically. Antennae elongate, first four 
segments in a ratio of about 16:5:11:11, segment eleven 1.5 X 
as long as thick ; pubescence light brown, erect setae numerous, 
some of them nearly half as long as thickness of flagellum. Front 
broad, WF .7 X WH, 1.6 X HE ; ocelli small, far removed from 
eyes, forming an angle in front that is less than a right angle ; 
DAO .13 X WF; OOL 1.7 X WOT. Vertex extended far above 
eye tops, distance from tops of eyes to vertex crest, subequal 
to eye height. Front alutaceous, moderately shining, with shallow 
punctures which are separated from one another by from 1 to 2 
times their own diameters. Pro- and mesonota wholly alutaceous, 
including disc of scutellum, and with abundant small punc- 
tures which are fairly dense on sides of mesoscutum; notauli 
impressed on anterior .4 of mesoscutum. Propodeum 1.45 X as 
long as broad, in lateral view 2.1 X as long as high; spiracles 
elliptical, directed dorso-laterad ; dorsal surface with rather 
strong reticulate sculpturing. Mesopleurum with callus wholly 
alutaceous, not strongly convex ; anterior portion of mesopleurum 
with many large punctures. Discoidal vein of fore wing arising 
a short distance down on transverse median vein, pigmented to 
about length of basal vein. 

Remarks. — The only known specimen of this species was taken 
on oak foliage at about 11 in the morning in open, bushy coun- 
try. The small ocelli and dark coloration suggest that this is 
a diurnal species. 

16. Pseudisobrachium carolinianum new species 

Holotype. — ■ S , Grahamville, South Carolina, 1 Aug. 1952 
(J. Shuler) [Coll. H. K. Townes]. 

Description. — Length 4.5 mm.; LFW 3.3 mm. Head and 
thorax black, abdomen dark brown, basal tergite margined with 
paler brown ; mandibles yellow-brown, teeth rufous ; antennae 



EVANS : REVISION OF PSEUDISOBRACHIUM 255 

wholly ferruginous; legs bright castaneous except front coxae 
slightly infuseated ; wings hyaline, veins and stigma brown. 
Mandibles with five teeth, basal tooth slightly thicker and more 
rounded than the third and fourth teeth. Clypeus truncate apic- 
ally. Antennae of moderate length, first four segments in a ratio 
of about 23 :5 :11 :10 ; segment eleven about 1.4 X as long as thick ; 
pubescence and setae pale, the latter rather numerous and 
mostly about a third as long as the flagellar width. WF .65 X 
WH, 1.3 X HE; DAO .16 X WF; OOL 1.12 X WOT; ocelli 
forming an angle in front which is slightly less than a right 
angle. Vertex extended above eye tops a distance equal to about 
.8 X HE. Front alutaceous, weakly shining, with large, shallow 
punctures which are separated from one another by from 1 to 
2 times their own diameters. Pro- and mesonota moderately 
shining, alutaceous, the punctures moderately strong ; notauli 
strong on anterior half of mesoscutum. Propodeum measuring 
1.3 times as long as broad; spiracles elongate-elliptical, directed 
laterad ; disc with fairly strong sculpturing which shows some 
tendency to form transverse striae. Mesopleural callus aluta- 
ceous, not strongly convex ; anterior portion of mesopleurum with 
many large punctures. Fore wing with discoida] vein about as 
long as basal vein, weakly pigmented. 

Paratypcs. — S 6 $, Miami, Florida, 1-i July 1950 (F. G. 
Butchers) [CNC, MCZ, USNM]. 

Variation. — The paratypes differ from the type in having 
consistently larger ocelli and a narrower front ; the punctures 
of the body also tend to be somewhat weaker. It is conceivable 
that these represent a distinct species, but I think not. 

17. Pseudisobrachium gibbosum new species 

Holotype. - - £ , 8 mi. N. of Rodeo, Hidalgo Co., New Mexico, 
4000 feet elevation, 8 Sept. 1959 (at light H. E. Evans) [MCZ, 
No. 30276]. 

Description. — Length 2.8 mm. ; LFW 1.6 mm. Head, thorax, 
and abdomen dark brownish-fuscous ; mandibles ferruginous ; 
antennae dark brown ; legs dark brown except tarsi and apical 
part of tibiae paler ; wings very pale, stigma brown but veins 
nearly colorless. Mandibles with five teeth, the basal three 
teeth small, subequal. Clypeus broad basally, its sides tapering 
to a rather narrowly truncate apex. Antennae short, first four 
segments in a ratio of about 13 :4 :4 A ; flagellar segments, except 
the last, only very slightly longer than thick, pubescence of 
flagellum pale and rather conspicuous although short, erect setae 



256 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

short and few in number. Front broad, WF .74 X WH, 1.68 X 
HE; ocelli small, DAO .15 X WF, OOL 1.1 X WOT; front 
angle of oeellar triangle approximately a right angle. Vertex 
extended above eye tops a distance about equal to eye height. 
Front strongly shining, very weakly alutaceous, with a few 
strong punctures on upper half. Pro- and mesonota also strongly 
shining, the mesoscutum with a few small but deep punctures; 
notauli weakly impressed on anterior half of mesoscutum. Pro- 
podeum short, measuring 1.3 X as long as broad; spiracles small, 
subcircular, opening laterad ; median carina present, disc with 
fine reticulate sculpturing over entire dorsal surface, stronger 
anteriorly. Mesopleurum shining and very weakly alutaceous 
except on the extreme anterior part, which is heavily sculptured ; 
central portion of mesopleurum convex, not only the callus 
but the areas beneath and behind it, with a single large pit in 
the center. Discoidal vein of fore wing absent. 

18. Pseudisobrachium otiosum new species 

Holotype. — <$ , Superior, Pinal Co., Arizona, 7-17 July 1948 
(II. K. Gloyd) 4 [USNM, No. 65159]. 

Description. — Length 2.8 mm.; LFW 1.9 mm. Head nearly 
black ; thorax and abdomen dark brown ; apical half of mandibles 
yellowish, teeth rufous ; antennae light yellowish-broAvn, includ- 
ing scape; front coxae brown, legs otherwise straw-yellow ; wings 
hyaline, clothed with pale setulae, stigma light brown, veins 
nearly colorless. Mandibles with five teeth, the basal three teeth 
small, subequal. Clypeus truncate apically. Antennae rather 
short, first four segments in a ratio of about 14 :4 :6 :6 ; flagellar 
segments each slightly longer than thick, segment eleven 1.3 X 
as long as thick ; pubescence of flagellum pale, erect setae pale, 
numerous, less than half as long as width of flagellum. WF 
.7 X AVH, 1.52 X HE ; ocelli small, DAO .15 X WF, OOL equal 
to WOT ; ocelli forming an angle in front which is slightly less 
than a right angle. Vertex extended above eye tops a distance 
equal to about .8 X HE. Front strongly alutaceous, weakly 
shining, with shallow, inconspicuous punctures. Pro- and meso- 
nota alutaceous, weakly punctate, moderately shining, notauli 
impressed only on the anterior .2 of mesoscutum. Propodeum 



* This is the data given on the label of the type, except that the collector's 
name is given as I). K. G. Lloyd. Almost certainly this specimen was taken at 
a light trap by II. K. Gloyd at the Boyce Thompson Southwestern Arboretum, 
4 mi. W. of Superior, as I have seen much other material taken hv him there 
on those dates I INHS]. 



EVANS: REVISION OF PSEUDISOBRACHIUM 257 

rather short, measuring 1.3 X as long as broad, in lateral view 
2.2 X as long as high ; spiracles small, elliptical, directed dorsad ; 
median carina well developed, disc shining, with weak reticulate 
sculpturing. Mesopleurum wholly alutaceous, without strong 
punctures and with the callus poorly differentiated. Discoidal 
vein of fore wing absent. 

Paratype.— ARIZONA: 1 $ , same data as type [MCZ] ; 1 $ , 
Boyce Thompson Southwestern Arboretum, 4 mi. W. Superior, 
28 Sept, 1949 (light trap, B. W. Benson) [INHS]. 

Variation. — One paratype is slightly larger, measuring 3 mm. 
in length, the fore wing 2.2 mm. In one paratype the third and 
fourth mandibular teeth are much smaller than the basal tooth, 
while in the second paratype the fourth tooth is so minute that 
the mandibles appear four-toothed except upon very close in- 
spection. 

19. Pseudisobraciiium testaceipes Kieffer 

Pseudisobrachium testaoeipes Kieffer, 1906, Berlin Eut. Zeitschr., 50:240. 
[Type: $, San Marcos, Nicaragua (Coll. Baker) (Pomona College, 
Claremont, Calif.)]. — Kieffer, 1914, Das Tierreich, 41:480. 
Description of holotype. — Length 2.5 mm.; LFW 1.8 mm. 
Head piceous, thorax dark castaneous, abdomen dark castaneous 
except light yellowish-brown on sides of basal segments; apical 
two-thirds of mandibles yellowish-brown, teeth rufous; scape 
light brown, flagellum dull castaneous ; legs pale castaneous ; tarsi 
straw-colored; wings hyaline, stigma light brown, veins amber; 
setulae of wing membrane brownish. Mandibles with five teeth, 
basal three teeth small, subequal. Clypeus narrowly truncate 
apically (actually very weakly arcuately concave). Antennae 
short, first four segments in a ratio of about 17:5:7:6, each 
flagellar segment very slightly longer than thick, segment eleven 
1.1 X as long as thick ; flagellar pubescence coarse, semi-erect, 
erect setae numerous and prominent. WF .68 X WH, 1.43 X 
HE; ocelli small, DAO .14 X WF; ocelli in a compact triangle, 
OOL 1.4 X WOT ; vertex extended above eye tops a distance 
equal to about .7 X HE. Front alutaceous, moderately shining, 
punctures shallow, inconspicuous. Pro- and mesonota (including 
scutellar disc) strongly alutaceous, almost beaded, however rather 
shining, punctures shallow and inconspicuous; notauli weakly 
impressed on anterior .4 of mesoscutum. Propodeum about 
1.4 X as long as broad ; disc with some short carinae arising from 
base in addition to usual median carina, posterior part of disc 



258 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

alutaceous, somewhat shining ; spiracles elliptical, directed dor- 
sad. Mesopleural callus strongly convex, shining although mod- 
erately alutaceous, remainder of mesopleurum alutaceous, ob- 
scurely punctate. Fore wing with discoidal vein present as a 
pigmented streak about as long as the basal vein. 

Remarks. — I have seen no specimens of this species other 
than the type. It is a reasonably distinctive species, but has 
doubtless escaped attention from collectors because of its small 
size. 

20. Pseudisobrachium pallidum new species 

Holotype. — S , Wellton, Yuma Co., Arizona, 9 Aug. 1917 
(Cornell Univ. Biol. Exp., J. C. Bradley) [CU]. 

Description. — Length 2.5 mm., LFW 1.8 mm. Entire body 
light brown ; mandibles light yellowish-brown ; antennae light 
yellowish-brown, flagellum with a tinge of rufous ; legs straw- 
colored; wings very pale, setulae pale, stigma tinged with 
brown, veins colorless. Mandibles with basal three teeth small, 
basal tooth with its inner margin arching into the inner mandibu- 
lar margin (Fig. 17). Clypeus broad basally, tapering to a 
rather narrowly truncate apex. Antennae short, first four seg- 
ments in a ratio of about 3:1:1:1, segment three about 1.2 X 
as long as broad, segment eleven about as long as broad ; flagellar 
pubescence pale, subappressed, erect setae virtually absent. 
Front very narrow, WF .59 X WH, 1.06 X HE; ocelli of mod- 
erate size, DAO .20 X HE ; OOL .74 X WOT. Eyes somewhat 
bulging, vertex elevated above eyes a distance equal to about 
half HE. Front shining though regularly alutaceous, punctures 
weak and scarcely noticeable. Pro- and mesonota alutaceous 
although somewhat shining, without noticeable punctures ; notauli 
absent. Propodeum very short, about 1.35 X as long as wide; 
disc shining, with weak sculpturing basally ; spiracles small, 
subcircular, directed laterad. Mesopleurum rather convex, shin- 
ing but wholly alutaceous, callus not well differentiated ; punc- 
tures inconspicuous. Discoidal vein of fore wing absent. 

Paratype. — 1 $ , same data as type [CU]. 

Variation. — The single paratype is very similar to the type 
in every respect, including size. 

21. Pseudisobrachium obscurum new species 

Holotype. — $ , Pine Springs, Culberson Co., Texas, 13-16 
July 1955 (at light, E. G. Matthews) [MCZ, No. 30278]. 



EVANS : REVISION OF PSEUDISOBRACHIUM 259 

Description. — Length 4.6 mm. ; LFW 3.1 mm. Head and 
thorax black, abdomen dark brown, with indistinct banding witb 
paler brown toward base ; apical half of mandibles yellowish, 
teeth rufous ; scape brown, flagellum bright reddish -brown ; 
front coxae nearly black, middle and hind coxae and femora 
medium brown, remainder of legs light brown ; wings hyaline, 
covered with pale setulae, stigma light brown, veins nearly color- 
less. Basal three teeth of mandibles subequal (Fig. 16). Clypeus 
truncate apically. Antennae rather short, first four segments 
in a ratio of about 18 :5 :10 :9, segment eleven 1.5 X as long as 
thick ; antennal pubescence extremely fine, pale, and closely ap- 
pressed, erect setae short and few in number. WF .66 X WH, 
1.20 X HE; ocelli slightly enlarged, DAO .21 X WF; front 
angle of ocellar triangle slightly greater than a right angle, OOL 
only .6 X WOT. Vertex broadly rounded, almost squared off, 
distance from eye tops to vertex crest equal to somewhat more 
than half HE. Front alutaceous, weakly shining, punctures 
numerous but small and inconspicuous. Pronotum and mesoscu- 
tum alutaceous, moderately shining, with numerous small punc- 
tures ; notauli weakly impressed on anterior fourth of meso- 
scutum; scutellar disc shining. Propodeum about 1.3 X as long 
as broad ; spiracles small, elliptical, directed dorsad ; disc alu- 
taceous but without other sculpturing except for median carina. 
Mesopleurum alutaceous, weakly punctate in front, callus not 
strongly differentiated. Discoidal vein of fore wing distinct, 
although very weakly pigmented like the rest of the veins. 

Paratypes. — TEXAS : 10 $ $ , same data as type [MCZ, CU, 
USNM]. CHIHUAHUA: 1 s, Chihuahua, 12 Aug. 1951 (at 
light, II. E. Evans) [MCZ]. ARIZONA: 2 S S ,1 mi. S. Portal 
Cochise Co., 12 Aug., 5 Sept. 1959 (at light, H. E. Evans) 
[CU, MCZ]; 1 8, Post Canyon, Pinaleno Mts., Graham Co., 
5-6000 feet, 16 July 1917 (W. M. Wheeler) [MCZ] ; 1 $ , Tuc- 
son, 15 July 1937 (Bryant) [CAS] ; 3 $ $ , Oracle, Pinal Co, 
25 July 1917 (W. M. Wheeler) [MCZ] ; 101 $ $ , Boyce Thomp- 
son Southwestern Arboretum, 4 mi. W. Superior, Pinal Co, May- 
Oct. (at light, H. K. Gloyd and B. W. Benson) [INHS, USNM, 
MCZ, CU], BAJA CALIFORNIA: 1 $ , 25 mi. S. Santa Rosalia, 
25 July 1938 (Michelbacher & Ross) [CAS] ; 1 S , Comondu, 22 
July 1938 (Michelbacher & Ross) [CAS] ; 1 $ , 5 mi. S. San 
Miguel, 20 July 1938 (Michelbacher & Ross) [CAS] ;6 U, 
Venancio, 17 July 1938 (Michelbacher & Ross) [CAS, MCZ] ; 
2 $ $ , La Paz, 7 Oct. 1941 (Ross & Bohart) [CAS] ;4 U, 
Santiago, 8 July 1938 (Michelbacher & Ross) [CAS] ; 1 S, 



260 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

5 mi. S. Miraflores, 10 July 1938 (Michelbacher & Ross) [CAS]. 

Variation in males. — The 135 paratypes range in size from 
2.1 to 4.4 mm., fore wing from 1.6 to 3.1 mm. In some of the 
smaller Arizona specimens the head and thorax are much more 
shining and less alutaceous than in the type and most other 
specimens. In some Arizona specimens the basal mandibular 
tooth is thicker than the third and fourth teeth, more as in 
Figure 17. Some of the Arizona specimens have the abdomen 
light brown, occasionally the whole body light brown ; these 
may, of course, be somewhat teneral. In the specimen from 
Chihuahua the antennae are darker than usual, while in one of 
the two specimens from Cochise County, Arizona, the antennae 
are very dark brown and the legs darker than usual. Head meas- 
urements do not exhibit an undue amount of variation (Table 
III). 

Female (assigned here tentatively). — Sacaton, Pinal Co., Ari- 
zona, 1935, from soil in cotton field (L. D. Christenson) [USNM]. 

Description of female. — Length of body 1.8 mm., of head 
.5 mm., of thorax .9 mm. Head castaneous ; thorax pale cas- 
taneous; abdomen, legs, antennae, clypeus, mandibles, and sides 
of head anterior to eyes, light yellowish-brown. Mandibles rather 
slender, basal two teeth small and situated back along inner mar- 
gin of mandible (Fig. 36). Clypeus truncate apically, median 
ridge strong, not prolonged beyond margin. Head 1.35 X as long 
as wide ; sides nearly parallel, weakly convergent behind to a 
broad, straight vertex ; occipital carina obsolete dorsally. Eye a 
fairly large, whitish facet which contrasts well with the brown- 
ish head. Front punctate except along a narrow median band, 
punctures generally a bit longer than wide, separated from one 
another mostly by about their own diameters ; surface of front 
weakly alutaceous between punctures, though rather strongly 
shining ; under surface of head weakly alutaceous, with small, 
rather evenly spaced punctures. Pronotal disc 1.6 X as long as its 
posterior width, about as long as maximum width of thorax ; meso- 
notum 1.4 X as long as wide, .68 X as long as maximum width 
of thorax; propodeum 1.5 X as long as wide. Pro- and mesonota 
weakly alutaceous, moderately shining, with small, widely spaced 
punctures which are largely absent medially; propodeum weakly 
alutaceous, obscurely punctate on sides; spiracles subcircular, 
opening dorso-laterad. Mesopleurum weakly alutaceous, punc- 
tures numerous but not very strong. Body with abundant pale 
setae ; coxae, femora, and tibiae each with a few fairly long pale 
setae. 



EVANS : REVISION OF PSEUDISOBRACHIUM 



261 



Other females. — ARIZONA : 4, same data as preceding 
[USNM, MCZ] ; 1, Huachuca mts., 14 Sept. 1938 (R. H. Cran- 
dall) [UA]. 

Variation in females. — Variation in size, color, and standard 
measurements in this series is slight. One specimen from Sacaton 
is smaller than the other three, the head measuring only .42 mm., 
thorax .75 mm. In this specimen the head and thoracic dorsum 
are only very obscurely alutaceous and therefore more strongly 
shining than in the other three. The specimen from the Huachuca 
mts. is slightly larger than any of the others, the body measuring 
2.6 mm., head .63 mm., thorax 1.1 mm. ; in all other respects it 
agrees very well with description presented above. 

Remarks. — The males of this species bear a close resemblance 
to those of flavinervis Fouts, another nocturnal, deserticolous 
species which occurs over much the same range. This resem- 
blance extends to the shape of the ocellar triangle and nature of 
the antennal pubescence as well as to features which usually 
characterize desert species (pale wings, large ocelli, etc.). Pre- 




Map 2. — Distribution of P. obscurum, males indicated by solid circles, 
supposed females by hollow circles. Distribution of P. armarium, males 
indicated by solid squares, supposed females by hollow squares. 



262 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



sumably these resemblances must be the result of convergence, 
since on the basis of mandibular structure the two forms belong- 
to different species-groups. In general, the front is less distinctly 
punctate in dbscurum than in flavinervis. The distribution of 
this species is shown on Map 2. 



Prolongatum Species-group 

Within this complex of three species the usually reliable 
character of the mandibular dentition breaks down. The basal 
mandibular tooth of the male is broad, its inner margin arching 
into the inner margin of the mandible. The third and fourth 
teeth are small, in some specimens of two of the species very 
small, even connate, and in a number of specimens of 'prolonga- 
tum (about 10 per cent of those examined) these two teeth are 



Species and locality 



aztecum 



No. 



LFW 



TABLE IV 
WF/HE O0L/W0T 



DAO/WF 



Norway Bay, Que. 4 

Ottawa, Ont. and vie. 2 
St. John, N. B. 2 

Bridgetown, Nova Scotia 49 



Bar Harbor, Me. 
Stratton, Me. 
Houghton, Me. 
Westerly, R. I. 
Lake George, N. Y. 
Oneonta, N. Y. 
Princeton, N.J. 
Mt. Holly Spr., Pa. 
Plummer's Isl., Md. 
Takoma Park, Md. 
Washington, D C. 
Bolivar, W. Va. 
Hamrlck, N. C. 
Crabtree Mds., N. C. 
Mt. Plsgah, N. C. 
Mt. Mitchell, N. C. 
Sioux City, Iowa 
Chllllvack, B.C. 
Tenlno, Wash. 
Union Gap. Wash. 
Spokane Falls, Wash. 



1 
96 

7 

1 

1 

2 
14 
12 

4 



3.4 (3.2-3.7) 
3.6 (3.5-3.7) 
3.1 

3.3 (2.6-3.7) 

3.4 

3.0 

3.6 

3.1 

3.8 

3.7 

3.6 

3.5 

3.3 (2.5- 

3.1 (2.5-: 

3.7 

2.6 

3.0 (2.9-3.1) 

3.5 (2.9-3.9) 

3.6 (3.0-4.1) 
3.6 (3.2-3.9) 
3.0 

3.5 
3.6 
2.7- 
3.5 



-3.7) 
-3.7) 



1.65 (1 
1.63 (1 
1 .6? (1 
1.58 (1 
1.72 
1.68 
1.70 
1.55 
1.75 
1.62 
1.51 
1.52 
1.60 (1 

1.66 (1 
1.54 
1.63 
1.68 (1 

1.66 (1 

1.67 (I 
1.65 (1 
1.56 
1.58 
1.62 
1.60 
1.56 



63-1.69) 
60-1 .66) 
65-1.73) 
53-1.64) 



50-1.75) 
62-1 .70) 



61-1.75) 
62-1 .72) 
63-1 .70) 
60- 1.70) 



1.42(1. 

1.29(1. 

1.41 (1 

1.30 (1 

1.32 

1.52 

1.38 

1.25 

1.52 

1.33 

1.32 

1.32 

1.35 (1 

1.30 (1 

1.18 

1.35 

1.41 (1 

1.45 (1 

1.50(1 

1 .47 (1 

1.46 

1.22 

1.28 

1.15 

1.25 



36-1. 4E) 
26-1.31) 
40-1.42) 
26-1.33) 



24-1 

24-1 



.43) 
.39) 



34-1.48) 
35-1.58) 
37-1.64) 
02-1 .60) 



.14 

.14 

.14 

.14 (.13-. 15) 

.14 

.14 

.14 

.15 

.13 

.13 

.14 

.13 

.14 (.12-. 15) 

.13 (.12-. 14) 

.14 

.13 

.14 

.14 (.13- 

.14 (.13- 

.14 

.12 

.13 

.13 

.14 

.14 



.15) 
.15) 



Ant. 11 L/W 



Cuernavaca, Mor. 


1 


4.1 


1.23 




.60 


.27 




1.6 


arenarium 


















Pine Barrens, N.J. 


5 


4.3(3.8-5.1) 


.96 (.93- 


1.01) 


.61 (.54-. 70) 


.25 (.23- 


.26) 


1.9(1.7-2.0) 


Kill Devil Hills, N. C. 


3 


3.4 (3.1-3.6) 


.98 (.96- 


.9?) 


.65 (.61-67) 


.24 (.23- 


.25) 


1.6(1.3-1.8) 


Tryon, N. C. 


1 


3.9 


1.00 




.65 


.22 




1.8 


Clarka Co., Ga. 


1 


3.6 


1.07 




.68 


.22 




2.0 


"Georgia* 


2 


3.4 (3.2-3.6) 


.96 (.95- 


.97) 


.62 (.60-. 64) 


.23 




1.6(1.5-1.7) 


Orlando, Fla. 


1 


4.1 


1.24 




.89 


.18 




1.6 


Camp Rucker, Ala. 


2 


4.3 (4.2-4.4) 


.96 (.95- 


■ 97) 


.69 (.63-. 70) 


.23 (.22- 


.24) 


1.7(1.6-1.8) 


Camp Shelby, Miss. 


2 


4.2 (4.1-4.3) 


.99 (.98- 


1.00) 


.69 (.68-. 70) 


.23 (.22- 


.24) 


1.9 


Urbana, III. 


15 


4.3 (3.8-4.8) 


.98 (.95- 


1.02) 


.64 (.55-. 71) 


.25 (.24- 


.26) 


1.8(1.6-1.9) 


prolongatum 



















1.6(1.5-1.7) 

1.6(1.5-1.7) 

1.6 

1.6(1.5-1.8) 

1.7 

1.8 

1.8 

1.6 

1.7 

1.6 

1.7 

1.5 

1.7(1.3-1.8) 

1.6(1.5-1.7) 

1.6 

1.8 

1.6(1.5-1.7) 

1.6(1.5-1.7) 

1.7(1.5-1.8) 

1.6 

1.6 

1.9 

1.5 

1.5 



actually fused. Thus this group is distinctly intermediate be- 
tween the three species-groups having five-toothed mandibles and 
the two having four-toothed mandibles. 



EVANS : REVISION OF PSEUDISOBRACHIUM 263 

22. Pseudisobrachium aztecum new species 

Holotype. — $ , Cuernavaca, Morelos, Mexico, 5500 feet eleva- 
tion, 6 June 1959 (at light, H. E. Evans) [MCZ, No. 30279]. 

Description. — Length 5.3 mm. ; LFW 4.1 mm. Head and 
thorax black, abdomen dark brown, basal segments suffused with 
paler; apical half of mandibles ferruginous; antennae brown; 
legs bright yellowish-brown except front coxae infuscated ; wings 
subhyaline, veins brown, stigma dark brown. Mandibles with 
five teeth, the fifth tooth broad, continuous with the inner mar- 
gin (Fig. 19). Clypeus truncate apically. Antennae fairly long, 
first four segments in a ratio of about 14 :4 :7 :7, segment eleven 
1.6 X as long as thick; pubescence of flagellum pale and ap- 
pressed, erect setae short, sparse, and inconspicuous. WF .62 X 
WH, 1.23 X HE ; ocelli large, anterior ocellus nearly .2 mm. 
in diameter, DAO .27 X WF ; OOL .6 X WOT. Vertex extended 
above eye tops a distance equal to only slightly over half eye 
height. Front alutaceous, moderately shining, with abundant 
small punctures. Pro- and mesonota strongly shining, non-alu- 
taceous, with abundant small punctures which are much more 
sparse medially; notauli strong on anterior .6 of mesoscutum. 
Propodeal disc dull and with fine sculpturing in front, more 
smooth and shining behind ; spiracles elongate, directed dorsad ; 
propodeum 1.5 X as long as broad. Mesopleurum shining, non- 
alutaceous, the anterior portion punctate. Discoidal vein of fore 
wing arising a short distance down on the transverse median 
vein, pigmented to about length of basal vein. 

23. Pseudisobrachium arenarium new species 

Holotype. — $ , Lebanon State Forest, Burlington Co., New 
Jersey, 19 Aug. 1958 (at light, H. E. Evans and D. F. Bene- 
way) [MCZ, No. 30280]. 

Description. — Length 5.5 mm. ; LFW 4.3 mm. Head and 
thorax black, abdomen brown, basal segments suffused with light 
brown ; apical half of mandibles light brown, teeth rufous ; an- 
tennae medium brown; legs bright castaneous, except front 
coxae blackish ; wings faintly tinged with brown, veins and 
stigma brown. Mandibles with five teeth, third and fourth teeth 
small, fifth tooth very broad and blunt (Fig. 20). Clypeus 
truncate apically. Antennae fairly long, first four segments in a 
ratio of about 14:3:8:7, segment eleven 1.9 X as long as thick; 
pubescence of flagellum pale, rather rough and suberect, erect 
setae numerous, most of them about half as long as thickness 
of flagellum. Front very narrow, WF .52 X WH, .93 X HE ; 



264 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

ocelli rather large, anterior ocellus measuring about .15 mm. in 
diameter, DAO .26 X WF ; OOL .54 X WOT. Vertex extended 
above eye tops a distance equal to about half HE. Front alu- 
taceous, weakly shining, with large, shallow punctures which are 
separated from one another by from one to two times their own 
diameters. Pro- and mesonota alutaceous, punctate, weakly shin- 
ing ; notauli sharply impressed on anterior half of mesoscutum ; 
scutellar disc shining. Propodeum 1.5 X as long as broad, its 
disc weakly sculptured except at the end of the median carina, 
where there is a smooth and shining area; spiracles elongate- 
elliptical, directed dorsad. Mesopleurum alutaceous and with 
strong though shallow punctures, except that the callus is well- 
defined and somewhat shining. Discoidal vein of fore wing aris- 
ing a short distance down on transverse median, pigmented to 
slightly more than length of basal vein (as figured for prolonga- 
tion, Fig. 57). 

Paratypes. — NEW JERSEY: 2 $ $, same data as type 
[MCZ] ; 2 S S , Wrangle Brook Road, Lakehurst, N. J., 26 Aug., 
7 Sept. 1955-56 (D. M. Anderson, J. G. Franclemont) [CU]. 
NORTH CAROLINA : 3 <J <J , Kill Devil Hills, Dare Co., 27-29 
July 1955 (at light, K. V. Krombein) [KVK] ; 1 $ , Tryon, 
25 July (at light, AV. F. Fiske) [USNM]. GEORGIA: 1 8, 
Clarke Co., 25 Sept, 1959 (Richards) [CU] ; 2 $ & , no further 
data [ANSP]. ALABAMA: 2 $ $ , Camp Rucker, 12 Dec. 1942 
(J. G. Franclemont) [USNM]. MISSISSIPPI: 2 $ $ , Camp 
Shelby, nr. Hattiesburg, 30 July, 2 Sept. 1943 (C. D. Michener) 
[AMNH]. ILLINOIS: 15 $ $ , Urbana, Aug.-Oct. (at light, 
C. A. Hart, G. T. Reigel) [INHS, MCZ, CU]. 

Other specimens not designated paratypes : 3 $ 6 , without 
data [INHS] ; 1 $ tentatively assigned here from FLORIDA: 
Orlando, July 1927 (0. C. McBride, in light trap) [USNM]. 

Variation in males. — The 30 paratypes vary in size from 4.2 
to 6.8 mm., fore wing from 3.1 to 5.1 mm. (Table IV). The 
abdomen is lighter than the head and thorax in all specimens and 
in some is light reddish-brown. The mesopleural callus varies 
from moderately to very strongly shining. Some of the variation 
shown in Table IV appears to be clinal, specimens from the North 
having, for example, generally larger ocelli than those from the 
South. The specimen from Florida which I assign here, tenta- 
tively, has unusually small ocelli and unusually wide front. 
Among the other specimens, the smallest ocelli and widest front 
occurs in some of those from Georgia and North Carolina. It 
seems probable that the Florida specimen, different as it seems, 
merely represents the extreme of one or more clines. 



EVANS : REVISION OF PSEUDISOBRACHIUM 265 

Female (assigned here tentatively). — St. Charles, Mo., 1949, 
"in vial with Stig [matomma] pallipes" (M. Talbot) [USNM]. 

Description. — Length of body 4 mm., of head .92 mm., of 
thorax 1.7 mm. Entire body castaneous, head slightly darker 
than thorax, abdomen slightly paler than thorax; mandibles, 
elypens, and scape light castaneous, fiagellum dull yellowish- 
brown ; legs bright yellowish-brown. Mandibles with three teeth, 
as figured for prolongatum (Fig. 39), clypeus broadly subtrun- 
eate, its median carina sharp but not reaching apical margin. 
Head 1.15 X as long as wide, sides nearly parallel, posteriorly 
arcuately convergent to a broad, nearly straight vertex. Eyes 
each consisting of a single pale, fairly conspicuous facet. Front 
with a median impunctate streak, otherwise with close, elongate 
punctures, somewhat alutaceous, on lower sides somewhat striato- 
punctate ; under surface of head strongly alutaceous and rather 
weakly punctate. Pronotal disc 1.3 X as long as its posterior 
width ; mesonotum 1.4 X as long as wide ; propodeum about 1.4 X 
as long as wide. Pronotal disc weakly alutaceous and with fairly 
strong punctures except medially, where it is smooth and shining. 
Mesonotum and propodeum both wholly but rather weakly alu- 
taceous, both with a number of fairly strong punctures on the 
sides; punctate lateral parts of propodeum actually as large in 
area as median impunctate strip. Mesopleurum alutaceous, 
weakly punctate. Hairs of body and legs numerous, pale, of 
moderate length. 

Other females. - - PENNSYLVANIA : 1, Philadelphia, 20 
May 1939 (W. L. Brown, from nest of Proceratium sp.) [USNM]. 
NORTH CAROLINA: 1, Durham, June 1945, Duke forest 
(A. S. Pearse) [INHS]. 

Variation in females. — The females from Pennsylvania and 
North Carolina are slightly smaller than the one from Missouri 
(head length .88 and .82 mm., thorax length 1.6 and 1.5 mm., 
respectively). The resemblance to the Missouri specimen is very 
close in every respect, including measurements. 

Remarks. — The females assigned here tentatively have much 
in common with prolongatum, and it is possible that they fall 
within the range of variation of that species. However, on the 
basis of the available material they do appear to show constant 
differences from prolongatum and to inhabit a generally more 
southerly range, suggesting that they may be the females of 
arenarium (Map 2). It is interesting to note that two of the 
three records for this species indicate an association with poner- 
ine ants, while several records for prolongatum suggest a relation- 
ship with formicine and myrmicine ants. 



266 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

24. Pseudisobrachium prolongatum (Provancher) 

Bethylus prolongatus Provancher, 1881, Nat. Canad., 12: 265. [Type: $ 

(not female as stated), Cap Eouge, Que. (Quebec Prov. Museum, yellow 

label no. 944)]. 
Perisemus prolongus Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 72. 
Isobrachium magnum Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 36. [Type: 

$ , Spokane Falls, Wash. (USNM no. 10068)]. New synonymy. 
Isobrachium myrmecophilum Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 37. 

[Type: 9 , Beatty, Pa. (USNM no. 10069) (S allotype misassociated)]. 

New synonymy. 
Isobrachium mandibular e Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 38. 

[Type: 9, Retreat, Haywood Co., N. C. (USNM no. 14046) ($ allo- 
type misassociated)]. New synonymy. 
Isobrachium momtanum Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 39. 

[Type: 9, Helena, Mont., April 30, from nest of Formica rufibarbis 

(H. G. Hubbard) (USNM no. 14047) ($ allotype misassociated)]. 

New synonymy. 
Pseudisobrachium montanum Kieffer, 1908, Genera Insect., 76: 24. — Kief- 

fer, 1914, Das Tierreich, 41: 479. — Fouts, 1928, Proc. Ent. Soc. Wash., 

30: 122. 
Pseudisobrachium magnum Kieffer, 1908, Genera Insect., 76 : 24. — Kieffer, 

1914, Das Tierreich, 41: 479. — Fouts, 1928, Proc. Ent. Soc. Wash., 

30: 122. 
Pseudisobrachium myrmecophilum Kieffer, 1908, Genera Insect., 76: 24. 

—Kieffer, 1914, Das Tierreich, 41:479. —Fouts, 1928, Proc. Ent. Soc. 

Wash., 30:122. 
Pseudisobrachium mandibulare Kieffer, 1908, Genera Insect., 76: 24. — Kief- 
fer, 1914, Das Tierreich, 41: 480. —Fouts, 1928, Proc. Ent. Soc. 

Wash., 30: 122. 
Pseudisobrachium rugosulum Fouts, 1928, Proc. Ent. Soc. Wash., 30: 124. 

[Type: $ , Mount Holly Springs, Pa., Aug. 12, 1920 (swept from wheat 

stubble, R. Fouts) (USNM no. 62551)]. New synonymy. 
Pseudisobrachium agilis Whittaker, 1928, Trans. Ent. Soc. London, 76: 386. 
[Type: S, Chilliwack, Br. Col., Aug.-Oct., 1926-27 (O. Whittaker) 

(British Museum)]. New synonymy. 
Pseudisobrachium prolongatus Krombein, 1958, U. S. Dept. Agri. Monogr. 2, 

first suppl., p. 97. 
Plesiotype. — $ , "R.C." [Rideau Canal, Ottawa, Ont.], Aug. 
22, 1894 (Harrington) [CNC]. 

Description. — Length 4.3 mm. ; LFW 3.7 mm. Head and 
thorax piceous, abdomen shining brown, on sides of basal seg- 
ments light yellowish-brown ; apical half of mandibles yellowish- 
brown, teeth rufous; antennae eastaneous; front coxae brown, 
legs otherwise bright yellowish-brown; fore wing very faintly 
tinged with brown, veins and stigma dark brown. Mandibles 



EVANS : REVISION OP PSEUDISOBRACHIUM 267 

with five teeth, third and fourth teeth small and close together, 
fifth tooth broad, its margin continuous with that of inner man- 
dibular margin (Fig. 21). Antennae with first four segments in 
a ratio of about 23 :6 :11 :11, segment three 1.7 X as long as thick, 
segment nine 1.5 X as long as thick; flagellar pubescence pale, 
rather coarse, erect setae numerous, many of them nearly half 
as long as diameter of flagellum. WP .71 X WH, 1.6 X HE ; 
ocelli small, DAO .11 X WF ; OOL 1.26 X WOT ; ocelli in a com- 
pact triangle, front angle less than a right angle. Vertex broadly 
rounded a distance above eyes tops subequal to eye height. Front 
and vertex strongly alutaceous, weakly shining, with weak 
punctures which for the most part are separated by from 1 to 
1.5 X their own diameters. Pronotum alutaceous, with many 
weak punctures. Mesoscutum alutaceous, punctures rather strong 
and numerous, on the sides separated by no more than their own 
diameters ; notauli fairly strong on anterior .6 of mesoscutum ; 
scutellar disc strongly shining. Propodeum 1.5 X as long as wide ; 
median carina long; disc with weak, irregular sculpturing, spir- 
acles elliptical, directed laterad. Mesopleurum with callus con- 
vex, shining, weakly alutaceous ; remainder of mesopleurum alu- 
taceous, anteriorly with strong, close punctures. Discoidal vein 
of fore wing arising a short distance down on transverse median 
vein, pigmented to a distance exceeding length of basal vein 
(Fig. 57). 

Males examined. — QUEBEC : 4, Norway Bay, 26-31 Aug. 
1938 (Shewell, Hobbs) [CNC] ; 1, Aylmer, 18 Sept. [CNC]. 
ONTARIO: 1, Rideau Canal, Ottawa, 22 Aug. (Harrington) 
[CNC]. NEW BRUNSAVICK: 2, St. John, 9-18 Sept. (A. G. 
Leavitt) [USNM]. NOVA SCOTIA: 49 ^, Bridgetown, 2 
Sept,-1 Oct, (G. E. Sanders) [CNC]. MAINE: 1, Bar Harbor, 

1 Oct. 1941 (A. E. Brower) [USNM] ; 1, Stratton, Franklin Co., 
19 Aug. 1945 (J. C. Bradley) [CU] ; 1, Houghton, 18 Aug. 
1945 (J. C. Bradley) [CU]. RHODE ISLAND: 1, Westerly, 8 
Sept. 1937 (M. Chapman) [HKT]. NEW YORK: 1, Oneonta, 

2 Sept, 1935 (II. K. Townes) [HKT] ; 1, Lake George, 30 Aug. 
1893 (J. L. Zabriskie) [MCZ]. NEW JERSEY: 1, Princeton, 
28 Sept, 1945 (K. W. Cooper) [USNM]. MARYLAND: 96, 
Plummer's Isl., 29 Aug.-26 Sept. 1958-60, one on Solidago (K. V. 
Krombein, H. E. Evans) [MCZ, USNM, KVK] ; 7, Takoma 
Park, July-Oct. (H. & M. Townes) [HKT]. DISTRICT OF 
COLUMBIA: 1, Washington, 8 Sept. 1952 (R. Boettcher) 
[USNM]. WEST VIRGINIA: 1, Bolivar, 19 Sept. 1942 (H. K. 
Townes) [HKT]. NORTH CAROLINA: 12, Mt. Pisgah, 5000- 
5749 feet, 2-5 Sept. (II. & M. Townes) [HKT]; 14, Crabtree 



268 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Mds., Yancey Co., 21 Aug. 1950 (H. & M. Townes) [HKT] ; 4, 
Mt. Mitchell, 4000-6500 feet, 17-26 Aug. 1950 (H. & M. Townes) 
[HKT]; 2, Hamrick, 17 Aug. 1950 (H. & M. Townes) [HKT]. 
IOWA: 1, Sioux City (C. N. Ainslie) [USNM]. WASHING- 
TON: 1, Tenino, 25 Sept. 1897 (A. P. Morse) [USNM]; 1, 
Spokane Falls [USNM] ; 1, Union Gap, 27 July 1942 (L. J. 
Lipovsky) [KU]. BRITISH COLUMBIA: 1, Chilliwack, Aug.- 
Oct. 1926-7 (O. Whittaker) [Coll. R. M. Fouts]. 

Variation in males. — The 207 males examined exhibit a size 
range from 3.0 to 5.5 mm., fore wing 2.5 to 4.1 mm. The legs 
are bright yellowish-brown in the Ottawa plesiotype described 
above, in the single male from Iowa, and in most specimens from 
the Atlantic coastal plain from Maine to District of Columbia; 
otherwise they tend to be suffused with brown, most particularly 
in specimens from the Pacific Northwest and the Southern Appa- 
lachians. As shown in Table IV, the ocellar triangle tends to be 
less far removed from the eyes in specimens from the Pacific 
Northwest, most far removed in specimens from the southern 
Appalachians (but exceptions are not uncommon). In occasional 
specimens from the Northeast and from the Appalachians, the 
mesopleural callus is less prominent and only weakly shining. 
The most striking variation is in the mandibles. In most speci- 
mens the third and fourth teeth are separate, although small 
(as in Fig. 20). In a few specimens (including some from 
eastern Canada, New York, and Maryland) they are connate 
(Fig. 21). In eight of the specimens examined, these teeth are 
completely fused so that the mandibles are in fact four-toothed 
as they are in members of the species-groups which follow (Fig. 
22). These eight specimens are from New York (Oneonta), 
Pennsylvania (Mt. Holly Springs, type of rugosulum Fouts), 
and North Carolina (Hamrick and Mt. Pisgah). Dr. I. H. H. 
Yarrow of the British Museum (Natural History) has examined 
the type of agilis Whittaker, from British Columbia, and writes 
that in this specimen the mandibles are four-toothed. In a 
topotypic paratype in the collection of R. M. Fouts, they appear 
to be weakly five-toothed, although the mandibles are worn and 
it is difficult to be certain of this. 

Plesiallotype. — 9, Toronto, Ontario, 24 April 1894 [CNC]. 

Description of female. — Length 4 mm., LH .85 mm., LT 1.7 
mm. Head and thorax castaneous, abdomen light castaneous ; 
mandibles, clypeus, and antennae light castaneous; legs bright 
yellowish-brown. Mandibles with three teeth, basal tooth rather 



EVANS : REVISION OF PSEUDISOBRACHIUM 269 

small (Fig. 39). Clypeus broadly subtmncate, median carina 
strong but not quite reaching margin. Head 1.2 X as long as wide, 
sides weakly arcuate, head width greatest about midway, behind 
this sides more convergent to vertex, which is straight ; occipital 
carina absent dorsally. Eyes indicated by small, pale brown 
spots. Front with an impunctate streak medially, otherwise 
with rather dense, elongate punctures, anteriorly rather strongly 
striato-punctate ; surface between punctures moderately shining, 
somewhat alutaceous ; under surface of head more strongly 
alutaceous, punctures rather evenly spaced. Pronotal disc 1.4 X 
as long as its posterior width ; mesonotum 1.3 X as long as wide ; 
propodeum 1 .5 X as long as wide. Pronotal disc shining, weakly 
alutaceous behind, with abundant small punctures except med- 
ially; mesonotum bare and shining medially, laterally weakly 
alutaceous and with some small punctures ; disc of propodeum 
strongly polished, with a few weak punctures on the extreme 
sides; spiracles circular, directed dorso-laterally. Mesopleurum 
strongly alutaceous on sides, weakly punctate. Body and legs 
with numerous pale setae. 

Females examined. — ONTARIO : 1, Toronto, 24 April 1894 
[CNC] ; 1, Eastern part [CNC]. NEW HAMPSHIRE: 1, Pike, 
on Picea (E. J. Kraus) [USNM]. MASSACHUSETTS: 2, 
Lexington, 5 June 1955 (in nests of Aeanthomyops, W. L. 
Brown) [MCZ] ; 3, Forest Hills, May (Mann, Williams) [US- 
NM] ; 1, Lynn Woods Res., 12 Oct. 1949, (rotten log, K. 
Christiansen) [USNM]. CONNECTICUT: 1, West Rock Ridge, 
New Haven, in moss, 24 June 1950 (P. Bellinger) [USNM]. 
NEW YORK: 1, Tuxedo, 30 May 1925 (in clump of grass, Wm. 
T. Davis) [MCZ] ; 1, Ithaca, 1 Oct. 1957 (soil sample, E. F. 
Menhinick) [CU]. PENNSYLVANIA: 2, Beatty [USNM, 
MCZ]. DELAWARE: 1, Christiana, 31 May 1953 (tree hole, 
R. S. Howard) [MCZ]. DISTRICT OF COLUMBIA: 6, Wash- 
ington, May, July, Aug., one in nest with Camponotus penvsyl- 
vanicus [USNM]. VIRGINIA: 1, Falls Church, IS May (N. 
Banks) [MCZ] ; 1, Vienna, 25 Sept. 1927 (leaf litter, ' J. C. 
Bridwell) [USNM] ; 2, Occoqnan, 23 April 1925 (W. M. Mann) 
[USNM]. NORTH CAROLINA: 1, Duke Forest, Durham, 14 
April 1945 (in litter, A. S. Pearse) [INHS] ; 1, 4 mi. N. 
Cherokee, 2000 ft., 29 May 1957 (Berlese sample, W. R. M. 
Mason) [CNC]; 1, Haywood Co. [USNM]. KENTUCKY: 1, 
Bowen, 8 May 1947 [INHS]. ILLINOIS: 1, Oakwood, 29 Sept. 
1933 (in old hickory log, II. II. Ross) [TNHS] ; 1, Little Grassy 
Lake, Williamson Co., 10 Aug. 1958 (in litter, W. L. Brown) 



270 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



[MCZ]. IOWA: 2, Iowa City [USNM]. NORTH DAKOTA: 
1, Walsh Co., 23 June 1950 (with Formica sp., W. E. LaBerge) 
[USNM]. MONTANA: 1, Helena, 30 April (from nest of 
Formica rufibarbis, II. G. Hubbard) [USNM] ; 1, Assinniboine, 
April (with Formica rufibarbis) [USNM]. 

Variation in females. — Of the 35 females before me, the 
smallest is 3.4 mm. long, head .68 mm., thorax 1.3 mm.; the 
largest is 5.2 mm. long, head 1.0 mm., thorax 1.9 mm. The color 
of the head and thorax varies from dark castaneous to pale 
castaneous, in the latter case scarcely any darker than the 
abdomen. The specimen described above is about average in 
both size and color. In some specimens the head is somewhat more 
parallel-sided, head length about 1.3 X head width. The mandibles 
of most specimens resemble Figure 39, but in some the third 
tooth is relatively weak, and in a few the inner mandibular 
margin is sufficiently undulate so as to suggest a very weak 
fourth tooth (as in Fig. 37). 

Remarks. — I have studied the types of Ashmead's species 
myrmecophilum, mandibular 'e, and montanum and can find no 
noteworthy differences between them. The evidence that these 




Map 3. — ■ Distribution of P. prolongation, males indicated by solid circles, 
females by hollow circles. Distribution of P. flainnervis, males indicated by 
solid triangles, supposed female by a hollow triangle. 



EVANS : REVISION OF PSEUDISOBRACHIUM 271 

females go with prolong atum is purely circumstantial, based 
largely on a coincidence of ranges (Map 3) plus the rather large 
size of both males and females as compared with sympatric 
species. While I am reasonably sure that this association of 
sexes is correct, it is to be hoped that males and females will 
someday be taken in closer association than they have so far. 

I have not studied the type of Provancher's prolongation, 
but 0. Peck, K. V. Krombein, and W. R. M. Mason have all 
seen the type and sent me their notes on it, Dr. Mason speci- 
fically checked several characters and compared the specimen 
with drawings which I sent him. The type has five-toothed 
mandibles and in every way compares closely with the plesiotype 
described above. The type is lacking the abdomen, hind legs, 
middle legs except coxae, right fore wing, and tips of antennae. 

This is the most northerly in distribution of any species of 
the genus, the more southerly records (North Carolina) all 
being from moderate to high altitudes. It is partially sympatric 
with ashmeadi and to a more limited extent with several other 
species, but in the colder parts of its range it is the only repre- 
sentative of the genus. The relatively large size of the species 
plus the fact that it is reasonably common in the northeastern 
United States and eastern Canada probably accounts for its 
having been described so many times. 

Carbonarium Species-group 

To this group are assigned nine species, two of them relatively 
common eastern species, the remaining seven apparently un- 
common species occurring in western United States, Mexico, and 
Central America. Not only are species differences in this group 
decidedly unspectacular, but the group as a whole is only weakly 
separable from specimens of the preceding group having four- 
toothed mandibles and from certain species of the rufiventre 
group, particularly flaviventre. This is the most difficult section 
of the genus, and I have sometimes been rather arbitrary in de- 
ciding what to consider a species, particularly with respect to the 
rather limited western material. 

25. Pseudisobrachium carbonarium (Ashmead) 

TLpijris carbonarium Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 59. [Type: 

$ , Washington, D. C. (TJSNM no. 14063)]. 
Holepyris carbonarius Kieffer, 1906 [In Andre, Spec. Hymen. Eur., 9: 341]. 
— Kieffer, 1914, Das Tierreich, 41: 388. 



272 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Pseudisobrachiiim carbonarius Krombein, 1958, U. S. Dept. Agri. Monogr. 
2, first suppl., p. 97. 

Plesiotijpc— $, Bowie, Maryland, 27 Aug. 1944 (H. & M. 

Townes) [HKT]. 5 

Description. — Length 4 mm.; LFW 3.3 mm. Head and 
thorax pieeous, abdomen shining reddish brown, paler basally 
and apically ; mandibles yellowish-brown, teeth rufous ; antennae 
wholly bright yellowish-brown with a tinge of rufous; legs bright 
yellowish-brown except middle and hind coxae slightly darker 
and fore coxae nearly pieeous ; wings subhyaline, stigma brown, 
veins amber. Mandibles with four teeth, the basal tooth broad, 
not separated from inner margin (as in Fig. 23). Clypeus with 
its truncate apical margin about equal to length of third antennal 
segment. Antennae with first four segments in a ratio of about 
26 :6 :10 :10, segment three and segment eleven each about 1.4 X 
as long as thick; flagellar pubescence pale, moderately coarse, 
erect setae numerous but mostly less than half as long as thick- 
ness of flagellum. Front very broad, WP .74 X WH, 1.8 X HE ; 









TABLE V 








Species and locality 


No. 


LFW 


WF/HE 




00L/W0T 


DAO/WF Ant. 11 L/W 


carbonarium 
















Maryland and D . C . 


19 


2.7 (2.4-3.3) 1 


.80 (1.67-1 


.94) 


1.56 (1.45-1.70) 


.11 (.10-. 13) 1 


.3 (1.2-1.5) 


Virginia 


5 


2.7 (2.4-3.0) 1 


.85 (1.78-1 


.91) 


1.57 (1.50-1.71) 


.11 1 


.3 


West Virginia 


n 


2.6 (2.4-2.8) 1 


.80(1.78-1 


.82) 


1.58 (1.54-1.62) 


.12 1 


.4 


Table Rock, S. C. 


14 


2.7 (2.1-3.2) 1 


.80 (1.73-1 


.90) 


1.62 (1.53-1.69) 


.13 (.12-. 14) 1 


.4 


Greenville, S. C. 


3 


2.5 (2.3-2.7) 1 


.70 (1.65-1 


.75) 


1.67 (1.64-1.70) 


.12 1 


.3 


Tigervillc, S. C. 


2 


2.9 (2.8-3.0) 1 


.86 (1.80-1 


.92) 


1.57 (1.54-1.60) 


.12 1 


.3 


Pinnacle Pk. , Ga. 


1 


3.0 1 


.95 




1.76 


.10 1 


.4 


Elgin, Ala. 


1 


2.5 1 


.78 




1.50 


.12 1 


.3 


Illinois 


7 


2.5 (2.2-3.0) 1 


.79(1.70-1 


.93) 


1.57(1.44-1.74) 


.12 (.11-. 13) 1 


.5(1.4-1.6) 


Onaga, Kansas 


1 


3.0 1 


.84 




1.46 


.13 1 


.5 


minimum 
















Grant Co., N. Mex. 


1 


2.3 


1.47 




1.20 


.14 


1.0 


Cochise Co., Ariz. 


1 


3.0 


1.70 




1.35 


.12 


1.2 


Yavapai Co., Arlx. 


1 


2.3 


1.58 




1.38 


.13 


1.2 


mlnutlsslmum 
















Las Cruces, N. Mex. 


1 


1.7 


1.50 




1.17 


.15 


1.0 


Tucson, Ariz. 


2 


1.5 (1.3-1.7) 


1.72 (1.63-1 


.76) 


1.47 


.12 


1.0 


Punta Lobos, Baja Cal . 


1 


1.6 


1.65 




1.44 


.12 


1.1 


Cuernavaca, Morelos 


1 


1.9 


1.57 




1.26 


.12 


1.1 


Yepocapa, Guatemala 


1 


2.0 


1.53 




1.52 


.13 


1.2 


flsvlcornls 
















La Celba, Honduras 


2 


2.4 (1.9-2.6) 


1.49 (1.46-1 


.52) 


1.24 (1.18-1.30) 


.14 


1.0 


Granada, Nicaragua 


1 


2.7 


1.50 




1.22 


.14 


1.0 


navajo 
















Coconino Co. , Ariz. 


1 


2.8 


1.48 




.94 


.14 1 


.2 


Yavapai Co. , Ariz. 


1 


2.7 


1.61 




1.03 


.14 1 


.2 


hurdl 
















Canutillo, Durango 


6 


2.9 (2.6-3.2) 


1.82 (1.75-1 


.90) 


1.15 (1.06-1.23) 


.13 (.12-. 14) 1 


.1 (1.0-1.3) 


krombeinl 
















Albuquerque, N. Mex. 


1 


2.9 


1.52 




1.06 


.15 1 


.3 


White Sands, N. Mex. 


1 


2.7 


1.48 




1.00 


1 


.3 


6 I have studit 


Ml 


the type specimen oi 


' th 


is species, l>nt 


it is in rather 


poor 


condition : I have therefore eleeti 


Ml to b; 


:isc 


my description 


on ;i specimen 


com- 



pared with the type. 



EVANS : REVISION OF PSEUDISOBRACHITJM 273 

ocelli small, in a compact triangle far removed from eyes, DAO 
.11 X WF, OOL 1.46 X WOT. Distance from tops of eyes to 
vertex crest slightly greater than HE. Front weakly shining, 
very strongly alntaceons, actually beaded in appearance, punc- 
tures very shallow and inconspicuous. Pro- and mesonota, 
including scutellar disc, also strongly alutaceous and obscurely 
punctate ; notauli present on anterior .4 of mesoscutum. Pro- 
podeum about 1.35 X as long as broad, disc wholly covered with 
tine, reticulate ridges, median carina strong; sides of propodeum 
also with fine sculpturing; spiracles elliptical, directed dorsad. 
Mesopleurum wholly strongly alutaceous, callus scarcely elevated 
or differentiated ; anterior part of mesopleurum with shallow 
punctures. Discoidal vein of fore wing arising a short distance 
down on the transverse median vein, strong basally, then weak- 
ened but extending as a pigmented line for a distance greater 
than length of basal vein (much as in ashmeadi, Fig. 58). 

Specimens examined. — MARYLAND: 1 $, Bowie, 27 Aug. 
1944 (H. & M. Townes) [HKT] ; 5 $ $ , Takoma Park, 7-11 
Sept. 1942-43 (H. & M. Townes) [HKT] ; 12 $ $ , Plummer's 
Island, Sept.-Oct. (Krombein, Viereck, Evans) [USNM, KVK, 
MCZ]. DISTRICT OF COLUMBIA: 1 $, Washington [type, 
USNM]. VIRGINIA: 2 $ $ , Rosslyn, Aug., Nov. [USNM]; 
1 $ , Falls Church, Aug. [USNM] ; 1 6 , Dunn Loring, Aug. 
[HKT] ; 1 $ , Vienna, June (J. C. Bridwell) [USNM]. WEST 
VIRGINIA: 1 S , Cheat Mts., June [CM] ;H, Philippi, Sept. 
(G. E. Wallace) [CM]. SOUTH CAROLINA: 14 S $ , Table 
Rock, 17 Aug. 1952 (G. & L. Townes) [HKT] ; 3 $ $ , Green- 
ville, Aug., Oct. [HKT] ; 2 S $ , Tigerville, 26 Aug. 1930 (Oman, 
Tuthill) [KIT]. GEORGIA: 1 $, Pinnacle Pk., Rabun Co., 20 
Aug. 1913 [CU]. ALABAMA: 1 «J , Elgin, 6 July 1939 (R. H. 
Beamer) [KU]. ILLINOIS: 2 $ $, Marshall, 27 Sept. 1934 
(Frison & Ross) [INHS] ; 2 $ $ , Anvil Rock, 3 Oct. 1934 
(Frison & Ross) [INHS] ; 1 $ , Cave-in -Rock, 2 Oct. 1934 
(Frison & Ross) [INHS] ; 1 $ , White Heath, 10 Sept. 1889 (C. 
A. Hart) [INHS] ; 1 $ , Urbana, 7 Aug. 1891 [INHS]. KAN- 
SAS: 1 $, Onaga (Crevecoeur) [KSU]. 

Variation in males. — The 55 males examined vary in size 
from 2.5 to 4.1 mm., fore wing from 2.1 to 3.3 mm. Variation in 
color and in standard measurements (Table V) is unusually 
small for a wide-ranging species. 

Female (assigned here tentatively). — Pittsboro, N. C, 7 
Oct. 1948, Berlese funnel, leaf mould (D. L. Wray) [USNM]. 



274 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Description of female. — Length 3.1 mm., LH .60 mm., LT 
1.1 mm. Head rufo-castaneous, thorax castaneous, abdomen 
light yellowish-brown ; mandibles, clypeus, and antennae light 
castaneous, legs wholly light yellowish-brown. Mandibles as 
shown in Figure 37 ; clypeus broadly truncate apically and 
with median carina strong. Head 1.35 X as long as wide, sides 
subparallel but actually very weakly converging almost to poster- 
ior margin, where they are arcuately convergent to a broad, 
straight vertex. Eyes small, not contrasting to head, barely 
distinguishable. Front wholly alutaceous, though less strongly 
so above than antero-laterally ; punctures rather elongate, sep- 
arated from one another by about or less than their own maximum 
diameters ; under side of head strongly alutaceous, weakly punc- 
tate. Pronotal disc 1.3 X as long as its posterior width, mesono- 
tum 1.4 X as long as broad, propodeum 1.4 X as long as broad. 
Pronotum sparsely punctate, weakly alutaceous, though barely so 
in median area; mesonotum wholly weakly alutaceous, weakly 
punctate on sides ; propodeal disc wholly alutaceous, though very 
weakly so antero-medially, sides weakly punctate. Mesopleurum 
laterally strongly alutaceous, weakly punctate. Body hairs pale, 
mostly rather short, abundant over most of body and legs. 

Other females. — NORTH CAROLINA: 1, Fayetteville, 8 
May 1949 (leaf mould, D. L. Wray) [USNM]. KENTUCKY: 
1, Bowen, 8 May 1947 [INHS]. NORTH DAKOTA: 1, Minot, 
21 May 1954 (wheat stubble, C. Benton) [USNM]. 

Variation in females. — The females from Kentucky and North 
Dakota are nearly identical in size to the specimen described 
above; the Fayetteville, N. C, specimen is slightly larger (head 
length .63 mm., thorax length 1.2 mm.). The latter specimen 
has the head only 1.3 X as long as wide, while in the Kentucky 
and North Dakota specimens it is fully 1.4 X as long as wide. 
The Kentucky specimen has the head rather distinctly striato- 
punctate antero-laterally and the eyes are more distinct than 
in the other specimens. 

Remarks. — The females associated here tentatively are very 
similar to those of rufiventre. They are more heavily alutaceous 
than typical rufiventre females and also have a more elongate 
head ; both these features also separate the males of these two 
species and suggest this association of the females. 



EVANS: REVISION OF PSEUDISOBRACHIUM 



275 









TABLE VI 










Species and locality 


Mo. 


LFW 


WF/HE 




00L/W0T 


DAO/WF 


Ant. 11 1 


w 


ashmeadl 


















Marmora, Ont. 


1 


2.7 


1.42 




1.00 


.16 


1.4 




Massachusetts 


4 


2.2(1.9-2.6) 


1.63(1.56- 


1.68) 


1.32(1.23-1.38) 


.15(.14-.16) 


1.3(1.2- 


.4) 


E. Hartford, Conn. 


3 


2.8(2.6-3.0) 


1.61 (1.48- 


1.63) 


1.24(1.16-1.30) 


.13(.12-.15) 


1.3(1.1-: 


.4) 


Long Island, N. Y. 


5 


2.5(2.2-2.9) 


1.77(1.71- 


1.94) 


1.43(1.33-1.72) 


.12(.10-.14) 


1.2(1.1-1 


.3) 


Upstate New York 


14 


2.7(2.3-3.3) 


1.54(1.40- 


1.64) 


1.20(1.00-1.40) 


.14 ( .13-. 16) 


1.3(1.2-1 


■ 4) 


Cassvllle, N.J. 


I 


2.6 


1.76 




1.41 


.12 


1 .3 




Takoma Park, Md. 


20 


2.5(2.0-3.0) 


1.60(1.46- 


1.80) 


1.24(1.17-1.31) 


.13(.12-.14) 


1.2(1.0-1 


.3) 


Washington, D.C. and vie. 


27 


2.5(2.0-2.9) 


1.67(1.47-' 


.80) 


1.25(1.06-1.42) 


.13 ( . 12-. 15) 


1 .2(1.0-1 


.4) 


Hardy Co., W. Va. 


1 


2.2 


1.61 




1.37 


.13 


1.2 




North Carolina 


8 


2.9(2.1-3.5) 


1.57(1.47-' 


.68) 


1.28(1.10-1.50) 


.14 { .12-. 16) 


1.3(1.0-1 


.4) 


South Carolina 


22 


2.5(1.9-3.1) 


1.53(1.43-1 


.80) 


1.30(1.13-1.53) 


.13 < . T 0- . 1 4) 


1.2(1.0-1 


• 4) 


Georgia 


8 


2.5(2.2-2.9) 


1.60(1.44-1 


.70) 


1.30(1.00-1.50) 


.131. 11-. 15) 


1.2(1.1-1 


• 3) 


Ft. George, Fla. 


1 


2.6 


1.55 




1.50 


.14 


1.1 




Waldo, Fla. 


2 


2.5(2.2-2.8) 


1.42(1.36-1 


.48) 


1.02(1.00-1.04) 


.15 


1.0 




Coleta, Ala. 


4 


2.5(2.2-2.8) 


1.45(1.43-1 


.47) 


1.23(1.15-1.28) 


.15 


1.3(1.1-1 


• 4) 


Fulton, Miss. 


3 


2.7(2.3-3.0) 


1.65(1.58-1 


.77) 


1.36(1.22-1.64) 


.12 (.11-. 13) 


1.2(1.0-1 


■ 4) 


Urania, La. 


1 


2.7 


1.40 




1.14 


.15 


1 .3 




Hamilton Co., Tenn. 


2 


2.2(1.9-2.5) 


1.49(1.45-1 


.53) 


1.12(1.11-1.13) 


.14 


1 .1 (1 .0-1 


.2) 


Gatllnburg, Tenn. 


1 


2.4 


1.66 




1.40 


.13 


1.1 


Bjrberton, Ohio 


2 


2.8(2.4-3.2) 


1.61 (1.57-1 


.65) 


1.18(1.10-1.25) 


.14 


1.3 




Michigan 


2 


2.6 


1.56(1.53-1 


.59) 


1.22(1.17-1.27) 


.15 


1.3 




Illinois 


9 


2.5(2.0-3.0) 


1.55(1.45-1 


.70) 


1.15(1.00-1.30) 


.14(.13-.16) 


1.2(1.0-1 


.4) 


Joplln, Mo. 


1 


3.1 


1.46 




1.00 


.16 


1.3 




Palmyra, Mo. 


2 


2.4(2.3-2.5) 


1.49(1.44-1 


.53) 


1.35(1.30-1.40) 


.14 


1.3(1.2-1 


.4) 


Manhattan, Kansas 


1 


2.0 


1.41 




1.14 


.14 


1.1 




perslmlle 


















Lower Lake, Cal. 


6 


2.9(2.6-3.1) 


1.58(1.42-1 


• 76) 


1.00(.90-1.13) 


.16(.13-.17) 


1.5(1.4-1 


■ 6) 


Upper Lake, Cal. 


3 


3.2(3.1-3.3) 


1.57(1.54-1 


.60) 


1.10(1.07-1.12) 


.15(.14-.16) 


1.6 




Soda Bay, Lake Co., Cal. 


1 


2.8 


1.60 




1.13 


.15 


1.5 




Colusa Co. , Cal. 


1 


2.5 


1.57 




.86 


.16 


1.4 




Davis, Cal. 


1 


2.7 


1.55 




.91 


.18 


1 .6 




Sacramento, Cal. 


1 


3.5 


1.70 




.88 


.15 


1.4 





26. Pseudisobrachium AsiiMEADi new species 

Isobradliium myrmecophilum Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 
37. [£ allotype, Washington, D. C. (USNM) ; 9 holotype = P. 
prolongatum Prov.]. 
Isobrachium mandibular -e Ashmead, 1893, ibid., p. 38. [ $ allotype, Ft. 
George, Fla. (USNM); 9 holotype = P. prolongatum Prov.]. 

Holotype. — $ , Washington, D. C., 6 Sept. 1952 (Richard 
Boettcher) [USNM no. 65154]. 

Description. — Length 4.2 mm. ; LFW 2.8 mm. Head piceous; 
thorax dark reddish-brown ; abdomen castaneous, somewhat paler 
basally and apically; mandibles yellowish, teeth rufous; scape 
yellowish-brown, flagellum similar but with a tinge of rufous ; legs 
wholly yellowish-brown ; veins and stigma brown. Mandibles 
with four teeth, third tooth small, basal tooth broad, confluent 
with inner margin of mandible (Fig. 23). Clypeus with its 
truncate apical margin about as long as third antennal segment ; 
median clypeal carina straight in profile. Antennae with first 
four segments in a ratio of about 22 :5 :8 :8, segment three and 
segment eleven each about 1.3 X as long as thick; flagellar 
pubescence pale, coarse, erect setae numerous, some of them half 
as long as width of flagellum. WF .69 X WH, 1.58 X HE ; ocelli 



276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

small, forming a triangle the front angle of which is less than a 
right angle; OOL 1.32 X WOT. Distance from tops of eyes to 
vertex crest subequal to eye height (Fig. 1). Front weakly 
shining, alutaceons though scarcely beaded as in carbonarium; 
punctures shallow though clearly evident, separated from one 
another by about their own diameters. Pronotum moderately 
shining, alutaceous, punctate. Mesoscutum also moderately shin- 
ing, punctures small ; notauli distinct on anterior .4 ; scutellar 
disc strongly shining. Propodeum 1.4 X as long as wide, disc 
shining, with reticulate sculpturing in front which is nearly 
absent behind ; median carina well developed ; spiracles small, 
nearly circular, directed dorsad. Mesopleurum with callus moder- 
ately convex, shining but distinctly alutaceous ; remainder of 
mesopleurum only weakly shining, with large, shallow punctures. 
Discoidal vein weakly pigmented for a distance about equal to 
basal vein (Fig. 58). Genitalia as shown in Figure 63. 

Paratypes. — ONTARIO : 1 S, Marmora, 16 Aug. (J. Mc- 
Alpine) [CNC]. MASSACHUSETTS: 2 S $ , Holliston, Aug. 
(N. Banks) [MCZ] ; 1 $ , Forest Hills, 12 Oct, 1910 [MCZ] ; 1 
8 , S. Natick, 2 Sept. 1940 (J. C. Bradley) [CU]. CONNECTI- 
CUT: 3 $ $, East Hartford, 3-5 Sept. 1947 (sweeping, H. E. 
Evans) [CU, MCZ]. NEW YORK: 1 $, Poughkeepsie, 25 
Aug. 1936 (H. K. Townes) [HKT] ; 1 $ , RensTville, (K. W. 
Cooper) [USNM] ; 2 $ S , Hancock, 10 Aug. 1935 (H. K. 
Townes) [HKT] ; 1 $ , Freeville, Tompkins Co., 1 Sept. 1922 
[CU] ;1 $ , McLean Bogs, Tompkins Co., 30 Aug. 1953 (sweep- 
ing grass, H. E. Evans) [CU] ; 5 $ $ , Ithaca, Aug.-Sept. (at 
light, sweeping) [CU, MCZ] ; 1 $ , Minetto, 21 Aug. 1938 (W. 
T. M. Forbes) [CU] ; 1 £ , Buffalo, 14 Oct. (M. C. Van Duzee) 
[CAS] ; 1 $ , DeBruce, 23-26 Aug. 1912 [AMNH] ; 1 $ , 
Farmingdale, 29 Aug. 1938 (H. K. Townes) [HKT] ; 1 $ , 
Bethpage, Aug. 1938 (F. S. Blanton) [CU] ; 2 S S , Babylon, 
Aug., Oct. (F. S. Blanton) [CU, USNM] ; 1 $ , Selden, L. I., 
1 Oct. 1934 (sweeping flowers, F. S. Blanton) [USNM]. NEW 
JERSEY: 1 S, Cassville, 18 Aug. 1910 [AMNH]. MARY- 
LAND: 20 £ $, Takoma Park, July-Sept., 1942-43 (H. & M. 
Townes) [HKT, MCZ]; 13 $ $ , Plummer's Island, 31 Aug.- 

5 Oct., one on Solidago (K. Y. Krombein, H. E. Evans) [MCZ, 
USNM, KVK]. DISTRICT OF COLUMBIA: 2 6 6, Wash- 
ington, 6-8 Sept. 1952 (R. Boettcher) [USNM]. VIRGINIA: 

6 $ $ , Vienna, Aug. 1932 (J. C. Bridwell) [USNM] ; 1 $ , 
Clifton, Aug. 1932 (J. C. Bridwell) [USNM] ; 1 $ , Dunn 



EVANS : REVISION OF PSEUDISOBRACHIUM 277 

Loring, 28 Aug. 1949 (K. V. Krombein) [HKT] ■ 3 S 6 , Falls 
Church, Aug.-Oct. (N. Banks) [MCZ] ; 1 $ , [no specific 
locality] under stone, 22 Aug. 1895 [USNM]. WEST VIR- 
GINIA : 1 S , Lost River State Park, Hardy Co., 29 July-11 Aug. 
1957 (KVK) [KVK]. NORTH CAROLINA: 1 $, Pink Beds, 
23 July 1952 (G. & L. Townes) [HKT] ; 1 $ , Mooresville, 12 
Sept, 1949 (H., M. & G. Townes) [HKT]; 1 $, Smoky Mts., 
Bryson City, 2000 feet, 25 Aug. 1930 (F. Carpenter) [MCZ] ; 
1 $ , Flat Rock, 6 Sept. 1952 (G. & L. Townes) [HKT] ; 1 $ , 
Kill Devil Hills, Dare Co., 3 Aug. 1956 (K. V. Krombein) 
[KVK] ; 2 6 $ , Walnut, 20 Aug. 1930 (P. W. Oman) [KU] ; 
1 $, Hamrick, 29 Aug. 1950 (H., M. & D. Townes) [HKT]. 
SOUTH CAROLINA: 12 $ $ , Greenville, July-Sept. (L. & G. 
Townes) [HKT] ; 6 6 $ , Table Rock, 17 Aug. 1952 (G. & L. 
Townes) [HKT] ; 1 $ , Columbia, 11 Sept. 1951 (L. & G. 
Townes) [HKT] ; 3 $ S , Cleveland, 2 Aug. 1952 (G. & L. 
Townes) [HKT]. GEORGIA: 1 $, Pinnacle Pk., Rabun Co., 
20 Aug. 1913 [CU] ; 1 $, Spring Creek, Decatur Co., 16-29 July 
1912 [MCZ] ; 2 $ $ , Tifton [USNM] ; 1 $ , Prattsburg, 25 
July 1930 (L. D. Tuthill) [KU] ; 1 $ , Perty, 12 Aug. 1939 
(R. H. Beamer) [KU] ; 2 $ $ , Austell, 17 July 1910 [MCZ]. 
FLORIDA: 2 S $, Waldo, 18 Aug. 1930 (R. H. Beamer) 
[KU]. ALABAMA: 4 N, Coleta (H. H. Smith) [USNM]. 
MISSISSIPPI: 3 S 3, Fulton, 14 July 1930 [KU]. LOUISI- 
ANA: 1 S, Urania, 14 July 1943 [INHS]. TENNESSEE: 2 
$ $ , Hamilton Co., 20 Sept. 1939 (Turner) [USNM] ; 1 $ , 
Gatlinburg, 31 Aug.-4 Sept. 1940 (B. D. Burks) [INHS]. 
OHIO : 2 $ 6 , Barberton, 23 Aug. 1936 (L. J. Lipovsky) [KU]. 
MICHIGAN: 1 $, Bay Co., 22 July 1939 (R. R. Dreisbach) 
[RRD]; 1 S, Midland Co., 10 July 1952 (RRD) [RRD]. 
ILLINOIS : 1 $ , Fox L., 13 Aug. 1937 (Ross & Burks) [INHS] ; 

1 $ , Evergreen Park, 23 Aug. 1934 (DeLong & Ross) [INHS] ; 

2 $ $ , Oak Lawn, July, Aug. [INHS] ; 1 $ , Marshall, 27 
Sept. 1934 (Frison & Ross) [INHS] ; 1 $ , Seymour, 5 Aug. 
1939 (Ross & Riegel) [INHS] ; 1 $ , Mahomet, 3 Aug. 1937 
(Ross & Burks) [INHS] ; 1 $ , Principia College, Jersey Co., 2 
July 1943 (C. L. Remington) [USNM] ; 1 $ , Alto Pass, 13 Aug. 
1891 (C. A. Hart) [INHS]. MISSOURI: H,5 mi. N. Joplin, 
5 Aug. 1950 (at light, H. E. Evans) [MCZ] ; 2 S $ , Palmvra, 
16 Sept. 1939 (G. T. Riegel) [INHS]. KANSAS: 1 $, Man- 
hattan, 27 Sept. 1930 (D. A. Wilbur) [KSU]. 

Variation in males. — The 148 males examined vary in size 
from 2.2 to 4.2 mm., with the vast majority between 3 and 4 mm. ; 



278 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

LFW varies from 1.9 to 3.5. In some specimens the head and 
thorax are paler than in the type, and in a few the thorax and 
abdomen are piceous like the head ; in no case is the abdomen 
contrastingly colored as compared to the thorax. The legs are 
yellow-brown in most specimens, but occasional specimens from 
various parts of the range have the legs brownish (though never 
dark brown). There is considerable variation in head shape; 
while in most specimens the head is slightly higher than wide 
and the distance from the eye tops to the vertex crest subequal 
to the eye height, in numerous specimens from various parts of 
the range the head is as wide as or wider than high and the dis- 
tance from the eye tops to the vertex crest notably less than the 
eye height. As suggested in Table VI, most of the variation in 
this species shows little apparent correlation with geography. 
However, there is a tendency for specimens from the southern 
parts of the range to have shorter antennae. As in other species, 
local populations are sometimes rather distinctive in certain 
morphological characters. For example, five specimens from 
Long Island, N. Y., and one from New Jersey, all have the front 
unusually broad and the eyes proportionally very small. 

Female (assigned here "tentatively) . — MASSACHUSETTS : 
Arlington, 24 May 1953, Formica fusca nest in woods (W. L. 
Brown) [MCZ]. 

Description of female. — Length 2.9 mm., LH .50 mm., LT 
.95 mm. Head castaneous, thorax light castaneous, abdomen 
bright yellowish-brown ; mandibles, clypeus, and antennae light 
castaneous ; legs straw-colored. Mandibles with a weakly indi- 
cated fourth tooth (Fig. 38). Clypeus weakly emarginate apic- 
ally, median carina strong. Head 1.33 X as long as wide, sides 
subparallel, posteriorly arcuately convergent to a vertex which 
is weakly emarginate medially. Eyes no larger than a head 
puncture, not contrasting in color to head and barely distinguish- 
able. Front strongly shining, non-alutaceous, punctures very 
strong, separated by from 1-2 X their own diameters, except 
absent along median strip ; under side of head shining, weakly 
alutaceous, weakly punctate. Pronotal disc 1.4 X as long as its 
posterior width ; mesonotum about 1.5 X as long as wide ; pro- 
podeuni also about 1.5 X as long as wide. Pronotum strongly 
shining, non-alutaceous, with fairly strong, widely separated 
punctures except along median strip, which is bare. Mesonotum 
strongly shining and non-alutaceous except around edges. Pro- 
podeal disc very strongly shining, weakly alutaceous behind 
and with a few punctures on extreme sides. Mesopleurum weakly 



EVANS : REVISION OF PSEUDISOBRACHIUM 



279 



alutaceous, weakly punctate. Hairs of body and legs numerous, 
pale, of moderate length. 

Other females. — MASSACHUSETTS : 2, Forest Hills, one of 
them 4 May 1918 (W. M. Mann) [USNM]. NORTH CARO- 
LINA : 1, Richmond Co., 28 Feb. 1938 (peach orchard, W. F. Tur- 
ner) [USNM]. GEORGIA: 1, Upson Co., 7 March, 1938 (peach 
orchard, W. F. Turner) [USNM] ; 3, Peach Co., May, Aug., Dec. 
(soil of peach orchard, W. F. Turner) [USNM] ; 1, Fort Valley, 
1936 (Christenson) [USNM]. TENNESSEE: 1, Hamilton Co., 
16 Apr. 1940 (peach orchard, W. F. Turner) [USNM] ; 1, Roane 
Co., 22 Nov. 1937 (in soil, W. F. Turner) [USNM]. MISSIS- 
SIPPI: 1, Lincoln Co., 14 July 1936 (W. F. Turner) [USNM]. 

Variation in females. — In the 12 females assigned here, head 
length varies from .47 to .53 mm. (mean .50) ; LH/WH varies 
from 1.26 to 1.33 (mean 1.30) ; thorax length varies from .90 
to 1.0 mm. (mean about .95). Some of the Georgia specimens 
have the body hairs unusually dense and elongate. In the speci- 
mens from Forest Hills, Mass., the under side of the head is 
quite strongly alutaceous, but in most specimens it is only 
weakly so. 

Remarks. — This is apparently one of the commoner species 




Map 4. — Distribution of P. ashmeadi, males indicated by solid circles, 
supposed females by hollow circles. Distribution of P. minimum, males 
indicated by solid triangles (females unknown). Distribution of P. per- 
svmile, males indicated by solid squares, possible female by a hollow square. 



280 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

of the genus. Males have often been misidentified as myrme- 
cophilum, but the type of that species is a female which I feel 
confident goes with Provancher's prolongatum. The females 
which I believe go with ashmeadi show relatively little variation 
in size, color, and body features, although the males exhibit 
considerable variation. It may well be true that the males 
assigned here represent more than one species and that the 
females go with only one of those species. (See Map 4.) 

27. Pseudisobrachium minimum new species 

Holotype. — $ , City of Rocks State Park, Grant Co., N. Mex., 
17 July 1959 (K. V. Krombein) [USNM, no. 65160]. 

Description.- — Length 3.0 mm.; LFW 2.3 mm. Head black; 
thorax dark brownish-fuscous ; abdomen dark, shining brown, a 
little paler on sides of basal segments ; apical two-thirds of man- 
dibles light brown, teeth rufous ; scape brownish-fuscous, flagel- 
lum bright castaneous ; front coxae fuscous, remaining coxae 
and all femora medium brown, rest of legs light brown ; wings 
hyaline, veins and stigma light brown. Mandibles with four 
teeth, basal tooth prominent, somewhat thicker than third tooth 
(Fig. 24). Clypeus with median carina arched, apex narrowly 
truncate. Antennae with first four segments in a ratio of about 
30:9 :10:10, segment three 1.2 X as long as thick, segment eleven 
about as long as thick ; flagellar pubescence pale and moderately 
coarse, erect setae numerous but rather short. WF .71 X WH, 
1.47 X HE ; ocelli small, in a small triangle the front angle of 
which is less than a right angle ; OOL 1.20 X WOT. Distance 
from tops of eyes to vertex crest subequal to eye height. Front 
shining, rather weakly alutaceous, punctures small but numerous 
and well-defined, separated from one another by about or slightly 
more than their own diameters. Pronotum and mesoscutum 
moderately shining, alutaceous, obscurely punctate ; notauli pres- 
ent only on anterior .1 of mesoscutum; scutellar disc weakly alu- 
taceous, shining. Propodeum 1.4 X as long as broad, disc witli 
weak sculpturing over the entire surface, median carina long 
but rather weak ; spiracles small, subcircular, directed dorso- 
laterad. Mesopleurum with callus elongate, weakly alutaceous, 
shining ; anterior portion of mesopleurum with large but rather 
weak punctures. Discoidal vein of fore wing very weakly indi- 
cated by a fainly pigmented line. 

Paratypes. — ARIZONA : 1 S , Southwestern Research Sta- 
tion, 5 mi. W. Portal, Cochise Co., 5400 feet, 23 Aug. 1959 



EVANS: REVISION OF PSEUDISOBRACHIUM 281 

(H. B. Evans) [CU] ; 1 $ , Cottonwood, Yavapai Co., 28 July 
1956, swept from alfalfa (Butler & Gerhardt) [UA]. 

Variation. — The specimen from Cochise County, Arizona, is 
larger and considerably darker than the type, having the legs 
and antennae very dark brown. Although closer to the type in 
size and color, the Cottonwood specimen has the head and thorax 
more heavily alutaeeous and therefore somewhat less shining 
and less distinctly punctate ; in this specimen, also, the clypeal 
carina is more strongly arched. 

Remarks. — The distribution of this species is shown on Map 4. 

28. Pseudisobraciiium minutissimum new species 

Holotype. — S , Punta Lobos, 1 mi. SE of Todos Santos, Baja 
California, 25 Dec. 1958 (H. B. Leech) [CAS]. 

Description. — -Length 2.1 mm.; LFW 1.6 mm. Head black; 
thorax dark brownish-fuscous ; abdomen dark brown, somewhat 
paler basally and apically ; apical three-fourths of mandibles 
straw-colored, teeth rufous ; scape brown, nagellum light brown, 
very slightly darker apically ; coxae and femora medium brown, 
remainder of legs brown ; wings slender, hyaline, veins and stigma 
brown. Mandibles with four teeth, basal tooth slightly larger 
than third tooth (Fig. 26). Clypeus narrowly truncate apically, 
with a median ridge which disappears somewhat before apical 
margin. Antennae with first four segments in a ratio of about 
3 :1 :1 :1, segments three and eleven each barely longer than 
thick ; flagellar pubescence coarse, pale, erect setae numerous, 
mostly less than half as long as thickness of flagellum. WF .69 X 
WH, 1.65 X HE ; ocelli small, in a small triangle the front angle 
of which is less than a right angle; OOL 1.44 X AVOT. Eyes 
rather protuberant laterally, vertex rather narrowly rounded off 
a distance above eye tops slightly greater than eye height (Fig. 
4). Front shining, moderately alutaeeous, punctures small, shal- 
low, separated by about twice their own diameters. Vertex and 
thoracic dorsum more strongly alutaeeous and less shining than 
front ; mesoscutum obscurely punctate, notauli weakly developed 
on anterior .3; disc of scutellum shining. Propodeum 1.4 X as 
long as broad, disc shining, with reticulate sculpturing which is 
strongest basally; spiracles small, subcircular, directed dorso- 
laterad. Mesopleurum weakly alutaeeous, obscurely punctate, 
callus only slightly convex and slightly more shining than re- 
mainder of mesopleurum. Discoidal vein of fore wing evident 
as a very weakly pigmented streak. 



282 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Paratypes. — ARIZONA: 2 $ $ , Tucson, June, October (But- 
ler, Crandall) [UA]. NEW MEXICO: 1 $ , Las Cruces 
[USNM]. MORELOS: 1 $ , 12 mi. E. of Cuernavaca, 14 Aug. 
1954 (U. Kansas Exp.) [KU]. GUATEMALA : 1 $ , Yepocapa, 
1948-49 (H. T. Dalmat) [USNM]. 

Variation. — The six available specimens exhibit considerable 
variation in head measurements, as shown in Table V. The 
Guatemala specimen has the entire body medium brown, the 
wing veins very light brown ; in the two Tucson specimens the 
wing veins are also light brown and in all three of these speci- 
mens the discoidal vein is very faint. The specimen from Las 
Cruces, New Mexico, not only has the lateral ocelli relatively 
rather close to the eyes (OOL 1.17 X WOT), but the entire 
body is pale castaneous, the setulae on the wing pale, and the 
veins and stigma nearly colorless. Also, in this specimen the 
clypeal carina is high and arched basally, rather abruptly de- 
clivous just before the apical margin. Some of the color variation 
in this series may, of course, be due to the fact that some speci- 
mens are teneral ; in any case it is no greater than that found in 
several other species. 

29. Pseudisobrachium flavicornis (Kieffer) new combination 

Holepyris flavicornis Kieffer, 190(5, Berlin. Ent. Zeitschr., 50: 246 [Type: 

$, Granada, Nicaragua (Coll. Baker) (Pomona College, Claremont, 

Calif.)]. 
Lyssepyris flavicornis Kieffer, 1913, Boll. Lab. Zool. Portici, 7: 108 [Made 

type of new genus Lyssepyris]. — Kieffer, 1914, Das Tierreieh, 41: 

396-397. 
Description of holotype. — Length 3.7 mm. ; LFW 2.7 mm. 
Head piceous, thorax dark brown, abdomen medium brown with 
indistinct banding with lighter brown ; mandibles light yellowish- 
brown, teeth rufous ; antennae yellowish-brown, very slightly 
darker apically ; legs wholly bright straw-yellow ; wings hya- 
line, stigma brown, veins light brown. Mandibles with only three 
teeth, second tooth small, basal tooth broad, arching into inner 
mandibular margin (Fig. 28). Clypeus narrowly truncate apic- 
ally, median carina straight in profile. First four antennal seg- 
ments in a ratio of about 9 :2 :3 :3, segment three slightly longer 
than thick, segment eleven about as long as thick ; pubescence 
pale, moderately coarse, about as in ashmeadi. WF .70 X WH, 
1.50 X HE; ocelli small, DAO .14 X WF; OOL 1.22 X WOT; 
ocelli well separated, front angle of ocellar triangle slightly less 
than a right angle. Distance from tops of eyes to vertex crest 



EVANS : REVISION OF PSEUDISOBRACHIUM 283 

equal to slightly less than HE. Front moderately shining, aluta- 
eeous, punctures shallow and inconspicuous, separated from one 
another by slightly more than their own diameters. Pro- and meso- 
nota alutaceous, inconspicuously punctate ; notauli moderately 
strong on anterior .3 of mesoscutum, absent behind. Propodeum 
short, about 1.4 X as long as broad ; disc alutaceous, with strong 
median and lateral carinae ; spiracles small, subcircular, directed 
dorsad. Mesopleural callus large, slightly depressed midway of 
its length, shining and non-alutaceous ; posterior margin of meso- 
pleurum below callus also shining, remainder of this sclerite 
weakly alutaceous and with rather strong punctures. Discoidal 
vein of fore wing weakly pigmented. 

Other males examined. — HONDURAS : 2 A A , La Ceiba, 14 
Aug. and 2 Dec. 1916 (F. J. Dyer) [USNM, AMNH]. 

Variation. — One of the two Honduras specimens [AMNH] 
is strikingly similar to the type in every respect, including size. 
The other specimen is considerably smaller (LFW 1.9) and has 
four-toothed mandibles, about as in ashmeacli; this specimen is 
also generally more w T eakly punctate than the other two. 

30. Pseudisobrachium navajo new species 

Holotype. — A , Tuba City, Coconino Co., Arizona, 27 July 
1954 (at light, H. E. and M. A. Evans) [MCZ, No. 30282]. 

Description. — Length 4.0 mm. ; LFW 2.8 mm. Head black, 
thorax piceous, abdomen dark brown, paler basally and apically ; 
mandibles light brown, teeth rufous ; antennae light brown, 
tinged with rufous beyond third segment; front coxae dark 
brown, legs otherwise light yellowish-brown ; wings hyaline, with 
pale setulae, stigma light brown, veins nearly colorless. Mandi- 
bles with four teeth, basal tooth slightly exceeding third tooth 
(Fig. 25). Clypeus narrowly truncate apically, median carina 
high and arched basally, but abruptly declivous well before 
margin. First four antennal segments in a ratio of about 
20 :6 :7 :7, segment three 1.4 X as long as thick, segment nine 
1.2 X as long as thick (one antenna broken off at segment five, 
the other at segment ten) ; flagellar pubescence coarse, pale, sub- 
erect, erect setae numerous, many of them half as long as width 
of flagellum. WF .71 X WH, 1.48 X HE; DAO .14 X WF; 
ocelli in a moderately broad triangle, the front angle about a 
right angle, OOL .94 X WOT. Vertex broadly rounded, in the 
middle nearly straight across ; distance from eye tops to vertex 
crest equal to .8 X HE. Front strongly alutaceous, somewhat 



284 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

shining, with numerous shallow but well-defined punctures, these 
separated by about or slightly more than their own diameters. 
Pronotum alutaceous, somewhat shining, obscurely punctate. 
Mesoscutum alutaceous and uniformly covered with small but 
distinct punctures; notauli rather weakly developed on anterior 
.3; scutellar disc shining, weakly alutaceous, weakly punctate. 
Propodeum 1.3 X as long as broad, disc with weak reticulate 
sculpturing and well-developed median and lateral carinae ; 
spiracles elliptical, directed dorso-laterad. Mesopleural callus 
convex and shining, remainder of mesopleurum less strongly 
shining, alutaceous, obscurely punctate. Discoidal vein of fore 
wing absent. 

Paratype. — 1 $ , Chino Valley, Yavapai Co., Ariz., 27 July 
1956 (swept from alfalfa, Butler and Gerhardt) [UA]. 

Variation. — In the paratype the basal mandibular tooth is 
slightly broader and shorter than the third tooth and the clypeal 
carina less abruptly declivous than in the type. In this specimen 
the notauli are absent and the mesopleurum more distinctly 
punctate anteriorly. 

31. Pseudisobrachium persimile new species 

Holotype. — S , Lower Lake, Lake Co., Calif., 8 Aug. 1958 
(light trap, R. E. Dolphin) [CAS]. 

Description. — Length 3.3 mm., LFW 3.1 mm. Head and thorax 
black, abdomen dark brown, paler on sides of basal segments ; 
apical half of mandibles light brown ; scape black, flagellum dark 
brown ; legs dark brown except tibiae medium brown, tarsi light 
brown ; wings hyaline, setulae brown, stigma brown, veins light 
brown. Mandibles with third tooth small, fourth tooth only 
slightly broader than third tooth. Clypeus truncate apically, 
median carina not arched in profile. Antennae rather long, first 
four segments in a ratio of about 20 :5 :8 :8, segment three about 
twice as long as thick, segment eleven 1.5 X as long as thick ; 
flagellar pubescence short although suberect, erect setae rather 
prominent, some of them half as long as width of flagellum. \VF 
.71 X WH, 1.67 X HE ; ocelli well separated, front angle of ocel- 
lar triangle less than a right angle, DAO .17 X WF ; OOL sub- 
equal to WOT. Head gradually narrowed behind eyes, vertex 
narrow but nearly straight across; distance from eye tops to 
vertex crest subequal to HE (Fig. 2). Front shining, alutaceous 
although not strongly so, punctures weak and barely visible. Pro- 
notum and mesoscutum also shining, moderately alutaceous, ob- 
scurely punctate ; notauli impressed on anterior half of meso- 
scutum ; disc of scutellum strongly shining. Propodeum elongate, 



EVANS : REVISION OF PSEUDISOBRACHIUM 285 

measuring 1.6 X as long as broad, disc strongly shining though 
with weak reticulate sculpturing, especially anteriorly ; spiracles 
subcircular, directed dorso-laterad. Mesopleurum elongate, cal- 
lus elongate and shining, remainder of mesopleurum somewhat 
alutaceous but without noticeable punctures. Fore wing with 
discoidal vein barely pigmented, arising a short distance down 
on transverse median vein. Abdomen slender. 

Paratype. — CALIFORNIA : 5 $ $ , same data as type but 
three of them 25 July 1958 [UCD, USNM, MCZ] ; 3 $ $ , Upper 
Lake, Lake Co., 25 July, 8 Aug. 1958 (light trap, R. E. Dolphin) 
[UCD, MCZ] ; 1 & , Soda Bay, Lake Co., 25 July 1958 (light 
trap, R. E. Dolphin) [UCD] ; 1 $ , College City, Colusa Co., 
16 July 1959 (light trap, J. Fowler) [UCD] ; 1 $ , Davis, Yolo 
Co., 24 June 1959 (light trap, F. E. Strong [UCD] ; 1 $ , Sacra- 
mento, 20 July 1933 (II. II. Kiefer) [CDAS] ; 1 $, Vacaville, 
Solano Co., 31 July 1954 (light trap, E. Mezger) [UCD]. 

Variation. — ■ The specimen from Vacaville, Davis, and Sacra- 
mento have the front and thoracic dorsum more heavily alu- 
taceous and more distinctly punctate than in the series from 
Lake Co. Variation in standard measurements is not unusual 
(Table VI). 

Female (assigned here tentatively). — Vernon, British Col- 
umbia, 9 May 1948 (E. S. Ross) [CAS]. 

Description of female. — Length 2.8 mm., LH .72 mm., LT 
1.2 mm. Entire body pale yellowish-brown; legs and antennae 
straw-colored. Mandibles with three teeth, the basal tooth weak, 
about as in Figure 41. Clypeus broadly truncate apically. Head 
1.23 X as long as wide, sides subparallel nearly to posterior 
margin, then arcuately convergent to a very broad, straight ver- 
tex. Eyes not discernible. Front wholly rather weakly alu- 
taceous, not strongly shining ; punctures absent from median 
strip, otherwise numerous, separated by about their own diam- 
eters, not especially strong and not notably elongated ; under 
side of head strongly alutaceous, weakly punctate. Pronotal disc 
1.4 X as long as its posterior width ; mesonotum and propodeum 
each about 1.5 X as long as maximum width. Pronotal disc 
weakly alutaceous, with weak, widely separated punctures ; meso- 
notum wholly alutaceous, not strongly shining, obscurely punc- 
tate on sides; propodeal disc weakly alutaceous, weakly punc- 
tate on sides. Mesopleurum strongly alutaceous, dull, obscurely 
punctate. Hair on body and legs short, pale, moderately abun- 
dant. 



286 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Remarks. — The males of this species have a general resem- 
blance to those of the sympatric Occident ale: they are elongate, 
dark in color, and have a similarly shaped head, although longer 
behind the eyes. However, the difference in the mandibular den- 
tition readily places these two species in different species-groups. 
The single female is from a locality far outside the known range 
of the males (Map 4). The only males I have seen from British 
Columbia belong to prolongation, the female of which is well 
known and considerably larger than the female described above. 
I am hesitant to describe this single female as new, and therefore 
assign it to persimile very tentatively on the assumption that 
the range of that species is much wider than present evidence 
indicates. 

32. Pseudisobrachium hurdi new species 

Holotype. — $ , 8 mi. S. of Canutillo, Durango, Mexico, 9 Aug. 
1951 (P. D. Hurd) [CAS]. 

Description. — Length 4.3 mm. ; LFW 2.9 mm. Head piceous, 
thorax and abdomen dark reddish-brown; apical half of mandi- 
bles yellowish, teeth rufous; scape dark brown, flagellum cas- 
taneous; legs medium brown except tarsi light brown; wings 
hyaline, veins and stigma light brown. Mandibles with third 
tooth small, fourth tooth broad and arching into inner mandibu- 
lar margin. Clypeus broadly truncate apieally, its truncate mar- 
gin longer than third antennal segment ; median carina high, 
weakly arched. Antennae with first four segments in a ratio of 
about 22:5:9:9, segment three 1.5 X as long as thick, segment 
eleven 1.2 X as long as thick; flagellar pubescence fine, pale, ap- 
pressed, erect setae mostly less than one fourth as long as width 
of flagellum. WF .72 X WH, 1.85 X HE ; ocelli small, DAO 
.12 X WF, in a broad triangle, front angle of about a right 
angle; OOL 1.22 X WOT. Vertex extended far above eye tops, 
distance from eye tops to vertex crest slightly greater than HE ; 
top of vertex nearly straight across (Fig. 3). Front alutaceous, 
moderately shining, with distinct punctures which are separated 
from one another by from one to two times their own diameters. 
Pronotum and mesoscutum shining, somewhat alutaceous, with 
sparse but distinct punctures; notauli impressed on anterior .3 
of mesoscutum ; scutellar disc strongly shining. Propodeum 1.4 X 
as long as broad, disc shining but wholly alutaceous, median 
carina strong; spiracles elliptical, directed dorso-laterad. Meso- 
pleurum shining, somewhat alutaceous, anteriorly with small 



EVANS : REVISION OF PSEUDISOBRACHIUM 287 

punctures ; callus slightly convex but not otherwise differing from 
remainder of pleurum. Fore wing with discoidal vein interstitial 
with median vein, distinct for a distance about equal to basal 
vein but only very weakly pigmented. 

Paratypes. — 5 $ S , same data as type (P. D. Hurd, H. E. 
Evans) [USNM, CIS, MCZ]. 

Variation. — The five paratypes resemble the holotype closely 
in size and coloration; there is some variation in the intensity 
of punctation on the head and thorax, and in two specimens the 
mesopleural callus is more strongly shining than in the type. 
The variation in measurements of body parts is not great (Table 
V). 

Remarks. — The six known specimens of this distinctive form 
were taken at Coleman lanterns which were placed on the ground 
in semi-arid grassland at about 6000 feet elevation. 

33. Pseudisobrachium KROMBEiNi new species 

jlolotype. — $, Albuquerque, N. Mex., 14-15 Aug. 1959 (at 
light, K. V. Krombein) [USNM, no. 65161]. 

Description. — Length 3.8 mm. ; LFW 2.9 mm. Head black, 
thorax piceous, abdomen dark, shining brown ; apical half of 
mandibles light brown; scape dark brown, flagellum dull brown- 
ish-ferruginous ; front coxae dark brown, remaining coxae and 
all femora medium brown, rest of legs light brown ; wings hya- 
line, setulae pale, veins and stigma light brown. Mandibles with 
third tooth very small, fourth tooth very broad, arching into 
inner mandibular margin (Fig. 27). Apical truncate margin of 
clypeus unusually broad, measuring nearly 1.5 X as long as 
length of third antennal segment (Fig. 50) ; median clypeal 
carina arched in profile. Antennae with first four segments in a 
ratio of about 20:5:8:8, segment three and segment eleven each 
about 1.3 X as long as thick; pubescence of flagellum pale, the 
setulae short but suberect, erect setae sparse and short. WF 
.68 X WH, 1.52 X HE; ocelli small, DAO .15 X WF; ocellar 
triangle broad, front angle a right angle; OOL 1.06 X WOT. 
Distance from tops of eyes to vertex crest equal to about .8 HE ; 
vertex nearly straight across. Front alutaceous, somewhat shin- 
ing, with small but distinct punctures which are separated from 
one another by 1-2 X their own diameters. Pronotum and meso- 
scutum strongly alutaceous but somewhat shining, punctures 
small and rather weak; notauli impressed on anterior .2 of 
mesoscutum ; scutellar disc strongly shining. Propodeum 1.4 X 
as long as broad, disc alutaceous but shining, median carina 
giving rise to a few weak transverse carinae ; spiracles small, 



288 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

elliptical, directed dorsad. Mesopleurum with callus elongate, 
shining and rather weakly alutaceous; anterior ventral part of 
mesopleurum rngoso-punctate. Discoidal vein of fore wing repre- 
sented only by a weak, unpigmented streak. 

Paratype.— 6 , White Sands, N. Mex., 27 June 1940 (D. E. 
Hardy) [KU]. 

Variation. — The single paratype is slightly smaller and paler ; 
the head is piceous, the thorax and abdomen medium brown, the 
legs and antennae light brown. The notauli are slightly longer, 
the mesopleural callus somewhat more alutaceous. Variation in 
head measurements is slight (Table V). The paratype has the 
front of the head somewhat damaged, making measurement of 
the anterior ocellus impossible ; however, the ocelli seem to be 
about the same size as in the type. 

Rufiventre Species-group 

To this group are assigned nine species, of which two are 
eastern and seven chiefly southwestern in distribution. There are 
no real differences between this group and the carbonariiim 
group ; arbitrarily I restrict this group to species with generally 
larger eyes and ocelli. However, this character is quite tenuous, 
as several species exhibit considerable variation in eye and ocellar 
size. Furthermore, flaviventre provides an almost perfect inter- 
mediate. The females, where known, do not differ in any notable 
fashion from those of the carbonarium group. 

34. PsEiiDisoBRACiiiUM flaviventre (Kieffer) new combination 

Eyyrix flaviventris Kieffer, 1904, Ark. Zool., 1: 526. [Type: &, Texas 

(Belfrage) (Naturhist. Kiksnms. Stockholm, no. 228)]. 
Xantepyris flavive?itris Kieffer, 1913, Boll. Lab. Zool. Portici, 7: 108. 
Xanthepyris flaviventris Kieffer, 1914, Das Tierreich, 41: 417. 

Description of holotype. — Length 4.0 mm.; LFW 2.9 mm. 
Head and thorax dark reddish-brown except prothorax somewhat 
lighter, abdomen bright, pale rufous; mandibles yellowish-brown, 
teeth rufous ; scape straw-colored, flagellum light reddish-brown ; 
legs, including all coxae, light straw-colored ; wings hyaline, with 
pale setulae, stigma light brown, veins almost colorless. Man- 
dibles with third tooth small, fourth tooth broad, rounded into 
inner mandibular margin. Clypeus truncate apically, median 
carina straight in profile. Antennae very short, first four seg- 
ments in a ratio of about 21 :5 :7 :6, segment three about 1.2 X 
as long as thick, segment eleven 1.1 X as long as thick ; pubescence 



EVANS: REVISION OF PSEUDISOBRACHUIM 



289 









TABLE VII 








Species and locality 


No. 


LFW 


WF/HE 




OOL/WOT 


DAO/WF 


Ant. 11 L/W 


flavl ventre 
















Dover, Del. 


1 


2.8 


1.33 




.84 


.18 


1.0 


Chestertown, Md. 


1 


2.6 


1.69 




1.13 


.13 


1.0 


Washington, D.C. and vie. 


3 


2.4(2.3-2.6) 


1.64(1.53- 


.72) 


1.26(1.18-1.32) 


.14 (.13-. 15) 


1.0( .9-1.1) 


Raleigh, N. C. 


2 


2.8 


1.34(1.31- 


.37) 


.85 ( .82- .88) 


.18 


1.1 


Aiken, S. C. 


1 


2.2 


1.46 




1.03 


.16 


1.0 


Roberta, Ga. 


2 


2.3 


1.41 (1.35-1 


.47) 


1.00( .97-1.30) 


.17 


1.2 


Prattsburg, Ga. 


1 


2.7 


1.28 




1.20 


.17 


1.0 


Okefenokee Sw., Ga. 


1 


2.3 


1.58 




1.06 


.16 


1.0 


No. Central Fla. 


8 


2.5(2.0-3.0) 


1.49(1.35-1 


.69) 


1.02 ( .86-1.19) 


.17 (.14-. 19) 


1.0 


Sebrlng, Fla. 


1 


2.1 


1.26 




.68 


.19 


0.9 


Okeechobee, Fla. 


1 


2.1 


1.46 




1.12 


.17 


1.0 


Everglades Nat. Pk, Fla. 


1 


2.7 


1.45 




.95 


.19 


1.1 


Cottondale, Ala. 


2 


2.0(1.8-2.2) 


1.34(1.32-1 


.36) 


1.06(1.00-1.12) 


.18 C.17-.19) 


1.1 


Tuscaloosa, Ala. 


8 


2.7(2.1-3.0) 


1.40(1.33-1 


.50) 


.88 ( .83- .96) 


.18 (.15-.21) 


1.1 (1.0-1.2) 


Eastern Texas 


5 


2.6(2.4-2.9) 


1.48(1.43-1 


.50) 


1.01 ( .95-1.07) 


.16(.15-.17) 


1.0 


Scott Co. , Kansas 


1 


2.6 


1.58 




1.03 


.15 


1.1 


Illinois 


4 


2.4(2.2-2.7) 


1.45(1.42-1 


.50) 


1.08(1.00-1.12) 


.18 (.17-. 20) 


1.1 


ruf! ventre 
















Massachusetts 


4 


3.0(2.5-3.4) 


1.36(1.28-1 


.48) 


.95 (.72-1.10) 


.19 (.17-.21) 


1.3(1.2-1.4) 


Bethany, Conn. 


I 


3.3 


1.20 




.95 


.22 


1.3 


Ithaca, N. Y. 


1 


3.1 


1.36 




1.00 


19 


1.3 


New Jersey 


6 


3.2(2.8-3.5) 


1.34(1.29-1 


.37) 


1 .03 (.94-1.13) 


.18 (.17-.19) 


1.4(1.3-1.5) 


Delaware 


3 


3.1 (3.0-3.3) 


1.33(1.23-1 


.43) 


1.01 (.94-1.10) 


.19 


1.4(1.3-1.4) 


Maryland 


11 


3.2(2.6-3.7) 


1.32(1.25-1 


.35) 


.02 (.89-1.10) 


.18 (1A-.21) 


1.3(1.2-1.4) 


No. Virginia 


12 


3.1 (2.5-3.4) 


1.31 (1.22-1 


.38) 


.97 (.87-1. 12) 


,19(.17-.21) 


1.4(1.2-1.5) 


West Virginia 


4 


3.3(2.9-3.6) 


1.38(1.33-1 


.43) 


.09(1.03-1.21) 


.17 


1.3 


Wake Co., N. C. 


2 


3.1 (2.8-3.3) 


1.22(1.20-1 


.24) 


.85 (.81-90) 


.19 


1.4 


Dare Co., N. C. 


10 


2.4(2.2-2.8) 


1.30(1.23-1 


.35) 


.04 (.96-1.12) 


.17(.16-.18) 


1.2(1.1-1.3) 


Experiment, Ga. 


2 


3.2 


1.20 




.83 (.73-. 93) 


.20(.18-.22) 


1.4 


Gainesville, Fla. 




2.5 


1.13 




.70 


.26 


1.2 


Orange Co., Fla. 




3.0 


1.31 




.04 


.17 


1.4 


Daytona Beach, Fla. 




2.1 


1.28 




.92 


.17 


1.3 


Miami, Fla. 




2.3 


1.18 




.78 


.25 


1.2 


Clay Co., Ala. 




2.9 


1.04 




.73 


.22 


1.5 


Tuscaloosa, Ala. 




2.8 


1.40 




.91 


.17 


1.2 


Harahan, La. 




2.5 


1.27 




.90 


.20 


1.3 


Comanche 
















Santa Cruz Co., Ariz. 


2 


3.1 (2.8-3.4) 


1.21 (1.17-1 


.25) 


.38 (82-. 94) 


.22 


1.4 


Pinal Co., Ariz. 


3 


2.7(2.6-2.8) 


1.14(1.12-1 


.17) 


.60 (.51-. 67) 


.24 (.23-. 25) 


1.7(1.4-2.0) 


Cochise Co. , Ariz. 


1 


2.9 


1.26 




.95 


.22 


1.6 


Ward Co., Texas 


1 


2.9 


1.04 




.52 


.30 


1.4 


pusillum 
















Shreveport, La. 


6 


2.0(1.6-2.5) 


1.24(1.19-1 


.28) 


.90 (.73-1.0) 


19 (.18-. 21) 


1.2(1.2-1.3] 



of flagellum pale, decidedly coarse and semi-erect, erect setae 
numerous, many of them nearly half as long as width of flagel- 
lum. Front rather broad, WF .71 X WH, 1.40 X HE ; ocelli 
relatively small, DAO .17 X WF ; ocelli in a rather broad tri- 
angle, front angle nearly a right angle ; OOL .88 X WOT. Head 
very slightly wider than high; vertex very broadly rounded, 
distance from tops of eyes to vertex crest equal to about .65 X 
HE ; anterior ocellus touching an imaginary line drawn between 
eye tops. Front alutaceous, somewhat shining, punctures shallow 
but rather well defined, separated from one another by about 
their own diameters. Pronotum alutaceous, moderately shining, 
punctures small but rather distinct. Mesoscutum shining, rather 
weakly alutaceous, distinctly punctate ; notauli strong on anterior 
.4 ; scutellar disc strongly shining. Propodeum short, only about 
1.2 X as long as broad, disc with reticulate sculpturing anter- 
iorly, median carina strong ; spiracles elliptical, directed dorsad. 



290 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Mesoscutum wholly somewhat shining:, but the callus especially 
so ; callus convex, well differentiated ; anterior part of meso- 
pleurum with numerous large punctures. Fore wing with dis- 
coidal vein evidenced by only a very faint, unpigmented line 
(Fig. 60). 

Males examined. — DELAWARE : 1, Dover, 4 Aug. 1932 
[USNM]. MARYLAND: 1, Chestertown, 30 July 1901 (E. G. 
Vanatta) [ANSP]. DISTRICT OF COLUMBIA: 1, Washing- 
ton (C. N. Ainslie) [USNM]. VIRGINIA: 1, Vienna, June 
(J. C. Bridwell) [USNM] ; 1, Arlington, 24 Aug. 1946 (R. H. 
Beamer) [KU]. NORTH CAROLINA: 2, Raleigh, 26 July 
1948 (M. W. King) [USNM]. SOUTH CAROLINA: 1, Aiken, 
23 June 1957 (W. R. M. Mason) [CNC]. GEORGIA: 1, Pratts- 
burg, 25 July 1930 (P. W. Oman) [KU] ; 2, Roberta, 6 Oct. 
1945 (P. W. Fattig) [INHS] ; 1, Okefenokee Swamp, 25 July 
1939 (R. H. Beamer) [KU]. FLORIDA: 3 Welaka, May (H. E. 
Evans) [CU] ; 1 Sebring, 31 Aug. 1942 (C. T. Parsons) 
[MCZ] ; 1, Old Town, 11 July 1939 (R. H. Beamer) [KU] ; 1, 
Sanford, 21 June 1933 (C. O. Bare) [KU] ; 2, Winter Park, 
May, July (at light, H. T. Fernald) [FSPB] ; 1, Okeechobee, 3 
April 1953 (J. C. Martin) [CNC] ; 1, Paradise Key, Everglades 
Nat. Pk., 5 April 1952 (G. S. Walley) [CNC]. ALABAMA: 2, 
Cottondale, 18 June 1957 (at light, W. L. Brown) [MCZ] ; 8, 
Tuscaloosa, July, Aug. (at light, B. D. Valentine) [CU, MCZ]. 
TEXAS : 4, St. Austin St. Pk., nr. Sealy, 14-16 June 1956 (at 
light, H. E. Evans and E. G. Matthews) [CU, MCZ] ; 1, Rich- 
mond, Fort Bend Co., 22 June 1917 [MCZ] ; 1, (Belfrage) 
[type, Stockholm Mus.]. KANSAS: 1, Scott Co., 14 Aug. 1951 
(at light, H. E. Evans) [KSU]. ILLINOIS: 2, Carterville, 
Williamson Co., Aug. 1958 (V. Cole) [UCD] ; 1, Urbana, 4 Sept. 
1945 (at light, H. H. Ross) [INHS] ; 1, Golconda, 7 July 1944 
(at light, Sanderson & Leighton) [INHS]. 

Variation. — The 43 specimens examined vary in length from 
2.1 to 4.2 mm., fore wing from 1.8 to 3.0 mm. The abdomen is 
brown in specimens from Maryland, District of Columbia, and 
Arlington, Virginia, while the whole body is very light brown 
in the specimen from Vienna, Virginia ; otherwise the rufous 
abdomen contrasts conspicuously with the head and thorax. The 
antennae show little variation in length and color, but there 
is much variation in the width of the front and in the length of 
the ocello-ocular line as compared to the width of the ocellar 
triangle (Table VII). Some of the latter variation is a result 



EVANS: REVISION OF PSEUDISOBRACHIUM 291 

of differences in the shape of the ocellar triangle, which, however, 
is seldom as compact as in rufivcntre or as broad as in rectangula- 
tum. 

Female (assigned here tentatively). — ALABAMA: Tusca- 
loosa Co., 6 April 1949, under stone on dry cut-over hillside 
(B. D. Valentine) [MCZ]. 

Description of female. — Length 3.2 mm., L1I .58 mm., LT 
1.1 mm. Entire body light castaneous; tips of mandibles rufous; 
antennae light castaneous, fading to dull yellowish-brown apic- 
ally; legs straw-colored. Mandibles very slender, with three 
apical teeth, the basal tooth very small (Fig. 41). Clypeus 
broadly truncate, median carina strong, not reaching apical 
margin. Head 1.25 X as long as wide, sides very weakly, gradu- 
ally convergent to near posterior margin, where they are more 
strongly, arcuately convergent to a broad, straight vertex. 
Eyes small, not larger than one of head punctures, barely 
distinguishable. Head strongly shining, weakly alutaceous ex- 
cept barely so medially and posteriorly, punctures large, poster- 
iorly separated by about or slightly more than their own 
diameters, anteriorly somewhat more crowded (punctures absent 
from median strip) ; under side of head alutaceous, punctures 
rather weak, evenly spaced. Pronotal disc 1.35 X as long as 
its posterior width ; mesonotum 1.4 X as long as wide ; propodeum 
1.4 X as long as wide. Pronotum strongly shining, with Avell 
separated, fairly strong punctures except along midline; meso- 
notum strongly shining, weakly alutaceous laterally ; propodeum 
very strongly polished, with a few weak punctures on extreme 
sides of disc. Mesopleurum weakly alutaceous, weakly punctate. 
Hairs of body and legs short, pale, abundant. 

Other females. --GEORGIA: 2, Peach Co., May, Sept. (in 
soil) [USNM] ; 3, Upson Co., March, July, Aug. (W. F. Turner) 
[USNM]. MISSISSIPPI: 1, Riclgeland, Madison Co., 20 May 
1959 (forest debris, Ross & Stannard) [INHS]. LOUISIANA: 
7, Bossier Par., Feb., May, Sept., Oct. (soil of peach orchard, W. 
F. Turner) [USNM] ; 1, Caddo Par., 13 Sept. 1937 (soil of peach 
orchard, W. F. Turner) [USNM]. TEXAS: 6, Bexar Co., July, 
Sept. (soil of peach orchard, W. F. Turner) [USNM]. ARKAN- 
SAS : 1, St. Francis Co., 13 June 1936 (in soil, W. F. Turner) 
[USNM]. ILLINOIS: 1, Little Grassy Lake, Williamson Co., 10 
Aug. 1958 (oak-hickory woods, in leaf litter in or near nest of 
Solenopsis of group molesta Say, W. L. Brown) [MCZ]. 

Variation in females. — Head length varies from .47 to .60 mm. 
(mean .56 mm.) ; LH/WH varies from 1.22 to 1.35 (mean 1.28) ; 



292 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



thorax length varies from 0.8 to 1.1 mm. (mean about 1.0 mm.). 
Smaller specimens tend also to be slightly more slender (pro- 
notum 1.5 X as long as wide, mesonotum 1.6, propodeum about 
1.5 X as long as wide). While most specimens agree closely in 
color with the one described above, in the Illinois specimen the 
head is medium castaneous, darker than the thorax and abdomen, 
while in the Mississippi specimen the whole body is rich medium 
castaneous. In some of the specimens from Bossier Parish, La., 
the head punctures are somewhat weaker and more widely spaced 
than described above. 

Remarks. — Collecting records indicate that the males are 
nocturnal. The species seems especially characteristic of the Gulf 




Map 5. — ■ Distribution of P. flaviventre, males indicated by solid circles, 
supposed females by hollow circles. Distribution of P. comanche, males indi- 
cated by solid triangles (females unknown). 



Coast states and ranges northward in smaller numbers to Illinois 
and to Delaware. This sex association is highly tentative and is 
based on coincidence of distribution of females (Map 5) as well 
as structure intermediate between ashmeadi and rufiventre (as 
is characteristic of the males). 

Separation of both sexes from the largely sympatric species 
ashmeadi and rufiventre is difficult. The females are about the 



EVANS : REVISION OF PSEUDISOBRACHIUM 293 

same size as those of ashmeadi but have the head more alutaceous 
and closely punctate, somewhat less so than in the slightly larger 
species rufiventre. In the males, the abdomen is usually rufous 
in flaviventre, occasionally rufous in rufiventre, and never rufous 
in ashmeadi. The antennae are only occasionally as short in 
rufiventre and ashmeadi as in flaviventre, and the mesopleurum 
is not of quite the same configuration. In ashmeadi and rufiventre 
the wings tend to have darker veins and setulae and the discoidal 
vein is distinctly pigmented ; however, occasional specimens are 
difficult to place on this character alone. 

35. Pseudisobrachium rufiventre (Ashmead) 

Isobrachium rufiventre Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 38-39. 

[Type: $ , Occoquaii Falls, Va., 5 Aug. 1885, under stone (T. Pergande) 

(USNM no. 2186); allotype: $, on same pin]. 
Pseudisobrachium rufiventre Kieffer, 1908, Genera Insect., 76: 24. — Kieff er, 

1914, Das Tierreich, 41: 480. — Fouts, 1928, Proc. Ent. Soc. Wash., 

30: 122. 
Pseudisobrachium flavicoxis Fouts, 1928, Proc. Ent. Soc. Wash., 30: 122. 

[Type: $, Baldwin, Kansas (USNM no. 62548)]. New synonymy. 
Pseiidisobradhium puncticcps Fouts, 1928, Proc. Ent. Soc. Wash., 30: 

123-124. [Type: $, Glen Echo, Md., 19 Aug. 1916 (at light, R. Fouts) 

(USNM no. 62550)]. New synonymy. 

Description of male allotype. — Length 3.3 mm., LFW 2.6 mm. 
Head and thorax dark reddish-brown, abdomen slightly paler 
and with indistinct banding with light brown, especially basally ; 
mandibles light brown, teeth rufous ; scape brown, flagellum dull 
reddish-brown ; legs wholly light brown ; wings hyaline, setulae 
brown, veins and stigma brown. Mandibles with third tooth 
small, basal tooth broad, as in other species of this complex. 
Apical margin of clypeus subtruncate. Antennae with first 
four segments in a ratio of about 18:5:8:8, segment three about 
1.4 X as long as thick, segment eleven 1.3 X as long as thick; 
flagellar pubscence unusually coarse and suberect, erect setae 
numerous, more erect though not very much longer than the 
pubescence. Front rather narrow, WF .64 X WH, 1.23 X HE ; 
ocelli of moderate size, DAO .21 X WF, in a compact triangle, 
posterior ocelli separated by hardly more than their own diam- 
eter ; OOL and WOT subequal. Eyes weakly bulging, head grad- 
ually contracted behind eyes to a rather broadly rounded vertex ; 
distance from eye tops to vertex crest about .8 X HE (Fig. 5). 
Front alutaceous, moderately shining, obscurely punctate. Pro- 
and mesonota also moderately shining though alutaceous, meso- 
scutum with shallow punctures which, on the sides, are separated 



294 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

by little more than their own diameters ; notauli strong on an- 
terior half of mesoscutum. Propodeum about 1.5 X as long as 
broad, in lateral view twice as long as high ; spiracles elliptical, 
directed dorso-laterad ; disc somewhat shining, weakly alutaceous, 
with some reticulate sculpturing at extreme base. Mesopleurum 
shining, weakly alutaceous, callus convex though not well sep- 
arated or differing notably from mesopleurum below and behind 
it ; anterior part of mesopleurum only weakly punctate. Fore 
wing with discoidal vein arising slightly below junction of basal 
and transverse median veins, pigmented to a distance greater 
than length of basal vein. 

Males examined. — MASSACHUSETTS: 1, Cambridge, Aug. 
1932 (J. Bequaert) [MCZ] ; 2, Holliston, 3-7 Sept. (N. Banks) 
[MCZ] ; 1, Framingham (C. A. Frost) [MCZ]. NEW YORK: 

1, Ithaca, 11 Aug. 1937 (at light, P. P. Babiy) [CU]. CON- 
NECTICUT: 1, Bethany, 16 Oct. 1960 (found dead in light 
shade, H. E. Evans) [MCZ]. NEW JERSEY: 2, Pemberton, 
5 Aug. 1939 (H. K. Townes) [HKT] ; 1, Butler, 1955 (R. Dor- 
land) [CU] ; 1, Ramsey, 29 July 1917 [AMNH] ; 1, Lakehurst, 
9 Aug. 1959 (D. Anderson) [CU] ; 1, Moorestown, 11 Aug. 1939 
(H. & M. Townes) [HKT]. DELAWARE : 1, Dover [USNM] ; 

2, Ship John L. H., Delaware Bay, 22 Aug. 1936 [USNM]. 
MARYLAND: 1, Snow Hill, 5 July 1933 (F. C. Bishop) 
[USNM] ; 2, Plummer's Island, 23 Sept. 1960 (H. E. Evans) 
[MCZ] ; 1, Annapolis, 1933 [USNM] ; 6, College Park, July, 
Aug. 1933 (F. C. Bishop) [USNM] ; 1, Glen Echo, 19 Aug. 1916 
(R. Fouts) [USNM]. WEST VIRGINIA: 2, French Creek, 
Upshur Co., Sept. 1938 (G. E. Wallace) [CM] ; 2, Shaver's Fork, 
Tucker Co., Oct. 1938 (GEW) [CM]. VIRGINIA: 6, Vienna, 
June-Aug. (J. C. Bridwell) [USNM] ; 4, Arlington, Aug., Sept. 
[KVK, MCZ] ; 1, Olney, 16 Aug. 1933 (F. C. Bishop) [USNM] ; 
1, Falls Church, 16 Aug. (N. Banks) [MCZ] ; 1, Occoquan Falls, 
5 Aug. 1885 [USNM]. NORTH CAROLINA: 2, Wake Co., July 
(H. & M. Townes) [HKT] ; 10, Kill Devil Hills, Dare Co., July, 
Sept. (at light, K. V. Krombein) [KVK]. GEORGIA: 2, Ex- 
periment, 11 Aug. 1929 (T. Bissell) [USNM]. FLORIDA: 1, 
Gainesville, 10 July 1957 (at light, H. V. Weems) [FSPB] ; 1, 
Orange Co., 21 Dec. 1929 (J. E. Sadler) [USNM] ; 1, Daytona 
Beach, 26 Julv 1945 (G. T. Riegel) [INIIS] ; 1, Miami, 14 Sept. 
1950 (F. G. Butchers) [CNC]. ALABAMA: 1, Clay Co., (II. H. 
Smith) [USNM] ; 1, Tuscaloosa, 5 July 1949 (B. D. Valentine) 
[MCZ]. LOUISIANA: 1, Harahan, 1 Sept. 1944 (C. L. Reming- 
ton) [INHS]. 



EVANS: REVISION OP PSEUDISOBRACHIUM 295 

Variation in males. — The 60 males examined vary in size from 
2.6 to 5.1 mm., fore wing- from 2.1 to 3.7 mm. The head is black or 
nearly so in several specimens. The color of the abdomen varies 
from dark brown with paler annulations basally to wholly 
rufous; the majority of specimens are intermediate, as in the 
allotype. The antennae and legs are frequently paler than in the 
allotype. There is a tendency for specimens from southern parts 
of the range to exhibit lighter colors, but not all specimens fall 
in with this trend. Variation in ocellar size, width of front, and 
antennal length is recorded in Table VII. In occasional speci- 
mens the front is more strongly punctate than described for the 
allotype. 

Description of female holotype. — Length 3.7 mm., LH .8 mm., 
LT 1.5 mm. Head and thorax bright rufo-castaneous, head 
slightly darker than thorax ; abdomen light orange-brown ; cly- 
peus and antennae light castaneous; legs yellowish-brown, ap- 
proaching amber. Mandibles with three teeth, basal tooth weak, 
as shown in Figure 40. Clypeus broadly truncate, median ridge 
strong. Head 1.18 X as long as wide, sides subparallel to just 
before posterior margin, where they are arcuately convergent to 
a broad, straight vertex. Eyes small, colored like head, barely 
distinguishable. Front somewhat alutaceous, weakly or barely 
so behind, more evidently so antero-laterally, where there is a 
weak tendency for the formation of longitudinal striae ; punc- 
tures elongate, absent from median strip but otherwise separated 
by about or slightly less than their own diameters, though less 
crowded anteriorly than in prolongatum; under side of head 
strongly alutaceous, rather weakly punctate. Pronotal disc rather 
short, 1.3 X as long as its posterior width ; mesonotum 1.5 X 
as long as wide ; propodeum about 1.4 X as long as wide. Pro- 
notum polished, barely alutaceous and not at all so medially, 
punctures strong but absent from median strip. Mesonotum 
weakly alutaceous and punctate, but center of disc smooth and 
shining. Propodeum shining, very weakly alutaceous, sides of 
disc with a number of well-defined punctures extending out onto 
median area more than in prolongatum. Mesopleurum alu- 
taceous, weakly punctate laterally. Body and legs with setae 
golden, mostly rather short. 

Other females. — VIRGINIA : 1, Barcroft, 9 Sept. 1934 (J. C. 
Bridwell) [USNM] ; 1, Alexandria, 24 June 1934 (J. C. Brid- 
well) [USNM] ; 1, Fort Lee, 10 April 1903 [MCZ]. 6 MASSA- 
CHUSETTS: 1, Blue Hills, Canton, 20 July 1956 (in nest of 

s No state is indicated on the label of this specimen ; there is also a Fort Lee 
in northern New Jersey. 



296 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



Formica obscuriventris under rock, W. L. Brown) [MCZj. 
KANSAS: 1, Baldwin [USNM]. 

Variation in females. — The Virginia specimens show only- 
very slight variation in size, color, and other characters. The 
Massachusetts specimen is somewhat smaller, length of body 
3 mm., of head .68 mm., of thorax 1.25 mm. It is somewhat 
lighter in color, the head being castaneous, the thorax and 
abdomen light castaneous; the mandibles, clypeus, and lower 
sides of the head are light yellowish-brown, the legs straw-colored. 
In this specimen the head punctures are slightly more widely 
spaced and for the most part less elongate. The Kansas specimen 
(type of flavicoxis Fouts), although from outside the otherwise 
known range of rufiventre, differs from the type of that species 
in no important way except that it is slightly paler, with the 
coxae bright yellowish as indicated by Fouts. 

Remarks. — The distribution of this species is shown on Map 
6. 




Map 6. — Distribution of P. rufiventre, males indicated by solid circles, 
females by hollow circles. Distribution of P. fouts-i, males indicated by 
solid triangles (females unknown). 



EVANS : REVISION OF PSEUDISOBRACHIUM 297 

36. Pseudisobrachium pusillum new species 

Holotype. — 6, Shreveport, La., 22 June 1949 (light trap, 
J. H. Robinson) [INHS]. 

Description. — Length 2.0 mm., LPW 1.8 mm. Head piceous, 
thorax dark castaneous, abdomen medium brown, paler on sides 
of basal segments; mandibles light brown basally, apical half 
straw-colored, teeth rufous ; antennae brown, flagellum somewhat 
paler beneath ; legs brown, tarsi and apices of tibiae paler ; wings 
hyaline, setulae dark, veins and stigma brown. Mandibles with 
four teeth, basal two teeth both strong, basal tooth somewhat 
thicker than third tooth. Clypeus truncate apically. Antennae 
with first four segments in a ratio of about 10 A :5 :5, segments 
three and eleven each about 1.2 X as long as thick; flagellar 
pubescence unusually coarse, suberect, erect setae also numerous. 
Front of moderate breadth, WF .61 X WH, 1.25 X HE ; front 
angle of ocellar triangle less than a right angle, ocelli only 
slightly enlarged, DAO .20 X WF ; OOL .95 X WOT. Distance 
from eye tops to vertex crest .8 X HE. Front alutaceous, mod- 
erately shining, obscurely punctate. Pro- and mesonota rather 
strongly alutaceous, including even scutellar disc, obscurely 
punctate ; notauli impressed on anterior .2 of mesoscutum. Pro- 
podeum about 1.7 X as long as wide, disc weakly alutaceous, 
depressed and carinate medially. Mesopleurum wholly aluta- 
ceous, obscurely punctate, callus small but convex. Fore wing 
with discoidal vein present, weakly pigmented. 

Paratopes. — LOUISIANA: 3 S $ , same data as type [INHS, 
MCZ], 1 $ , same data but 6 Aug. 1948 [USNM], 1 $ , same data 
but 28 July 1949 [CNC]. 

Variation. — The five paratypes show little variation in size 
and in standard measurements (Table VII). They do, however, 
exhibit considerable variation in the degree to which the vertex 
is extended above the eye tops ; in one specimen the distance from 
the eye tops to the vertex measures only about .6 X HE, in two 
others .7, in another .8 (like the type), in another .9. 

Female (assigned here tentatively). — ARKANSAS : Pike Co., 
12 May 1948 (peach orchard, Turner) [USNM]. 

Description. — Length 1.8 mm., LH .40 mm., LT .73 mm. 
Head light castaneous, thorax and abdomen light yellowish- 
brown ; antennae light castaneous, fading to dull straw-yellow 
apically ; legs straw-colored. Mandibles with three teeth, apical 
two teeth large and somewhat splayed out, basal tooth small. 
Clypeus broadly subtruncate apically, its median carina very 



298 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

sharp, reaching apical margin. Head 1.45 X as long as wide, 
sides weakly bulged midway, convergent behind to a broad, 
straight vertex. Eyes not distinguishable. Front shining, non- 
alutaceous, punctures small, shallow, separated for the most part 
by much more than their own diameters ; under side of head 
shining, non-alutaceous, weakly punctate. Pronotal disc 1.5 X 
as long as its posterior width ; mesonotum 1.6 X as long as wide ; 
propodeum about 1.5 X as long as wide. Pro- and mesonota 
strongly shining, weakly punctate except medially, in part ob- 
scurely alutaceous. Propodeal disc very strongly shining, non- 
alutaceous, impunctate. Mesopleurum alutaceous, obscurely punc- 
tate. Hairs of body and legs short and pale. 

Other females. — 1, same data as type [USNM]. 

Variation. — The second specimen measures 2.0 mm. long, head 
.42 mm., thorax .78 mm. The sides of the head are slightly more 
bulging, such that the head is only 1.37 X as long as wide. 

37. Pseudisobrachium comanche new species 

Holotype. — $ , Pefia Blanca, Santa Cruz Co., Arizona, 3950 
feet elevation, 24 Aug. 1959 (at light, H. E. Evans) [MCZ, No. 
30287]. 

Description. — Length 3.2 mm., LFW 2.8 mm. Head and 
thorax piceous, abdomen dark brown, paler on sides of basal seg- 
ments ; apical two-thirds of mandibles yellowish-brown, teeth ruf- 
ous ; antennae dull reddish-brown ; front coxae infuscated, legs 
otherwise bright reddish-brown ; wings hyaline, setulae brownish, 
veins and stigma brown. Mandibles with third tooth very small, 
fourth tooth broad ; clypeus subtruncate apically. Antennae elon- 
gate, first four segments in a ratio of about 17:5:7:7, segment 
three 1.7 X as long as thick, segment eleven 1.4 X as long as 
thick ; flagellar pubescence coarse, suberect, erect setae numerous, 
on basal segments some of them more than half as long as width 
of flagellum. Front narrow, WF .58 X WH, 1.17 X HE ; ocelli 
in a compact triangle, front angle less than a right angle, slightly 
enlarged, DAO .22 X WF ; OOL .82 X WOT. Eyes very prom- 
inent laterally, head contracted behind them to a rather narrowly 
rounded vertex ; distance from eye tops to vertex crest about .6 X 
HE. Front alutaceous, moderately shining, punctures shallow 
and inconspicuous. Pronotum and mesoscutum alutaceous, some- 
what shining, the latter somewhat more evidently punctate than 
the former ; notauli distinct on anterior .3 of mesoscutum ; scutel- 
lar disc shining. Propodeum about 1.6 X as long as broad, in 



EVANS : REVISION OF PSEUDISOBRACHIUM 299 

lateral view 2.6 X as long as high ; disc weakly sculptured, some- 
what polished ; spiracles elliptical, directed dorso-laterad. Meso- 
pleurum wholly alutaceous, with some large, shallow punctures 
anteriorly. Fore wing with discoidal vein lightly pigmented to 
about length of basal vein (Fig. 59). 

Paratypes. — ARIZONA : 1 6 , Madera Canyon, Santa Rita 
Mts., 4880 feet, 24 Aug. 1959 (J. G. Franclemont) [CU] ; 3 
£ S , 4 mi. W. Superior, Pinal Co., May, July (H. K. Gloyd) 
[INHS, USNM, MCZ] ; 1 £ , SW Research Station, 5 mi. W. 
Portal, Cochise Co., 5400 feet, 11 Sept. 1959 (H. E. Evans) 
[MCZ]. TEXAS: 1 6, 1 mi. E. Barstow, Ward Co., 10 July 
1956 (E. G. Matthews) [CU]. 

Variation. — The six paratypes range in size from 3.0 to 
3.9 mm., fore wing from 2.6 to 3.4 mm. There is considerable 
variation in ocellar size and antennal length (Table VII) ; the 
specimens from the Santa Ritas and the Chiricahuas not only 
have relatively small ocelli but also have the head longer behind 
the eyes (distance from eye tops to vertex crest about .8 X HE). 
Otherwise the series is of rather uniform appearance. 

Remarks. — The distribution of this species is shown on Map 5. 

38. Pseudisobrachium apache new species 

Holotype. — • <$ , Peiia Blanca, Santa Cruz Co., Arizona, 3950 
feet elevation, 24 Aug. 1959 (at light, H. E. Evans) [MCZ, No. 
30288]. 

Description. — Length 4.4 mm. ; LFW 3.7 mm. Head black, 
thorax very dark brown, abdomen medium brown with indistinct 
banding with light brown on posterior parts of segments ; apical 
half of mandibles light brown, teeth rufous ; scape dark brown, 
nagellum medium brown ; front coxae dark brown, legs other- 
wise straw-colored ; wings hyaline, stigma brown, veins light 
brown. Mandibles with four teeth, third tooth smaller than 
broadly rounded basal tooth. Clypeus with apical margin slightly 
convex (Fig. 46), median carina high, arched in profile. Anten- 
nae with first four segments in a ratio of about 25 :7 :12 :11, seg- 
ment three nearly twice as long as maximum width, segment 
eleven 1.7 X as long as wide ; pubescence of flagellum light, golden 
brown, suberect, moderately coarse, erect setae numerous but 
mostly rather short, WF .60 X WH, 1.10 X HE ; ocelli rather 
large, DAO .25 X WF ; posterior ocelli rather close together, 
front angle of ocellar triangle less than a right angle ; OOL 
.57 X WOT. Eyes strongly bulging, measuring 1.25 X as high 
as wide ; distance from eye tops to vertex crest equal to about 
half eye height. Front alutaceous, moderately shining, punctures 



300 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

small, shallow, relatively inconspicuous. Pro- and mesonota 
strongly polished, hardly at all alutaceous, punctures small but 
(especially on mesoscutum) sharply denned and rather evenly 
spaced; notauli strong on anterior half of mesoscutum. Pro- 
podeum 1.7 X as long as wide; disc shining though with weak 
microscopic sculpturing; spiracles narrowly elliptical, directed 
dorsad. Mesopleurum strongly shining, with some sculpturing 
anteriorly and posteriorly; callus elongate, very weakly alu- 
taceous. Discoidal vein of fore wing a scarcely pigmented streak 
which arises a short distance down on transverse median vein. 

Paratypes. — ARIZONA : 1 $ , Patagonia, Santa Cruz Co., 14 
Oct. 1927 (J. A. Kusche) [CAS] ;3 N, Globe, Gila Co., July 
1949 (at light) [USNM] ; 7 S $ , 4 mi. W. Superior, Pinal Co., 
May-Aug. 1946-49 (at light, H. K. Gloyd, B. W. Benson) [INHS, 
MCZ, CU]. 

Variation. — The 12 available specimens range in size from 
3.5 to 5.0 mm., fore wing from 3.0 to 3.9 mm. In one specimen 
from Globe the front is somewhat more shining than in the type, 
and in another specimen from that locality and one from Su- 
perior the pro- and mesonota are very weakly alutaceous; how- 
ever, in every case there is marked contrast between the front 
and the thoracic dorsum in this regard. The amount of variation 
in antennal length and head measurements is unusually small 
(Table VIII). 

39. Pseudisobrachium foutsi new species 

Isobrachium montanuvi Ashmead, 1893, Bull. U. S. Nat. Mus., 45: 39. 
[$ allotype, "Arizona" (USNM); $ holotype = P. prolongatum 
Prov.]. 

Holotype. — <5 , Port Isabel, Cameron Co., Texas, 23-27 June 
1956 (at light, H. E. Evans & E. G. Matthews) [MCZ, No. 
30289]. 

Description. — Length 4.6 mm., LFW 2.8 mm. Head dark 
reddish-brown, thorax castaneous, abdomen yellowish-brown ex- 
cept first tergite infuscated basally, all tergites somewhat paler 
apically ; mandibles yellowish-brown, darker basally and apically ; 
antennae light castaneous; legs wholly bright yellowish-brown; 
wings hyaline, with pale setulae, stigma light brown, veins pale 
amber. Mandibles with third tooth smaller than broad basal 
tooth; clypeus narrowly truncate apically (Fig. 29). Antennae 
with first four segments in a ratio of about 4 :1 :2 :2, segments 
three and eleven each about 1.5 X as long as thick; flagellar pu- 
bescence pale, coarse, suberect, erect setae numerous but not 



EVANS : REVISION OF PSETJDISOBRACHIUM 301 

much longer than pubescence. Front narrow, WP .62 X WH, 
1.10 X HE; ocelli enlarged, in a rather compact triangle the 
front angle of which is distinctly less than a right angle ; DAO 
.26 X WF; OOL .64 X WOT. Eyes only slightly bulging, head 
gradually narrowed behind eyes, distance from eye tops to 
vertex crest about .65 X HE. Front alutaceous, weakly shining, 
punctures shallow and inconspicuous. Pro- and mesonota also 
alutaceous and rather weakly shining, obscurely punctate except 
somewhat more evidently so on sides of mesoscutum; notauli 
weakly impressed on anterior .2 of mesoscutum; scutellar disc 
shining but alutaceous and weakly punctate. Propodeum 1.6 X 
as long as broad, disc shining, weakly alutaceous but with some 
reticulate sculpturing anteriorly ; spiracles elongate-elliptical, di- 
rected dorso-laterad. Mesopleurum wholly alutaceous, weakly 
punctate anteriorly, callus weakly differentiated. Fore wing 
with discoidal vein indicated by a faint, unpigmented streak which 
is interstitial with median vein (Fig. 61). 

Paratypes. — TEXAS : 23 8 8 , Port Isabel, Cameron Co., all 
same data as type except 2 8 8 20-23 June 1948 (at light, H. E. 
Evans) [MCZ, CU, USNM] ; 3 8 8 , Lolita, Jackson Co., 6 July 
1916 (at light, J. D. Mitchell) [USNM] ; 4 8 8 , Gillett, Karnes 
Co., 25 June 1917 (J. C. Bradley) [CU] ; 1 8 , Helotes, Bexar 
Co., 1 July 1917 (J. C. Bradley) [CU] ; 16 8 8 , Kerrville, Kerr 
Co., July 1952-53 (light trap, L. J. Bottimer) [USNM] ; 24 8 8 , 
Brazos River, Richmond, Ft. Bend Co., 22 June 1917 (at light, 
J. C. Bradley) [CU] ; 1 8 , Ft. Hood, Bell Co., 21 July 1955 
(E. G. Matthews) [CU]. 

Other material examined. — The following specimens are as- 
signed here tentatively but are not to be considered paratypes : 
TEXAS : 4 8 8 , Brownsville, Cameron Co., June, Sept., Oct. 
[USNM, CAS, KU] ; 1 8 , Mission, Hidalgo Co., 27 Sept, 1951 
(Cartwright & Gurney) [USNM] ; 1 8 , Knippa, Uvalde Co., 3 
July 1910 (F. C. Pratt) [USNM] ; 1 $ , Juno, Val Verde Co., 
3 July 1917 (J. C. Bradley) [CU] ; 1 8 , Lozier Canyon, Terrell 
Co., 8 July 1948 (at light in desert, Werner & Nutting) 
[USNM] ; 1 $ , Limpia Canyon, Davis Mts., 7 July 1917 (J. C. 
Bradley) [CU] ; 1 8, "Tex." [ANSP]. NEW MEXICO: 
2 8 8, Roswell, 17 Aug. 1951 (at light, H. E. Evans) [MCZ] ; 
2 $ 8 , Lordsburg, 13-17 July 1917 (J. C. Bradley, J. Bequaert) 
[CU,MCZ];5 8 8 , Deming, 12-17 July (W. M. Wheeler) [MCZ]. 
ARIZONA .- 1 8 , Portal, Cochise Co., 5 Sept. 1959 (at light, H. 
E. Evans) [CU] :H, Paradise, Cochise Co.. 17 Sept.. 1927 (J. 
A. Kusche) [CAS] ; 4 8 8 , Texas Canyon, Cochise Co., Sept., 



302 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Oct. (J. A. Kusche) [CAS] ; 6 $ £ , Willcox, 3 Sept. 1959 (at 
light, H. E. Evans) [CU, MCZ] ; 1 $ , Huaehuca mts., 14 Sept, 
1933 (R. II. Crandall) [UA] ; 2 $ $ , Safford, 23 June 1954 
(light trap, F. G. Werner) [UA] ; 10 & $ , 4 mi. W. Superior, 
Pinal Co., May, July (at light, H. K. Gloyd) [INHS] ; 2 $ $ , 
Canelo, Santa Cruz Co., 10 July 1957 (G. D. Butler) [UA] ; 
1 $ , Patagonia, Santa Cruz Co., 14 Oct. 1927 (J. A. Kusche) 
[CAS] ; 2 $ $ , Tucson, Aug. (R. H. Crandall) [UA] ; 1 $ , 
Tuba City, Coconino Co., 27 July 1954 (at light, H. E. & M. A. 
Evans) [CU]. CALIFORNIA: 32 $ $ , Blythe, Riverside Co., 
6-20 Aug. 1947 (light trap, J. W. MacSwain) [CIS, MCZ] ; 1 $ 
Thermal, Riverside Co., 17-18 Aug. 1927 [CU] ; 6 $ $ , Riverside, 
24 Sept. 1935 (H. H. Keifer) [CDAS] ; 1 $ , El Centro, Imperial 
Co., 11 Sept. 1959 (light trap, C. R. Waegner) [CDAS] ; 1 $ , 
Chula Vista, San Diego Co., 26 Sept. 1935 (H. H. Keifer) 
[CDAS]; 1 $, Lancaster, Los Angeles Co., 26 Aug. 1958 (in 
alfalfa field, E. I. Schlinger) [USNM] ; 1 $ , 8 mi. NW Winters, 
Yolo Co., 22 July 1959 (light trap, J. Fowler) [UCD]. COA- 
HUILA :H, Tanque de Malone, La Bahia, 20 June 1938 (R. H. 
Baker) [USNM]. 

Variation. — The amount of variation in this species is be- 
wildering, and I concede the possibility that I may be confusing 
several species under one name. Most specimens from eastern 
Texas agree reasonably well in size, color, and standard measure- 
ments, and I have little doubt that they are conspecific with the 
Port Isabel type ; I have therefore designated these specimens as 
paratypes. Specimens from southern and western Texas, Coa- 
huila, New Mexico, Arizona, and California may be in part or 
wholly conspecific with the type series; I have identified these 
specimens as font si but prefer not to designate them as paratypes. 

The smallest specimen in the entire series is from Lancaster, 
California (length 1.6 mm., LFW 1.6 mm.) while the largest is 
from Texas Canyon, Arizona (length 4.7 mm., LFW 3.6 mm.). 
Specimens from western parts of the range tend to be darker in 
color; for example, the series from Willcox, Arizona, and most 
California specimens have the entire body dark brown and the 
antennae and legs brown. On the other hand, the specimens 
from Blythe, California, have the abdomen rufous and the legs 
wholly straw-colored. The two specimens from Canelo, Arizona, 
are similarly colored, but in these the front is unusually strongly 
punctate. In two of the four Brownsville, Texas, specimens the 
head is rather long behind the eyes (distance from eye tops to 
vertex crest .7-. 9 X HE), and this is true of many Arizona and 
most California specimens, though there is much variation in 



EVANS : REVISION OF PSEUDISOBRACHIUM 



303 



this regard in most series from one locality. In some specimens 
(mainly of somewhat peripheral distribution, see Table VIII) 
the ocelli are enlarged only slightly. The broadest front occurs 
in the series from Riverside, California, although this is ap- 
proached by specimens from other California, Arizona, and New 
Mexico localities. The mandibles of some Arizona specimens have 
the basal tooth scarcely any wider than the third tooth (Fig. 
30). 

Remarks. — This nocturnal, chiefly deserticolous species is 
named for Robert M. Fouts, of Laredo, Texas, the only North 
American worker to have directed attention to these wasps up 
to this time. The female has yet to be discovered. The distribu- 
tion is shown on Map 6. 









TABLE VIII 








Species and locality 


No. 


LFW 


WF/HE 




00L/W0T 


DA0/WF 


Ant. 11 L/W 


foutil 
















Richmond, Tex. 


24 


2.6(2.1-3.0) 


1.3(1.05-1. 


21) 


.62 (.53-. 69) 


.28 (.25-. 31) 


1.2(1.1-1.3) 


Ft. Hood, Tex. 


1 


2.6 


1.06 




.50 


.30 


1.4 


Helotes, Tex. 


1 


2.9 


1.00 




.40 


.31 


. 


Gillett, Tex. 


4 


2.2(2.0-2.7) 


1.17(1.06-1 


.33) 


.64 (.47-. 75) 


.24 (.18-. 29) 


1.1 (1.0-1.3) 


Lolita, Tex. 


3 


2.6(2.3-2.9) 


1.22(1.14-1 


.35) 


.57 (.54-. 62) 


.27 (.25-. 28) 


1.3(1.2-1.4) 


Mission, Tex. 


] 


2.8 


1.34 




.88 


.18 


1.5 


Brownsville, Tex. 


4 


2.7(2.4-3.0) 


1.13( .93-1 


.27) 


.61 (.40-. 76) 


.28 (.23-. 39) 


1.3(1.0-1.5) 


Port Isabel, Tex. 


24 


2.4(1.9-2.8) 


1.20(1.13-1 


.30) 


.70 (.62-. 79) 


.24 (.23-. 26) 


1.2(1.0-1.4) 


Kerrvllle, Tex. 


16 


2.4(1.8-2.9) 


1.05 ( .95-1 


.16) 


.56(.39-.91) 


.29 (.20-. 33) 


1.3(1.1-1.5) 


Knlppa, Tex. 


1 


1.9 


1.22 




.70 


.20 


1.1 


Juno, Tex. 


1 


1.8 


1.22 




.80 


.20 


1.0 


Terrell Co., Tex. 


1 


1.9 


1.11 




.56 


.27 


1.3 


Jeff Davis Co., Tex. 


1 


2.5 


1.15 




.55 


.25 


1.3 


Roswell, N. Mex. 


2 


3.0(2.9-3.1) 


1.15 




.51 (.50-. 52) 


.26 (.25-. 27) 


1.2 


Lordsburg, N. Mex. 


2 


2.9 


1.28(1.18-1 


.38) 


.59 (.42-. 76) 


.26 (.22-. 30) 


1.3 


Demlng, N. Mex. 


5 


3.2(3.0-3.4) 


1.48(1.33-1 


.59) 


.71 (.64-. 77) 


.21 (.18-. 24) 


1.3(1.2-1.4) 


Cochise Co., Ariz. 


13 


2.9(1.9-3.6) 


1.28(1.12-1 


.48) 


.68 (.52-. 87) 


.23 (.17-. 27) 


1.3(1.1-1.5) 


Safford, Ariz. 


2 


2.8 


1.37(1.34-1 


.40) 


.73 (.69-. 76) 


.23 (.22-. 24) 


1.1 


Superior, Ariz. 


8 


2.4(1.9-2.8) 


1.26(1.17-1 


.36) 


.85 (.80-. 91) 


.22 (.21-. 24) 


1.3(1.2-1.4) 


St. Cruz Co., Ariz. 


3 


3.3(3.2-3.4) 


1.27(1.20-1 


.34) 


.77 (.70-. 85) 


.21 (.20-. 23) 


1.3(1.2-1.4) 


Tucson, Ariz. 


2 


2.9(2.8-3.0) 


1.16(1.07-1 


.25) 


.67 (.62-. 72) 


.24(.22-.26) 


1.2 


Coconino Co., Ariz. 


1 


2.5 


1.32 




.78 


.23 


1.1 


Blythe, Cal. 


32 


3.0(2.5-3.5) 


1.32(1.10-1 


• 54) 


.81 (.64-1.00) 


.20 (.17-. 23) 


1.3(1.1-1.4) 


El Centro, Cal. 


1 


2.5 


1.25 




.83 


.21 


- 


Riverside, Cal. 


6 


3.2(3.0-3.4) 


1.50(1.44-' 


.60) 


.90 (.78-1.06) 


.17(.16-.18) 


1.3 


Thermal, Cal. 


1 


2.8 


1.13 




.76 


.22 


1.1 


San Diego Co. , Cal. 


1 


2.6 


1.14 




.62 


.23 


1.3 


Los Angeles Co., Cal. 


1 


1.6 


1.00 




.52 


.29 


1.3 


Yolo Co., Cal. 


1 


2.7 


1.30 




.80 


.22 


1.4 


La Babla, Coahuila 


1 


2.3 


1.27 




.71 


.20 


1.2 


emarglnatum 
















Kerrville, Texas 


3 


2.6(2.5-2.7) 


1.30(1.26- 


1.33) 


.86 (.82-. 90) 


.I7(.16-.18) 


1.3(1.2-1.5) 


apache 
















Pena Blanca, Ariz. 


1 


3.7 


1.10 




.57 


.25 


1.7 


Patagonia, Ariz. 


1 


3.5 


1.16 




.62 


.25 


1.5 


Globe, Ariz. 


3 


3.4(3.0-3.7) 


1.16(1.10- 


1.26) 


.62 (.56-. 73) 


.26 (.24-. 28) 


1.6(1.4-1.8) 


Superior, Ariz. 


7 


3.5(3.1-3.9) 


1.16(1.13- 


1.19) 


.63 (.55-. 68) 


.25 (.24-. 26) 


1.6(1.5-1.7) 



40. PSEUDISOBRACHIUM EMARGINATUM new species 

Holotype.— S , Kerrville, Texas, 10-14 July 1953 (light trap, 
L. J. Bottimer) [USNM, no. 65155]. 

Description. — Length 3.4 mm., LFW 2.6 mm. Head and 
thorax black, abdomen dark brown, somewhat paler on sides of 
basal segments ; apical half of mandibles yellowish, teeth rufous ; 



304 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

scape dark brown, flagellum medium brown ; front coxae dark 
brown, remaining coxae and all femora medium brown, rest of 
legs light brown; wings hyaline, setulae light brown, veins 
and stigma very light brown. Mandibles with third tooth small, 
fourth tooth broad. Clypeus with median carina low, weakly 
arched; apical margin distinctly arcuately concave (Fig. 49). 
First four antennal segments in a ratio of about 17:5:8:7, seg- 
ment three and segment eleven each about 1.5 X as long as thick ; 
flagellar pubescence pale, very coarse and suberect, erect setae 
numerous and rather long. Front of moderate breadth, WF 
.64 X WH, 1.33 X HE ; ocelli only very slightly enlarged, DAO 
.17 X WF ; OOL .90 X WOT ; ocelli well separated, front angle 
of ocellar triangle less than a right angle. Eyes rather bulging, 
head narrowed behind the eyes, vertex rather evenly rounded off 
a distance above the eyes equal to about three fourths eye height. 
Front shining, weakly alutaceous, punctures close though shallow 
and rather inconspicuous. Pronotum and mesoscutum more 
strongly alutaceous and less shining, barely punctate; notauli 
weakly impressed on anterior .3 of mesoscutum; scutellar disc 
shining. Propodeum about 1.5 X as long as wide, disc wholly 
alutaceous though somewhat shining; spiracles small, elliptical, 
directed dorso-laterad. Mesopleurum wholly alutaceous, without 
strong punctures, callus large, moderately convex. Fore wing 
with discoidal vein represented only by an unpigmented streak. 

Paratypes. — 2 S $ , same data as type [USNM, MCZ]. 

Variation. — The two paratypes are similar to the type in size 
and coloration. However, in both of them the front is more 
strongly alutaceous and less shining than in the type, the anten- 
nae are shorter and somewhat lighter brown, and the legs are 
also lighter brown. 

41. PSEUDISOBRACHIUM RECTANGULATUM new species 

Holotype. — $ , Red River, Wilbarger Co., Texas, 5 July 1956 
(at light, H. E. Evans and E. G. Matthews) [MCZ, No. 30291]. 

Description. — Length 2.8 mm.; LFW 2.2 mm. Head dark 
reddish-brown, thorax medium brown, abdomen light brown 
basally, darker beyond; mandibles, clypeus, and antennae light 
reddish-brown; legs wholly light brown; wings hyaline, setulae 
pale, veins and stigma light brown. Mandibles as in preceding 
species. Clypeus narrowly truncate apically, its truncate apical 
margin very slightly longer than third antennal segment ; median 
carina weakly arched in profile. Antennae with first four seg- 
ments in a ratio of about 17:4:7:7, segment three about 1.4 X 



EVANS : REVISION OP PSEUDISOBRACHIUM 305 

as long as wide, segment eleven 1.2 X as long as wide ; flagellar 
pubescence pale, short, subappressed, erect setae short and sparse. 
WP .65 X WH, 1.31 X HE ; ocelli slightly enlarged, DAO .20 X 
WF ; ocelli in a broad triangle, front angle a right angle, OOL 
.75 X WOT. Vertex broadly rounded a distance above eye tops 
equal to about two-thirds X HE. Front alutaceous, moderately 
shining, with small, rather shallow punctures which are sep- 
arated from one another by somewhat more than their own diam- 
eters. Pronotum and mesoscutum alutaceous, moderately shin- 
ing, obscurely punctate ; notauli distinct for anterior .3 of 
mesoscutum ; scutellar disc weakly alutaceous, somewhat shining. 
Propodeum about 1.5 X as long as broad, disc alutaceous, median 
carina strong ; spiracles small, subcircular, directed dorso-laterad. 
Mesopleurum wholly altuaceous, callus not especially well defined. 
Fore wing with discoidal vein indicated only by an unpigmented 
streak which is interstitial with medial vein. 

Paratypes. — TEXAS: 2 $ $, same data as type [CU, 
USNM] ; 1 $ , Victoria, 19 Aug. 1912 (J. D. Mitchell)' [USNM]. 
NEBRASKA: 1 $ , Sidney, 2 Aug. 1936 (H. H. Ross) [INHS]. 
KANSAS: 1 6 , Grant Co., 18 Aug. 1952 (at light, H. E. Evans) 
[MCZ]. 

The following additional specimen is assigned here tentatively 
and is not to be regarded a paratype : NUEVO LEON : 1 $ , 
Vallecillo, 2-5 June 1951 (at light," H. E. Evans) [MCZ]. 

Variation. — The two topotypic paratypes resemble the type 
closely, showing only slight differences in ocellar size. The 
Kansas and Nebraska specimens are somewhat darker in color 
and the front is broader; the Nebraska specimen is a little more 
noticeably punctate on the head and thorax than the type, 
the Kansas specimen less so. The specimen from Victoria, Texas, 
has shorter and slightly paler antennae but is otherwise close 
to the type. In the Nuevo Leon specimen the front is narrow 
and somewhat more shining and more distinctly punctate than in 
the. rest of the series; the ocelli are rather large and close to- 
gether ; the distance from the eye tops to the vertex crest is equal 
to only slightly more than half the eye height ; the mesopleural 
callus is large and somewhat convex and shining. Altogether 
this specimen presents a somewhat different appearance and may 
well represent a distinct species. 

42. PSEUDISOBRACHIUM FLAVTNERVIS Fonts 

Psewlisobrachium flavinervis Fouts, 1928, Proc. Ent. Soc. Wash., 30: 123. 
[Type: $, Lone Star, Calif., 21 Sept. 1927 (taken from soil) (USNM, 
no. 41217)]. 



306 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

Description of type. — Length 4.0 mm. ; LFW 3.2 mm. Head 
piceous, thorax dark reddish-brown, abdomen brown, obscurely 
banded with light brown ; mandibles light brown, teeth rufous ; 
scape light brown, flagellum bright, pale castaneous ; front coxae 
brownish, legs otherwise straw-colored; wings very pale, setulae 
pale, stigma very light brown, veins nearly colorless. Mandibles 
with third tooth small, basal tooth broad, arching into inner 
mandibular margin (Pig. 31). Clypeus narrowly truncate 
apically, median carina barely arched in profile. Antennae with 
first four segments in a ratio of about 20:6:8:7, segment three 
and segment eleven each about 1.3 X as long as thick ; pubescence 
of flagellum very pale, short, subappressed, erect setae very 
sparse, short, and inconspicuous. Front narrow, WP .63 X WH, 
1.10 X HE ; ocelli large ; DAO .28 X WF ; OOL .44 X WOT, 
latter very broad, front angle greater than a right angle. Dis- 
tance from eye tops to vertex crest equal to only about half eye 
height; eyes rather long, not strongly bulging, rather weakly 
hairy (Fig. 6). Front alutaceous, moderately shining, with small 
but well-defined punctures which are separated from one another 
by slightly more than their own diameters. Pro- and mesonota 
alutaceous, somewhat shining, only rather obscurely punctate ; 
notauli impressed on anterior .3 of mesoscutum ; disc of scutellum 
shining, with a few punctures. Propodeum 1.45 X as long as 
broad ; disc alutaceous, with a strong median carina ; spiracles 
small, elongate, directed somewhat dorsad. Mesopleurum alu- 
taceous, with some small punctures antero-ventrally ; callus not 
especially strongly defined. Discoidal vein of fore wing repre- 
sented by a faint, unpigmented streak which arises well below 
junction of basal and transverse median veins (Fig. 62). 

Specimens examined. — CALIFORNIA : 1 $ , Lone Star, Fres- 
no Co., 21 Sept. 1927 [USNM] ;2 n, Mojave, Kern Co., 25 
July 1947 (R. H. Beamer) [KIT] ; 2 $ $ , Llano, Los Angeles 
Co., 12 July 1956 (E. I. Schlinger) [UCD] ; 2 $ $ , Nr. Hinc- 
ley, San Bernardino Co., 1 Aug. 1927 (Rehn, Pate, Rehn) 
[ANSP] ; 1 S , Victorville, 15 Aug. 1927 (J. C. Bradley) [CU] ; 
2 $ $ , Thermal, Riverside Co., 17 Aug. 1927 (J. C. Bradley) 
[CU] ; 12 $ $ , Blythe, Riverside Co., 6 July, 20 Aug. 1947 
(light trap, Barr & MacSwain) [CIS] ; 10 $ $ , Holtville, Im- 
perial Co., Sept., Oct, 1959 (light trap, C. R. Waegner) [CDAS] ; 
6 $ $ , El Centro, 12 Oct. 1959 (light trap, C. R, Waegner) 
[CDAS] ; 6 $ S , Imperial, 19-22 Oct. 1959 (light trap, C. R. 
Waegner) [CDAS] ; 2 $ $ , Calexico, 22 Oct, 1959 (C. R. Waeg- 
ner) ; 1 $ , Winterhaven, 3 Sept, 1959 (light trap, C. R. Waeg- 
ner) [CDAS] ; 15 $ $ , Imperial Co., June 1912 (J. C. Bridwell) 



EVANS : REVISION OF PSEUDISOBRACHIUM 307 

[TJSNM]. BAJA CALIFORNIA : 2 $ $ , San Felipe, June 1939 
(Michelbacher & Ross) [CAS] ; 1 $ , Coyote Cove, Conception 
Bay, 29 June 1938 (Michelbacher & Ross) [CAS]. SONORA : 
4 $ $ , Hermosillo, 25 June 1959 (at light, H. E. Evans) [CU, 
MCZ]. ARIZONA: 1 $, Quartzite, Yuma Co., 20 Aug. 1927 
(J. C. Bradley) [CU] ; 8 $ £ , Wellton, Yuma Co., Aug., Sept. 
[CU, MCZ] ; Palomas, Yuma Co., 8 Aug. 1917 (J. C. Bradley) 
[CU] ; 4 $ $ , Gila Bend, 17 Sept. 1938 (R. H. Crandall) [UA] ; 

4 6 <$ , Phoenix, Aug., Sept. [CU, UA] ; 2 $ £ , Higley, 29 July 
(at light, J. Bequaert) [MCZ] ; 7 $ $ , Roosevelt Lake, 11 Aug. 
1949 [USNM] ; 135 S S , 4 mi. W. Superior, June-Oct. (light 
trap, Gloyd, Benson) [INHS, USNM, MCZ] ; 1 $ , Florence 
[USNM] ; 4 3 $ , Maricopa, 17 Oct. 1927 (J. A. Kusche) [CAS] ; 

5 S $ , Tucson, June-Aug. [CAS, UA] ; 5 S S , Sabino Canyon, 
Santa Catalina Mts., July, Aug. [KVK, UA] ; 1 $ , Organ Pipe 
Nat, Mon., 8 May 1955 (light trap, J. Eden) [UA] ; 1 S , Babo- 
quivari Canyon, W. side Baboquivari Mts., 25 July 1952 [CAS]. 
NEVADA: 1 S, Las Vegas, 30 June 1940 (A. L. Melander) 
[MCZ]. TEXAS -.1$, Valentine, 12 July 1938 (R. H. Beamer) 
[KIT]. 

Variation. — The 247 males examined show a size range from 
2.0 to 5.2 mm., with most specimens between 3.5 and 4.5 mm. ; 
fore wing size ranges from 1.6 to 3.7 mm. Some specimens are 
lighter in color than the type, and in some specimens from 
southern California and from western Arizona the abdomen is 
distinctly paler than the head and thorax. In many specimens 
from these areas the clypeus is light yellowishJorown, contrasting 
to the much darker front ; this is true, for example, of the series 
from Gila Bend and from Roosevelt Lake, Arizona. In the 
majority of specimens the front is rather strongly shining and 
only weakly alutaceous, but in every specimen the punctures are 
distinct. Variation in width of front, ocellar size, and other 
critical measurements is moderate (Table IX). Specimens from 
the extreme southern and eastern ends of the range are rather 
remarkably different in certain measurements. The specimen 
from Conception Bay, Baja California, has the vertex elevated 
far above the eye tops and broadly squared off; the ocelli are 
unusually small and the ocello-ocular line nearly as great as 
the width of the ocellar triangle. One of the specimens from 
northern Baja California shows tendencies in these directions, 
and thus it seems possible that the Conception Bay specimen 
merely represents the end of a cline. The specimen from Valen- 
tine, Texas, has a remarkably broad front and unusually strong 



308 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



notauli. Despite this variation, flavinervis is one of the more 
distinctive species of the genus by virtue of the light, smooth 
antennae, the rather long, weakly hairy eyes, and the punctate 
front. 

Female (assigned here tentatively). — ARIZONA: 4 mi. W. 
Superior, 15 June 1948 (light trap, H. K. Gloyd) [INHS]. 

Description of female. — Length of body 2.7 mm., LH .6 mm., 
LT 1.1 mm. Head dark castaneous, thorax and abdomen medium 
castaneous ; apical half of mandibles, clypeus, and antennae light 
yellowish -brown ; legs yellowish-brown. Mandibles with four 
teeth, third tooth and particularly fourth tooth very small 
(about as in Fig. 38). Clypeus weakly emarginate medially, 
median carina strong. Head 1.4 X as long as wide, sides weakly, 
gradually convergent to near posterior margin, where they are 
more abruptly, arcuately convergent. Eyes small, paler than 
background, no larger than a head puncture. Punctures of front 
absent from median strip, well separated laterally and posteriorly 
(mostly by slightly more than their own diameters), anteriorly 
decidedly crowded (separated by less than their own diameters) ; 
front alutaceous and with a weak tendency to be longitudinally 
striate (less evident on sides than submedially) ; under side of 
head strongly alutaceous, punctate. Pronotal disc 1.4 X as long 
as wide; mesonotum slender, 1.7 X as long as wide; propodeum 
slender, gradually tapered anteriorly, about 1.7 X as long as 









TABLE 


IX 








Species and locality 


No. 


LFW 


WF/HE 




00L/W0T 


DAO/WF 


Ant. 


11 L/W 


flavinervis 


















Lone Star, Cal. 


1 


3.2 


1.10 




.44 


.28 


1.3 




Mojave, Cal. 


2 


3.1 (3.0-3.2) 


1.18(1.16-1 


.20) 


.45 


.26 (.24-, 28) 


1.2 




Llano, Cal. 


2 


2.8 


1.13 




.47 (.44-. 50) 


.25 (.24-. 26) 


1.3 




Hlncley, Cal. 


2 


3.0(2.9-3.2) 


1.13 




.39 (.38-. 40) 


.25 


1.3 




Vlctorvllie, Cal. 


1 


3.1 


1.18 




.49 


.23 


1.3 




Thermal, Cal. 


2 


2.7(2.5-2.9) 


1.10(1.08-1 


.12) 


.58 (.53-. 64) 


.22 (.20-. 24) 


1.3 




Blythe, Cal. 


12 


2.4(2.0-2.8) 


1.09(1.05-1 


.13) 


.55 (.50-. 60) 


.24(.22-.26) 


1.2(1 


,1-1.3) 


El Centre and vie. 


39 


2.9(2.2-3.3) 


1.07( .97-1 


.18) 


.47 (.38-. 56) 


.25 (.23-. 30) 


1.3(1 


.2-1.5) 


Wlnterhaven, Cal. 


1 


2.9 


1.12 




.63 


.23 


1.3 




San Felipe, Baja Cal. 


2 


2.9(2.6-3.2) 


1.10(1.00-1 


• 20) 


.57(.46-.68) 


.25 (.23-. 27) 


1.3(1 


.2-1.4) 


Conception Bay, B.C. 


1 


2.5 


1.29 




.89 


.17 


1.3 




Hermostllo, Sonora 


4 


2.4(2.0-2.7) 


1.12(1.05-1 


.17) 


.52 (.46-. 55) 


.23 (.22-. 25) 


1.1 




Yuma Co., Ariz. 


11 


2.6(2.1-3.0) 


1.12.(1.04-1 


.21) 


.51 (.45-. 67) 


.24 (.21-. 25) 


1.3(1. 


1-1.4) 


Maricopa Co., Ariz. 


10 


2.7(2.3-3.2) 


1.10( .98-1 


.23) 


.46 (.37-. 59) 


.24 (.22-. 25) 


1.3(1 


1-1.4) 


Gila Co., Ariz. 


7 


2.5(2.4-2.6) 


1.18(1.12-1 


.24) 


.58 (.54-. 63) 


.23 (.22-. 24) 


1.3(1, 


2-1.4) 


Pinal Co., Ariz. 


135 


2.5(1.6-3.1) 


1.13(1.00-1 


.24) 


.57 (.48-. 62) 


.23 (.20-. 26) 


1.3(1. 


1-1.4) 


Pima Co., Ariz. 


12 


2.7(2.3-3.2) 


1.20(1.11-1 


.27) 


.59 (.46-. 72) 


.22 (.20-. 26) 


1.2(1, 


1-1.3) 


Las Vegas, Nev. 


1 


3.0 


1.07 




.40 


.27 


1.2 




Valentine, Texas 


1 


2.5 


1.52 




.78 


.19 


1.2 




rectangulatum 


















Sidney, Nebr. 


1 


2.9 


1.40 




.72 


.22 


1.2 




Grant Co. , Kansas 


1 


2.5 


1.42 




• 82 


.19 


1.2 




Wilbarger Co., Tex. 


3 


2.1 (2.0-2.2) 


1.31 (1.29-1 


.32) 


.77 (.75-. 79) 


.21 (.19-. 22) 


1.2 




Victoria, Tex. 


1 


2.4 


1.30 




.69 


.20 


1.0 




Valleclllo, N. Leon 


1 


2.5 


1.08 




.60 


.24 


1.1 




macrops 


















Port Isabel, Tex. 


2 


2.9 


.98 (.94-1. 


02) 


.29 (.28-. 30) 


.36 (.35-. 37) 


1.2 




San Juan, Tex. 


1 


2.8 


1.05 




.33 


.36 


1.2 





EVANS : REVISION OF PSEUDISOBRACHIUM 309 

wide. Pronotal disc strongly shining, with sparse, rather weak 
punctures ; mesonotum wholly although weak alutaceous ; pro- 
podeum also wholly alutaceous, although very weakly so medially 
and strongly shining here, sides obscurely punctate. Mesopleu- 
ruin alutaceous, obscurely punctate. Hairs of body and legs 
short, pale. 

Remarks. — The female described above was taken in a light 
trap at the same time and place as a long series of males (also 
some males of obscurum) . I feel fairly certain that it properly 
belongs with flavinervis, less certain as to how to clearly dis- 
tinguish it from other females of this complex. The distribution 
of this species is shown on Map 3. 

43. Pseudisobrachium macrops new species 

Holotype. — S , Port Isabel, Cameron Co., Texas, 23-27 June 
1956 (at light, H. E. Evans and E. G. Matthews) [MCZ, No. 
30292]. 

Description. — Length 4.4 mm. ; LFW 2.9 mm. Head black, 
thorax dark reddish-brown, abdomen bright rufo-castaneous ; 
mandibles light brown, teeth rufous ; scape light brown ; flagellum 
dull rufo-castaneous; legs, including all coxae, pale straw-col- 
ored; wings hyaline, veins and stigma brown. Mandibles with 
third tooth small, fourth broad and arching into inner mandibu- 
lar margin. Clypeus truncate apically, median carina weakly 
arched in profile. First four antennal segments in ratio of about 
20 :5 :9 :9, segment three 1.5 X as long as thick, segment eleven 
1.2 X as long as thick; pubescence of flagellum pale, rather 
coarse, semi-erect, erect setae fairly numerous, toward the base 
some of them nearly half as long as thickness of flagellum. WF 
.56 X WH, 0.94 X HE ; ocelli remarkably large and convex, 
DAO .37 X WF ; OOL .30 X WOT, actually less than diameter 
of an ocellus ; front angle of ocellar triangle greater than a right 
angle. Vertex elevated above eye tops a distance equal to about 
half HE ; eyes distinctly more hairy than in flavinervis. Front 
strongly alutaceous, rather weakly shining, punctures shallow 
and less conspicuous than in flavinervis, separated from one an- 
other by from 1 to 1.5 X their own diameters. Mesoscutum 
somewhat more strongly shining and more distinctly punctate 
than pronotum ; notauli strong on anterior .4 of mesoscutum ; 
disc of scutellum strongly shining. Propodeum 1.35 X as long 
as broad, disc weakly alutaceous, shining, median carina strong ; 
spiracles elliptical, directed somewhat dorsad. Mesopleural callus 



310 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY 

elongate, moderately convex, shining, somewhat alutaceous ; an- 
terior and ventral portions of mesopleurum more strongly alu- 
taceous and with some punctures. Fore wing with discoidal vein 
only very weakly indicated, arising somewhat below junction of 
basal and transverse median veins. 

Paratypes. — TEXAS: 1 $, same data as type [CU] ; 1 S, 
San Juan, Hidalgo Co., 28 June 1938 (L. W. Hepner) [KU]. 

Variation. — In both paratypes the front is relatively a little 
wider than in the type. In the San Juan specimen the mesoscu- 
tum is not noticeably more shining or punctate than the pronotum 
and the punctures of the front are smaller. 

Anomalous Males 

Under this heading are considered three species known from 
males only, none of which fit well into any of the six species- 
groups recognized. The first two of these species possess five- 
toothed mandibles and genitalia of unusual form as well as 
certain unique features ; I would expect these two species to 
have evolved from a primitive stock independently of each other 
and of other known species. The third species, superbum, has 
only three mandibular teeth and the median carina of the pro- 
podeum replaced by some weak, irregular longitudinal rugae ; 
the genitalia of this species are of the conventional Pseudiso- 
brachium type. I would regard this as one of the more highly 
evolved members of the genus. 

44. Pseudisobrachium petiolatum new species 

Holotype. — $, Tabernilla, Canal Zone, 27 April 1907 (Au- 
gust Busck) [USNM, no. 65385]. 

Description. — Length 3 mm. ; LFW 2 mm. Head piceous ; 
thorax and abdomen castaneous, petiole somewhat infuscated ; 
legs light brown ; antennae medium brown ; fore wing very faintly 
clouded, veins and stigma brown. Mandibles with five sharp 
teeth in an oblique series (Fig. 7). Clypeus broadly rounded 
apically, its median carina sharp except at base and apex. An- 
tennae very long, flagellum clothed with semirecumbent pubes- 
cence of moderate length plus a few longer, suberect setae ; first 
four antennal segments in a ratio of about 4 :1 :4 :3 ; antennal 
segment eleven about twice as long as thick. WF .56 X WH, 
about equal to HE ; OOL 1.10 X WOT, ocelli of moderate size, 
in a compact triangle, anterior ocellus situated well above eye 



EVANS: REVISION OF PSEUDISOBRACHIUM 311 

tops. Front strongly alutaceous but somewhat shining, the punc- 
tures shallow and relatively inconspicuous. Occipital carina 
complete. Pronotum long, collar subt'oveolate. Mesoscutum alu- 
taceous, weakly shining; notauli strong on anterior .7, absent 
behind; scutellum wholly alutaceous, anterior groove and lateral 
foveae rather shallow. Propodeum 1.6 X as long as wide, median 
carina weak, extending for about half its length ; lateral carinae 
strong, the groove above them subfoveolate ; spiracles very large, 
subcircular, directed dorsad. Mesopleurum wholly alutaceous, 
the front half roughened by large punctures. Fore wing with 
discoidal vein strong, arising well down on transverse median 
vein; basal vein erect, reaching subcosta far basad of stigma 
(Fig. 52). First abdominal segment with a rather long petiole 
(Fig. 68). Genitalia with the parameres deeply divided into two 
lobes; digitus and cuspis complex, the latter somewhat disc- 
shaped, with marginal setae ; basis volsellaris weakly developed 
and without any evidence of a vannus ; aedoeagus simple, deeply 
divided apically, one half somewhat angled (probably an arti- 
fact) (Fig. 65). 

Remarks. — This unusual species has a clypeus and genitalia 
unlike those of any other species, but more like those of other 
genera of Pristocerini. The complete occipital carina suggests 
Propristocera, and the petiolate abdomen is much like that of 
P. laevigata. However, the hairy eyes, type of antennal pubes- 
cence, and many features of the genitalia suggest that this species 
properly belongs in Pseudisobrachium. 

45. Pseudisobrachium anomalum new species 

Holotype.— $, Arlington, Va., July 19, 1952 (K. V. Krom- 
bein) [USNM, no. 65386]. 

Description. — -Length 3.5 mm.; LFW 2.2 mm. Head black; 
thorax and abdomen dark brownish-fuscous ; legs pale castaneous, 
except front coxae infuscated ; mandibles yellowish, infuscated 
basally, teeth rufous ; antennae yellowish-brown, somewhat 
darker beyond basal four segments ; wings hyaline, stigma brown, 
veins very light brown. Mandibles with five strong teeth in 
an oblique series (Fig. 8). Median lobe of clypeus weakly emar- 
ginate apically. Antennae rather short, flagellum with rather 
coarse, semirecumbent pubescence and a few erect setae which 
are nearly half as long as width of flagellum ; first four antennal 
segments in a ratio of about 15:5:7:6; segment eleven about 
1.3 X as long as thick. WF .66 X WH, 1.35 X HE ; OOL 1.3 X 



312 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

WOT, ocelli small, in a compact triangle, the front ocellus well 
above level of eye tops. Front rather strongly shining, but 
slightly duller and more noticeably alutaceous anteriorly and also 
behind ocellar triangle ; punctures strong, especially so above 
the eyes, separated from one another by about their own diam- 
eters. Pronotum short, rather roughly punctate. Mesoscutum 
strongly shining, especially between notauli, where it is not at all 
alutaceous ; mesoscutal punctures strong, very sparse medially, 
more crowded laterally; notauli impressed on anterior .9 of 
mesoscutum. Scutellum shining, punctate, the basal groove deep, 
the lateral foveae shallow. Propodeum 1.3 X as long as broad; 
median carina distinct, dorsal surface otherwise completely 
covered with a reticulum of rather strong ridges ; spiracles 
small, elliptical, directed laterad. Mesopleural callus shining, 
non-alutaceous, the remainder of the mesopleura more or less 
alutaceous and pitted. Fore wing with transverse median vein 
subereet, weakly arched; discoidal vein absent (Fig. 54). Abdo- 
men sessile, rather short and broad. Aedoeagus strongly com- 
pressed, of complex structure ; inner arm of paramere very 
slender, digitiform ; vannus strong, bearing the radiating ridges 
characteristic of all species of Pscudisobrachium except petiola- 
tum (Figs. 64, 67). 

Parages. --NEW JERSEY: 1 $, Moorestown, 23 July 
1939 (H. & M. Townes) [HKT]. FLORIDA: 1 $, Okaloosa 
Co., 31 July 1955 (F. W. Mead) [FSPB]. ILLINOIS: 1 $, 
Alto Pass 13 Aug. 1891 (C. A. Hart) [INHS]. 

Variation. — The paratypes are all slightly smaller than the 
type (LFW 2.0-2.1 mm.). In the New Jersey specimen the head 
and thorax are dark castaneous, the abdomen somewhat lighter ; 
in the Illinois specimen the entire body is castaneous ; in the 
Florida specimen the body is wholly nearly black and the femora 
and coxae somewhat infuscated. The antennae of the Florida 
specimen are bright yellowish-brown with a tinge of rufous. 
In the New Jersey and Florida specimens the antennae are 
shorter than in the type, segment eleven measuring 1.1-1.2 X 
as long as thick. WF measures .67-.68 X WH, 1.39-1.45 X HE. 
In the Florida specimen the ocellar triangle is rather wide, OOL 
only 1.1 X WOT. In the New Jersey specimen the posterior 
half of the mesoscutum is weakly alutaceous, even between the 
notauli, as are the mesopleural calli ; this specimen is lacking 
all but the basal three segments of the gaster. In the New Jersey 
and Illinois specimens the pronotum has a shallow transverse 
groove before the posterior margin. 



EVANS : REVISION OF PSEUDISOBRACHIUM 313 

16. Pseudisobrachium superbum new species 

Holotype. — S , Cano Saddle, Gatun Lake, Panama, 17 May 
1923 (K. C. Shannon) [USNM, no. 65387]. 

Description. — Length 5.2 mm.; LFW 3.8 mm. Head and 
thorax shining' black, propodeum piceous, abdomen piceous ex- 
cept suffused with pale reddish-brown laterobasally ; mandibles 
yellow, teeth rufous ; scape yellow, fiagellum light reddish-brown ; 
legs entirely straw -yellow, tarsi slightly darker than the rest; 
wings subhyaline, veins and stigma brown. Mandibles slender, 
with only three apical teeth, the basal tooth rounded (Fig. 32). 
Clypeus unusually large, broadly truncate apically. Antennae 
of moderate length ; first four segments in a ratio of about 
15:3:11:8, segment eleven twice as long as thick; flagellar pu- 
bescence pale, appressed, erect setae numerous, short, some of 
them extremely thin. WF .61 X WH, 1.15 X HE ; ocelli of mod- 
erate size, DAO .16 X WF ; ocelli in a compact triangle, the front 
angle less than a right angle ; OOL 1.18 X WOT. Vertex nar- 
rowly rounded, extending above eye tops a distance equal to 
slightly more than half the eye height; occipital carina com- 
plete. Front strongly shining, non-alutaceous, with small but 
sharply defined and deep punctures; these punctures are close 
together on the lower front, but on the vertex they are separated 
by about three times their own diameters. Pronotal disc unusu- 
ally flat, unusually wide in front, shining and with strong, sparse 
punctures. Mesoscutum strongly shining, non-alutaceous, with 
only a few punctures, these mostly on the sides; notauli very 
strong, complete. Disc of scutellum strongly shining medially, 
laterally with some minute punctures. Propodeum about 1.3 X 
as long as broad, its entire dorsal surface crossed by close trans- 
verse ridges, medio-basally with several irregular longitudinal 
ridges, but without a well-defined single median carina ; spiracles 
nearly circular, directed dorsad. Mesopleurum shining, non- 
alutaceous, with large, widely separated punctures except on the 
callus, which is smooth. Genitalia differing from the usual form 
of the genus only in that the dorsal branch of the paramere is 
rather suddenly contracted subapically (Fig. 66). Discoidal 
vein of fore wing weakly pigmented for a short distance, inter- 
stitial with median vein; stigma unusually large (Fig. 51). 

Paratype. — 1 $, Pacora, Canal Zone, May 13, 1953 (F. S. 
Blanton) [USNM]. 

Variation. — The single paratype is slightly larger than the 



314 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

type, measuring 6 mm. in length, fore wing 4 mm. The pro- 
podeum is darker than in the type, nearly black, while the 
abdomen is uniformly dark reddish-brown. There are no other 
noticeable differences, and head measurements are virtually 
the same as in the type. 

Unassigned Females 

The remaining five species are known from females only. In 
all probability some of these names will fall in synonymy when 
the sexes have been properly associated. 
47. Pseudisobrachium gigas new species 

Holotype. — 5 , Barro Colorado Island, Canal Zone, Sept. 
1941 (J. Zetek) [USNM, no. 65156]. 

Description. — Length 6.5 mm., LH 1.4 mm., LT 2.4 mm. 
Head and thorax dark reddish-brown, almost piceous, abdomen 
dark reddish brown, blotched with paler brown, especially lat- 
erally and apically ; mandibles light castaneous, teeth rufous ; 
antennae bright castaneous, segments 3-12 with an apical annulus 
of dark brown ; legs wholly bright castaneous ; sting-palps yellow- 
ish. Mandibles with four strong teeth (Fig. 33). Clypeus with 
a strong median ridge which is continued past the truncate 
apical margin as a sharp median tooth. Head 1.15 X as long 
as wide ; sides nearly parallel until just before posterior margin, 
where they are arcuately contracted ; vertex straight across, occi- 
pital carina obsolete dorsally. Eye in form of a single distinct 
lens which is white and strongly contrasting to the brown head ; 
eye larger than one of punctures of head. Head punctures 
strong, slightly longer than wide, rather evenly spaced, space 
between them, on the average, about 1-1.5 X greatest width of a 
puncture ; space between punctures flat, polished, non-alutaceous 
and with no sculpturing whatever. Pronotum relatively flat, sides 
of disc rounded; disc 1.15 X as long as wide; pro- and mesonota 
shining, non-alutaceous, devoid of punctures medially but with 
sparse punctures laterally; mesopleura shining and with evenly 
spaced punctures. Propodeum with disc strongly shining, non- 
alutaceous, with a few punctures on the extreme sides, sides sub- 
carinate ; spiracles circular, directed dorso-laterad. 

Paratype. — 9 , Barro Colorado Island, Canal Zone, 29-31 
March 1924 (J. C. Bradley) [CU]. 

Variation. — The paratype is remarkably similar to the type 
in size, color, and all structural details. The measurements pre- 
sented for the type apply equally well to this specimen. 



EVANS : REVISION OF PSEUDISOBRACHIUM 315 

Remarks. — • The male of this species must be a remarkably 
large Pseudisobrachium indeed. It will doubtless be found to 
have five-toothed mandibles and may well run to the crassum 
group in the keys presented here. 

48. Pseudisobrachium zeteki new species 

Ilolotype. — 2 , Barro Colorado Island, Canal Zone, July 
1941 (J. Zetek) [USNM, no. 65157]. 

Description. — Length 3.6 mm., LH .75 mm., LT 1.4 mm. 
Head and thorax dark reddish-brown, abdomen yellowish-brown, 
each of basal five tergites with darker bands basally and apically ; 
mandibles yellowish-brown, suffused with rufous basally and 
apically; antennae castaneous, flagellum fading to straw-yellow 
apically; legs bright yellowish-brown except front coxae brown- 
ish. Mandibles with four strong teeth, third tooth actually 
slightly smaller than basal tooth (Fig. 34). Clypeus with median 
ridge very strong, though not continued past the apical margin 
as a tooth. Head 1.3 X as long as wide; sides nearly parallel 
until just before posterior margin, where they are arcuately 
contracted ; vertex straight across, occipital carina prominent on 
crest of vertex. Eye a single large amber-colored facet which 
contrasts moderately with the brown head. Punctures of head 
strong, anteriorly separated by no more than their own diameters, 
posteriorly more widely spaced, largely absent from center of 
upper front; surface between punctures very strongly polished, 
non-alutaceous. Pronotum less flattened than in gigcts, rather 
slender; disc 1.28 X as long as wide; mesonotum 1.6 X as long 
as wide ; propodeum 1.6 X as long as wide. Thoracic dorsum 
wholly strongly shining, non-alutaceous ; punctures small, re- 
stricted to sides of nota and propodeal disc. Mesopleurum with 
sparse punctures, strongly shining except somewhat alutaceous 
below; sides of propodeum also somewhat alutaceous. 

Remarks. — The male of this species may be cly peat urn or, less 
probably, rettenmeyeri or coopcri. Another possibility is super- 
bum, which also has a complete occipital carina, but only three- 
toothed mandibles. 

49. Pseudisobrachium manni new species 

Ilolotype.— 2, Mixico, 7 Guatemala, 24 May (W. M. Mann) 
[USNM, no. 65158] . 

1 I have been unable to find this locality on maps available to me. There 
are two village** called Mixco, one in the central highlands in the state of 
Guatemala, one in the eastern lowlands in the state of Izabal. 



316 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Description. — Length 4.0 mm., LH .9 mm., LT 1.5 mm. Head 
black, thorax piceous, grading into black dorsally, abdomen fer- 
rugino-castaneous ; mandibles, clypens, and scape bright castane- 
ous, flagellum dull castaneous, apical segment yellowish ; front 
coxae piceous, legs otherwise bright straw-yellow. Mandibles 
relatively slender, two apical teeth strong, third tooth slightly 
smaller and somewhat recessed, basal tooth small, much recessed 
(Fig. 35). Clypeus broadly truncate apically, median elevation 
strong but somewhat round-topped, smoothly declivous well be- 
fore apical margin. Head 1.25 X as long as wide ; sides nearly 
parallel, behind arcuately convergent to vertex, which is straight ; 
occipital carina obsolescent dorsally. Eye a single large facet 
which is pale and strongly contrasting to the black head. Front 
with strong punctures except along midline ; punctures anteriorly 
separated by less than their own diameters, laterally (around 
eyes) distinctly striatopunctate ; punctures posteriorly slightly 
smaller, separated from one another by about their own diam- 
eters ; surface of front between punctures strongly polished, non- 
alutaceous or very nearly so ; under surface of head with strong, 
rather evenly spaced punctures. Pronotal disc 1.2 X as long as 
wide ; mesonotum 1.6 X as long as wide ; propodeum short, only 
1.3 X as long as wide. Thoracic dorsum strongly shining, pro- 
notum and propodeum weakly alutaceous posteriorly ; pronotum 
with numerous punctures, well distributed but absent from mid- 
line ; mesonotum and propodeum with punctures absent from 
broad median area. Mesopleurum alutaceous and with large, 
almost contiguous punctures. Sides of propodeum minutely alu- 
taceo-striolate ; propodeal spiracles subcircular, opening laterally. 

Remarks. — The male of this striking form may possibly be 
a member of the crassum group such as perpunctatum or dalmati. 
On the other hand, the mandibles are more like those of supposed 
females of the occidentals and obscurant groups. However, no 
members of either of these groups are currently known from 
Guatemala. 

50. PSEUDISOBRACHIUM PAUCIPUNCTATUM Fouts 

Pseudisobrachium paucipunctata Fouts, 1928, Proc. Ent. Soc. Wash., 30: 
122. [Type: $, Salt Lake, Utah, 13 June (USNM, no. 62549)]. 
Description of type. — Length 2.9 mm., LH .6 mm., LT 1.1 mm. 
Head and thorax castaneous, abdomen pale yellowish-brown; 
mandibles, clypeus, and scape light yellowish-brown, flagellum 
straw-colored ; legs wholly straw-colored. Mandibles as figured 



EVANS: REVISION OF PSEUDISOBRACHIUM 317 

for obscurum (Fig. 36). Clypeus somewhat emarginate apically, 
median carina strong. Head 1.36 X as long as wide, sides slightly 
but perceptibly narrowed to just before posterior margin, where 
they are arcuately contracted to a broad, straight vertex ; occipi- 
tal carina obsolete dorsally. Eyes barely contrasting to front. 
Front moderately alutaceous anteriorly, more weakly so behind ; 
punctures small, somewhat elongate, separated by about their 
own diameters except more sparse medially and posteriorly; 
under side of head alutaceous, weakly punctate. Pronotal disc 
1.6 X as long as wide; mesonotum 1.7 X as long as wide; pro- 
podeum 1.6 X as long as wide. Pronotum shining, weakly alu- 
taceous, punctures small and well separated, absent from median 
area; mesonotum shining, weakly alutaceous, with some weak 
lateral punctures. Propodeum wholly weakly alutaceous, some- 
what shining, weakly punctate laterally; spiracles circular, di- 
rected dorso-laterad. Mesopleurum alutaceous, weakly punctate. 
Body and legs clothed with short, pale setae. 

Other females. — CALIFORNIA : 1, Yucaipa, 11 May 1938 
(peach orchard, Christenson) [USNM]. 

51. Pseudisobrachium costaricanum new species 

Holotype. — $ , Hamburg Farm, Santa Clara Prov., Costa 
Rica, 30 Sept. 1926, F. Nevermann) [USNM, no. 65388]. 

Description. — -Length 3.4 mm., LH .82 mm., LT 1.4 mm. 
Head and thorax piceous, abdomen bright rufo-castaneous ex- 
cept segments more yellowish apically and laterally ; mandibles 
yellowish, teeth rufous; scape bright yellowish, flagellum dull, 
pale yellowish-brown ; front coxae reddish-brown, legs otherwise 
bright yellowish-brown. Mandibles slender, with three teeth, 
basal tooth large and projecting (Fig. 42). Clypeus with apical 
margin strongly emarginate, median ridge very strong and pro- 
jecting as a small tooth over apical margin. Head 1.22 X as 
long as wide, sides slightly but perceptibly narrowed to just 
before posterior margin, where they are arcuately contracted to 
a broad, straight vertex ; occipital carina obsolete dorsally. Eye 
consisting of a large, white facet which contrasts strongly to 
head color. Front strongly shining, non-alutaceous, with elongate 
punctures which are separated on posterior part by less than 
their own diameters (except more sparse medially), on anterior 
part separated by much less than their own diameters, on sides 
somewhat striato-punctate ; under side of head shining, with 
punctures rather evenly spaced. Pronotal disc 1.4 X as long as 



318 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

wide ; mesonotum 1.5 X as long as wide ; propodeum 1.4 X 
as long as wide. Dises of pro- and mesonota strongly polished, 
non-alutaceous over most of central area, with weak punctures 
which are largely absent from central area ; disc of propodeum 
strongly polished, obscurely alutaceous behind. Sides of meso- 
pleurum strongly alutaceous, weakly punctate. Body hairs pale, 
moderately long, those on the coxae and femora shorter than 
those on body and on apical parts of legs. 

Paratype. — 1 9, same data as type [USNM]. 

Variation. — The paratype is somewhat smaller, length of body 
3.1 mm., of head .7 mm., of thorax 1.2 mm. The head is a bit 
more slender, measuring 1.28 X as long as broad; the punctures 
are somewhat less crowded than in the type, especially anteriorly. 
In all other respects the paratype resembles the type very 
closely. 

LITEEATUBE CITED 

ASHMEAD, W. H. 

1893. Monograph of the North American Proctotrypidae. Bull. U. S. 
Nat. Mus., 45: 472 pp., 18 pis. 
Bruch, C. 

1917a. Insectos mirmecophilos. Physis, Bev. Soc. Arg. Cienc. Nat., 3: 

141-149. 
1917b. Nuevas capturas de insectos mirmecofilos. Physis, Eev. Soc. 
Arg. Cienc. Nat., 3: 458-465. 
Chitty, A. J. 

1906. Pseudisobrachium cantianum : a species of Bethylinae (Procto- 
trypidae) new to science. Ent. Mon. Mag., 42: 148-151. 
Duchaussoy, A. 

1916. Nouveaux Bethylides de l'Afrique Nord et de 1 'Europe orientale. 
Alger. Bull. Soc. Nat. Hist., 7: 109-126. 
Fouts, E. M. 

1928. Notes on the Bethylinae with descriptions of one new Cuban 
and twelve new North American species. Proc. Ent. Soc. Wash., 
30: 121-132. 

KlEFFER, J. J. 

1906. Beschreibung neuer Proctotrypiden aus Nord- und Zentral- 
amerika. Berlin. Ent. Zeitschr., 50: 237-290. 

1914. Bethylidae. Genus Pseudisobrachium Kieffer. Das Tierreich, 41: 
472-484. 

1922. Philippine Serphidae (Proctotrypidae). Philipp. Jour. Sci., 20: 
65-103. 
Mann, W. M. 

1915. Some myrmecophilous insects from Hayti. Psyche, 22: 161-166. 
Eeid, J. A. 

1941. The thorax of the wingless and short-winged Hymenoptera. 
Trans. E. Ent, Soc. London, 91 : 367-446. 



PLATES 



PLATE 1 

Heads of male Pseudisobrachium, anterior view with antennae omitted 
(not drawn to same scale). 
Fig. 1. P. ashmeadi n. sp., paratype 
Fig. 2. P. persimile n. sp., paratype 
Fig. 3. P. hurdi n. sp., paratype 
Fig. 4. P. minutissimum n. sp., holotype 
Fig. 5. P. rufivenire (Ashmead), allotype 
Fig. 6. P. flavinervis Fouts. 



PLATE 2 

Apex of mandibles of male Pseudisobrachium (not drawn to same scale). 
Fig. 7. P. petiolatum n. sp., holotype 
Fig. 8. P. anomalum n. sp., holotype 
Fig. 9. P. micheneri n. sp., holotype 
Fig. 10. P. blomi n. sp., holotype 
Fig. 11. P. olypeatum n. sp., holotype 
Fig. 12. P. cooperi n. sp., holotype 
Fig. 13. P. crassum n. sp., paratype 
Fig. 14. P. occidentale n. sp., paratype 
Fig. 15. P. brunneum n. sp., paratype 
Fig. 16. P. obscurum n. sp., holotype 
Fig. 17. P. pallidum n. sp., holotype 
Fig. 18. P. michoacanum n. sp., holotype 
Fig. 19. P. aztecum n. sp., holotype 
Fig. 20. P. arenarium n. sp., holotype 

Fig. 21. P. prolongatum (Provancher), plesiotype, Ottawa, Ont. 
Fig. 22. P. prolongatum (Provancher), specimen from Mt. Pisgah, N.C. 
Fig. 23. P. ashmeadi n. sp., paratype 
Fig. 24. P. minimum n. sp., holotype 
Fig. 25. P. navajo n. sp., holotype 
Fig. 26. P. minutissimum n. sp., holotype 





8 





10 





12 




13 




14 






17 







21 




22 





24 




25 




26 



PLATE 3 

Figs. 27-32, apex of mandibles of male Pseudisobrachium. Figs. 33-42, 
apex of mandibles of female Pseudisobrachium. Figs. 43-50, clypeus of 
male Pseudisobrachium. Figures not drawn to same scale. 
Fig. 27. P. krombeini n. sp., holotype S 
Fig. 28. P. flavicornis (Kieffer), $ 
Fig. 29. P. foutsi n. sp., holotype $ 
Fig. 30. P. foutsi n. sp., $ from Willcox, Arizona 
Fig. 31. P. flavinervis Fouts, $ 
Fig. 32. P. superbum n. sp., holotype $ 
Fig. 33. P. #i<7as n. sp., holotype 9 
Fig. 34. P. setelci n. sp., holotype $ 
Fig. 35. P. manni n. sp., holotype $ 
Fig. 36. P. obscurum n. sp. (?), 9 
Fig. 37. P. carbonarium (Ashmead) (?), $ 
Fig. 38. P. ashmeadi n. sp. (?), 9 

Fig. 39. P. prolongatum (Provancher), 9 plesiallotype, Toronto, Ont 
Fig. 40. P. rufiventre (Ashmead), 9 
Fig. 41. P. flaviventre (Kieffer) (?), 9 
Fig. 42. P. costaricanum n. sp., holotype 9 
Fig. 43. P. clypeatum n. sp., holotype $ 
Fig. 44. P. rettenmeyeri n. sp., holotype <5 
Fig. 45. P. foutsi n. sp., paratype 5 
Fig. 46. P. apache n. sp., holotype <$ 
Fig. 47. P. cooperi n. sp., holotype <5 
Fig. 48. P. texanum n. sp., holotype 5 
Fig. 49. P. emarginatum n. sp., holotype 5 
Fig. 50. P. krombeini n. sp., holotype (5 








28 







39 






34 





38 



40 





42 




43 




44 



A 

45 





46 



47 





48 



49 



50 



PLATE 4 

Fore wings of male Pseudisobracliium, not drawn to same scale. 
Fig. 51. P. superbum n. sp., holotype 
Fig. 52. P. petiolatum n. sp., holotype 
Fig. 53. P. texanum n. sp., paratype 
Fig. 54. P. anomalum n. sp., holotype 
Fig. 55. P. occidentale n. sp., paratype 
Fig. 56. P. castaneum n. sp., paratype 
Fig. 57. P. prolongatum (Provaneher) 
Fig. 58. P. ashmeadi n. sp., paratype 
Fig. 59. P. oomanche n. sp., paratype 
Fig. 60. P. flaviventre (Kieffer) 
Fig. 61. P. foutsi n. sp., paratype 
Fig. 62. P. flavinervis Fouts 



PLATE 5 

Abdominal structures of male Pseudisobrachium, not drawn to same scale. 
Fig. 63. Genitalia of P. ashmeadi n. sp., paratype, ventral aspect on left side, 

dorsal on right 
Fig. 64. Paramere and volsella of P. anomalum n. sp., holotype, dorso-mesal 

aspect 
Fig. 65. Genitalia of P. petiolatum n. sp., holotype, ventral aspect, right 

paramere and volsella omitted 
Fig. 66. Inner arm of paramere of P. superb um n. sp., holotype 
Fig. 67. Aedoeagus and paramere of P. anomalum n. sp., holotype, lateral 

aspect 
Fig. 68. Base of abdomen of P. petiolatum n. sp., holotype 




68 



Bulletin of the Museum of Comparative Zoology 

AT HARVARD COLLEGE 
Vol. 126, No. 3 



THE CARABID BEETLES OF NEW GUINEA 

PART I. CICINDELINAE, CARABINAE, 
HARPALINAE THROUGH PTEROSTICHINT 



By P. J. Darlington, Jr. 



With Four Plates 



CAMBRIDGE, MASS., U.S.A. 
PRINTED FOR THE MUSEUM 

May 'A, 1962 



Publications Issued by or in Connection 
with THE 

MUSEUM OF COMPARATIVE ZOOLOGY 
AT HARVARD COLLEGE 



Bulletin (octavo) 1863 - - The current volume is Vol. 12(5. 

Breviora (octavo) 1952 — No. 156 is current. 

Memoirs (quarto) 1864-1938 -- Publication was terminated with 
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Johnsonia (quarto) 1941 -- A publication of the Department of 
Mollusks. Vol. 4, no. 40 is current. 

Occasional Papers of the Department op Mollusks (octavo) 
1945 — Vol. 2, no. 28 is current. 

Proceedings of the New England Zoological Club (octavo) 
1899-1948 — Published in connection with the Museum. Publication 
terminated with Vol. 24. 

The continuing publications are issued at irregular intervals in num- 
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Zoology, Cambridge 38, Massachusetts. 



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and future volumes will be published under Museum auspices. 



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Bulletin of the Museum of Comparative Zoology 
AT HARVARD COLLEGE 

Vol. 126, No. 3 



THE CARABID BEETLES OF NEW GUINEA 

PART I. CICINDELINAE, CARABINAE, 
HARPALINAE THROUGH PTEROSTICHINI 



By P. J. Darlington, Jr. 



With Four Plates 



CAMBRIDGE, MASS., U.S.A. 
PRINTED FOR THE MUSEUM 



No. 3 The Carabid Beetles of New Guinea 1 
Part I. Cicindelinae, Carabinae, Harpalinae through 

Pterostichini 

By P. J. Darlington, Jr. 

CONTENTS 

Page 

Introduction 323 

Purpose ; sources of material 323 

Disposition of material ; abbreviations 324 

Previous work on Carabidae of New Guinea and neigh- 
boring areas 324 

History and background of present work 325 

Policies and methods : stages of faunal taxonomy .... 328 

Methods : details 330 

Localities 330 

Ecology 331 

Subspecies 331 

Variation 332 

Taxonomic section 333 

Subfamily Cicindelinae 333 

Collyrini : Tricondyla 334 

Megacephalini : Megacephala 335 

Cicindelini 336 

Prothyma 336 

Caledonomorpha 336 

Distipsidera 337 

Therates 338 

Cicindela 340 

Subfamily Carabinae 351 

Ozaenini : Pseudozaena 351 

Paussini 353 

Arthropterus 354 

Euplatyrhopalus 354 

i Work aided by two fellowships of the John Simon Guggenheim Memorial 
Foundation, 1947 and 1956 ; see pages 327 and 328. 



322 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Scaritini 355 

Geoscaptus 355 

Syleter 356 

Clivina 358 

Subfamily Mormolycinae 397 

Mormolycini : Mormolycc 397 

Subfamily Harpalinae 397 

Apotomini : Apotomus 397 

Bembidiini 398 

Bembidion 399 

Tachys 400 

Limnastis . 484 

Trechini 487 

Perileptns 488 

Perileptodes 489 

Panagaeini 492 

Trichisia 493 

Dischissus 494 

Microschemus 495 

Pterostichiui 497 

Morion 500 

Mecyclothorax 505 

Brachidius 507 

Caelostomus 508 

Cosmodiscus 513 

Homalonesiota 516 

Abacetus 517 

Lesticus 521 

Rhytiferonia 533 

Prosopogmus 536 

Paraloma 538 

• Platycoelus 541 

Haploferonia 547 

Loxandrus 549 

Nebrioferonia 557 

Tiferonia 560 

Catadromus 563 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 323 

INTRODUCTION 

Purpose; sources of material. This is, in taxonomic sequence, 
the first part of a survey of the beetles of the family Carabidae 
of the island of New Guinea. However, it is not the first-pub- 
lished part of the survey. Part 11, covering the Agonini (Aneho- 
menini), has already appeared." Reasons for publishing Part II 
before Part I are given on page 90 of Part II. The present part 
covers the subfamilies and tribes of Carabidae that precede the 
Agonini in the Junk-Schenkling Coleopterorum Catalogus (Pars 
86 by Horn, 1926, and Pars 92, 97, 98, 104, and 112 by Csiki 
1927-1933). Part HI will cover the groups that follow the 
Agonini. These three parts, arranged as numbered, will then 
cover the whole family Carabidae in the order of the Catalogus. 
A fourth part will be necessary for statistics and discussion of 
the New Guinean carabid fauna as a whole and of its geographi- 
cal relationships and origins, and to describe additional species 
that accumulate during the course of the work. 

1 have already listed (Part II, pp. 90-91) the principal sources 
of material used in this survey. Three notable additional lots 
have been received since then. One is a collection of more than 
1,000 specimens of mostly small Carabidae collected in New 
Guinea by the Hungarian entomologist Dr. L. Biro from 1896- 
1902, loaned me for study by the Hungarian National Museum, 
Budapest, through the kindness of Dr. Z. Kaszab. Another is the 
collection made in New Guinea and elsewhere by Dr. E. O. 
Wilson in 1954-1955. This collection is not large (Dr. Wilson 
was concentrating on ants) but it includes a number of inter- 
esting species from new localities. The third is a lot of about 
1400 specimens recently collected in New Guinea by Dr. J. L. 
Gressitt and others ( W. W. Brandt, E. J. Ford, Jr.*, and T. C. 
Maa) for the Bishop Museum. The Bishop Museum is now 
accumulating large collections of insects from New Guinea as 
well as from other parts of the Pacific-Australian area, and 
more Carabidae will certainly be forthcoming, but not in time 
to be included in this part of my work. They will have to b-.' 
dealt with in Parts III and IV. Besides the principal collections 
described here and in Part II, I have received specimens or 
assistance from many other museums and individuals, who will 
be listed in Part IV. 



-Bulletin of the Museum of Comparative Zoology, Vol. 107, No. 3, pp. S7-25L 
with 4 plates, August 1952. 



324 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Disposition of material; abbreviations. Unless otherwise noted, 
specimens collected by myself are in the Museum of Compara- 
tive Zoology, here usually abbreviated as M.C.Z. ; by Miss Chees- 
man, in the British Mus(eum) ; by Toxopeus, returned to the 
Leiden Mus(eum), for distribution; and by Biro, in the Hun- 
garian National Mus(eum). The United States National Mu- 
seum (Washington) is abbreviated as U.S.N.M. ; the American 
Museum of Natural History (New York), as A.M.N. H. Other 
abbreviations are, I think, immediately intelligible. 

Previous work on Carabiclae of New Guinea and neighboring 
areas. Very little has been published on the New Guinean cara- 
bid fauna as such, although a number of species have been 
described or recorded from the island casually or in revisions 
not primarily concerned with New Guinea. However two refer- 
ences are worth giving because they mark efforts of students of 
adjacent carabid faunas to extend their work to New Guinea. 
One is a two-part contribution by an Australian, Sloane, on 
"New Carabiclae from German New Guinea and its Depend- 
encies" and "Further Carabiclae from German New Guinea and 
its Dependencies" published in Deutsche Entomologischcs Zeit- 
schrift for 1907, pp. 177-185, 467-474. Twenty-odd species are 
described or referred to in this work. The other is Andrewes' 
"Catalogue of Indian Insects. Part 18 — Carabidae" (Cal- 
cutta : Government of India, Central Publications Branch, 
1930), which includes all species known from New Guinea in 
many of the Oriental genera (indicated by asterisks) but not in 
all genera. 

Something more should be said about the work done by Sloane 
and Andrewes not on the fauna of New Guinea but on adjacent 
related faunas. 

T. G. Sloane (1858-1932) was an Australian pastoralist, man- 
ager of a sheep station in New South Wales. Study of Car- 
abidae was his avocation. He published about 60 papers on 
Australian carabids between 1881 and 1923, with final notes on 
paussids published posthumously in 1933. His papers include a 
number of important revisions and keys as well as descriptions 
of more than 600 new species. He worked in isolation and was 
not able to study the older types of Australian Carabidae in 
European museums. However his work was done with care and 
good judgment, and went a long way toward putting Australian 
Carabidae in practical working order. His collection was, unfor- 
tunately, allowed to deteriorate during the last years of his life. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 325 

It is now in good care at the Division of Entomology, Com- 
monwealth Scientific and Industrial Research Organization, at 
Canberra. 

H. E. Andrewes (who died in 1950 at the age of 87) was at 
one time employed in the Indian Forest Service but was forced 
to leave India because of eye trouble. He retired from business 
at an early age and deliberately set himself the task of revising 
the Carabidae of the Oriental Region. For this purpose he 
accumulated a large private collection, and worked primarily 
on it and on the collection of the British Museum — he lived 
in London after his retirement. The results of his work were 
122 papers, published from 1919 to 1947, including a series of 
important revisions, faunal lists (Philippines, Sumatra, Mt. 
Kinabalu), the useful catalogue and bibliography of "Indian" 
Carabidae referred to above, two fine volumes on Carabidae in 
the Fauna of British India (unfortunately he was not able to 
complete this work), and descriptions of hundreds of new- 
species. In the course of his work he was able to visit most 
European museums and see the types of most known Oriental 
Carabidae. He was therefore able to put almost all the old 
names in their proper places, and to bring practical taxonomic 
order to the whole Oriental carabid fauna. He did this with 
good, conservative judgment. He left the Oriental carabids in 
shape for further work. His collection has now been added 
to the already outstanding collection of Oriental Carabidae at 
the British Museum. 

Without Sloane's work on Australian Carabidae and An- 
drewes' on Oriental ones it would be impossible to understand 
the nature and geographical relationships of the New Guinean 
carabid fauna. 

History and background of the present work. I was a mem- 
ber of the Harvard Australian Expedition of 1931-1932, and as 
a result of it I began to accumulate a working collection of 
Australian Carabidae at the Museum of Comparative Zoology. 
During that trip I reached the middle part of the Cape York 
Peninsula and felt the pull of New Guinea, although I could not 
go there then. 

During World War II I did go to New Guinea, as an army 
entomologist, and saw most of the occupied localities from Milne 
Bay to Sansapor. Two principal collections of Carabidae were 
made on the island while I was temporarily free of army duties. 
The first was at Dobodura, inland from Oro Bav and not far 



326 



BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 



from Buna, on the north side of Papua. I spent more than 
four months in hospital there. The hospital was in an opening 
in interior rain forest (different from and much richer than 
coastal forest) beside a fine, small river. My right arm was in 
a cast for the first two or three months, but I had a fresh pair 
of heavy army pajamas (almost the equivalent of an untailored 
linen suit) daily, heavy army shoes, enough vials of alcohol, and 
an understanding hospital commander who gave me permission 
to come and go as I liked. I was there from March to July, 
1944, long enough to cover many square miles of rain forest, 
swamp, and grassland. Most of my collecting was done one- 
handed, but practice improved that. It was during this time 
that I gradually learned most of the habitats of Carabidae in 
New Guinea. 

My second principal collection on New Guinea was made on 
the Bismarck Range, North-East New Guinea, where I was 
fortunate enough to be able to go during a two-weeks leave in 
October, 1944, thanks to the courtesy and aid of ANGAU, the 
Australia New Guinea Administrative Unit of the mandated 
area. I flew in from Lae in a C-47, which picked up vegetables 
for army hospitals. The plane landed at the Kerowagi airstrip, 
one of about seven grass airstrips made on the Bismarck Range 



MT. WILHELM SUMMIT 



KEROWAG 




KUNDIAWA 



Sketch map of route to Mt. Wilhelm, Bismarck Bange, North-East New 
Guinea. Scale: one inch = eight miles. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 327 

before the war for use of prospectors and missionaries. The 
strips lie a mile or more above sea level. From Kerowagi I went 
by J ee P to Kundiawa, from there on foot up the valley of the 
Chim River to Toromanbanau, and thence to the summit of Mt. 
Wilhelm, the highest peak in the British half of New Guinea, 
reputedly 15,400 feet. During this trip I walked first through 
grassy, inhabited valleys at altitudes of from about 5,000 to 
7,500 ft., then (above Toromanbanau) through mountain rain 
forest which became low, dense, moss-covered cloud forest 
toward timberline (about 11,000 ft.), then up steep slopes of 
tussock grass past two mountain lakes of glacial origin, and 
finally onto the rocks of the summit. Collecting of Carabidae, 
especially in the mountain forest, was very fine. This is not 
the place to describe this trip in more detail, but I give here a 
sketch map of my route, for the information of other collectors 
and of taxonomists working on my material. 

During the war I collected also some useful material in the 
Philippines. After the Avar, as a preliminary to actual study of 
New Guinean (and Philippine) Carabidae, I made an effort to 
bring together at the Museum of Comparative Zoology a basic 
collection of Carabidae of the Oriental Region and Indo-Austral- 
ian Archipelago, and to become familiar with them. Before the 
war the Museum had purchased several thousand specimens of 
carabids from South India, from Mr. P. Susai Nathan. This 
and some other, older material was put in order and partly 
determined, and additional Oriental Carabidae were obtained 
from other sources. Especially useful were specimens secured 
by exchange from the late H. E. Andrewes (see above), which 
were not numerous but which represented particularly import- 
ant, identified genera and species, and a good set of Javan 
Carabidae received by exchange from Mr. C. J. Louwerens. All 
this would still have left me with a very fragmentary collection 
and insufficient familiarity with Oriental Carabidae if I had 
not been able to spend six months at the British Museum during 
the winter of 1947-48. This was made possible by a fellowship 
from the John Simon Guggenheim Memorial Foundation. At 
the British Museum I was able to see, besides other important 
material, Andrewes' collection of Oriental Carabidae including 
most of his types, and 1 arranged an exchange that added about 
a thousand identified species to the Museum of Comparative 
Zoologv's Oriental carabid collection. 



328 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

As a further preparation for work on New Guinean (and 
Philippine) Carabidae, I compiled an up-to-date list of the 
species of the family known from the Indo-Australian Archi- 
pelago, with a chronological bibliography of each species. The 
list is based on the Junk-Schenkling Catalogue, with additional 
names and references added from other sources. 

Special preparations have been made to relate the Carabidae 
of New Guinea to those of Australia too. The working collection 
of Australian carabids at the M.C.Z., begun in 1931-1932 (see 
above), has been increased by purchase and exchange and by 
a second trip to (eastern) Australia, partly supported by a sec- 
ond fellowship from the Guggenheim Foundation, from Decem- 
ber 1956 to June 1958. During this time my wife, son, and I 
were in the field almost continuously, traveling and living in a 
small truck, except for four months (May through August 
1957) when we were in winter quarters in Canberra. At Can- 
berra, incidentally, I was able to see and study Sloane's collec- 
tion of Australian Carabidae, with most of his types. Our 
field work in 1956-1958 covered the forested eastern edge of 
Australia from southern Tasmania to northern Cape York. The 
Carabidae we obtained in North Queensland, especially on the 
Cape York peninsula in January and May-June 1958, have 
proved especially useful for comparison with New Guinean 
forms. 

In order to complete preparation for work on Australian 
Carabidae, I have compiled a list and bibliography of the 
Australian species based on the Junk-Schenkling Catalogue, 
amplified and brought up to date. 

Policies and methods: stages of faunal taxonomy. Three stages 
can be recognized in taxonomic study of a fauna of any ani- 
mals, although the stages are usually not sharply separated. The 
first stage (comparable to alpha-taxonomy) is the random de- 
scription of species as they happen to fall into the hands of 
specialists. The descriptions are likely to be widely scattered 
in different journals, and the specimens on which they are based 
are likely to be widely scattered in museums and private collec- 
tions. Some of the descriptions are likely to be poor and some 
species are likely to be put in wrong genera or even Avrong tribes. 
Nevertheless, even this initial stage in faunal taxonomy serves 
a useful purpose. The scattered descriptions are listed in the 
Zoological Record, and they advertise to interested persons 
some preliminary information about the nature of the fauna 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 329 

and where specimens from it are preserved. However, there is 
no great loss and often much gain if the first stage is omitted 
and if taxonomists can go directly to the second stage, which 
is to put the fauna as a whole in preliminary order by appropri- 
ate taxonomic methods. This is equivalent to beta-taxonomy. 

By appropriate taxonomic methods I mean methods like those 
of Sloane and Andrewes, and many comparable workers in other 
parts of the world. The methods are essentially subjective : com- 
parison of specimens, detection of similarities and differences, 
and reaching of conclusions based primarily on the personal 
judgment and experience of the taxonomist, not on statistical 
analysis or other objective tests. This kind of taxonomy is not 
necessarily typological. Many taxonomists who practice it are 
conscious that they are dealing with samples of populations 
and are deeply interested in variations in populations. The 
great advantage of this kind of taxonomy is that it can be done 
comparatively rapidly, so that one man can cover a whole large 
fauna in a reasonable time and determine its general composition 
and geographical relationships and something of its origin and 
evolution. This general information is important. To get it — 
that is, to get an understanding of the fauna as a whole — the 
second-stage taxonomist has to bypass too-difficult cases, includ- 
ing eases that cannot be solved with the material immediately 
available or that require time-consuming study of situations 
outside the actual fauna being studied. 

Third-stage taxonomy (equivalent to gamma-taxonomy) is the 
critical study of selected cases that cannot be solved promptly 
and satisfactorily by personal judgment. Some cases require 
world-wide revisions for their solution. Others require laborious 
dissections or study of larvae. Still others require complex 
statistical treatment. And finally some cases may require genetic 
breeding. This kind of work reveals relationships and sibling 
species that second-stage taxonomists miss, and confirms (or 
reverses) second-stage taxonomists' conclusions in many details. 
In other words third-stage taxonomy solves the problems that 
second-stage taxonomy leaves. But good second-stage taxonomy 
shows where problems are and presents them for third-stage 
treatment. 

The present work is second-stage taxonomy. It is an attempt 
to classify all the Carabidae known from New Guinea (but there 
are probably hundreds of species still to be discovered especially 
in the mountains) and in general to put the whole fauna in 



330 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

order within practical limits. The limits are imposed mainly by 
time. If I should set standards too high or allow myself to be 
delayed by details, I would never finish this work as a whole. 

Methods: details. My methods of making descriptions, citing 
localities, making measurements, etc. are described in the intro- 
duction of Part II of the present work (pp. 91ff.). However, 
although I shall try to be reasonably consistent in editorial 
details, I shall not follow a single model exactly but shall vary 
treatment according to the requirements of each group. More- 
over, I expect to vary the treatment to fit the importance or 
interest of each group. I have already (Part II) treated the 
Agonini more thoroughly than I plan to treat any other tribe 
of Carabidae in New Guinea, because I have a special interest 
in this tribe and because the Agonini is the principal tribe that 
has radiated on the high mountains of New Guinea. The present 
part, Part I, includes some smaller groups of special interest, 
including the Pterostichini, some of which have undergone wing 
atrophy and become flightless. On the other hand, the Cicinde- 
linae are outside my usual sphere of interest and (in New 
(iuinea) include no known flightless species except Tricondyla 
aptera, and I have treated them comparatively briefly. But 
these are special cases. The bulk of the present part of my 
work is concerned with small ground -living Carabidae (Clivina, 
Tachys, etc.) which, in New Guinea, have been poorly known 
until now, but of which special collecting methods have j'ielded 
much new material. These will be given something like average 
treatment. Part III will include a larger proportion of previ- 
ously known species, most of them winged, including Harpalini 
which are dominant especially in open areas (but a few occur 
in rain forest), and also many Lebiini which are dominant arbo- 
real Carabidae especially in rain forest (but some occur in 
open areas). The circumstances under which my collecting was 
done, particularly the difficulty of using a net one-handed, lim- 
ited my catch of arboreal forms, so that my material of them 
is comparatively scanty and will be treated comparatively 
briefly. 

Localities. A preliminary outline map of New Guinea with 
important carabid localities was included in Part II (p. 93). 
This map will probably be reproduced in Part IV, with addi- 
tional localities added. One locality, however, should be men- 
tioned now. It is Alfred Russell Wallace's "Dorey" or "Dory." 
Dorey is on the northeast corner of the Yogelkop, and Wallace 



DARLINGTON : CAKABID BEETLES OP NEW GUINEA 331 

did go there (from Ternate) and spent 3 or 4 months there, 
beginning- about the end of March, 1858. However, his specimens 
labeled from Dor(e)y include a number of common Oriental 
species of Carabidae that reach Celebes or the Moluccas but, 
except for Wallace's records, are not recorded from New Guinea. 
I have encountered so many such cases that I am convinced that 
Wallace accidentally mixed his collections and that many of 
his specimens labeled as from Dorey are really from Celebes 
or the Moluccas. Cases here noted include Clivina castanca and 
wallacci, Apotomus, Tachys sericeus and haliploides, Abacctus 
convexiusculus, and Loxandrus celebensis (see under L. medi- 
us). The names of these and other species, and higher groups, 
previously recorded from New Guinea on what I consider doubt- 
ful or erroneous grounds are put in parentheses in the following 
pages. 

Ecology. The ecological information accompanying my speci- 
mens is scanty. It was limited by war-time conditions, by the 
fact that I had little time for collecting and a limited supply of 
vials and alcohol, so that I often had to put many specimens 
together under one label for each locality. I kept a list of 
species recognized, but many of the smaller ones were not dis- 
tinguished in the field. Nevertheless, I can usually say whether 
a particular species was taken in rain forest or not, whether it 
was associated with open water or not, or whether it was 
arboreal. This is not ecology in the detailed, modern sense, but 
it permits a rough ecological classification of the species which 
is useful in analyzing the nature and history of the fauna. 

Subspecies. The concept of subspecies has been critically re- 
examined by several authors recently. The re-examination is 
good in itself, and some criticisms of the subspecies concept 
have been valid, although the criticisms apply more to conti- 
nental situations, where distributions are continuous, than to 
island populations, where distributions are interrupted. New 
Guinea is a very large island, on which three kinds of situa- 
tions occur that can be expressed by trinomials — - by use of 
subspecies. First, a population that is spread over the whole 
of New Guinea may be slightly different from related popula- 
tions on other islands. Second, a species that is widely distributed 
on New Guinea may occur in a habitat that is discontinuous on 
the island. Such a species may be broken into slightly different, 
geographically isolated subpopulations on different mountain 
tops, or in different river valleys, etc. Third, a species may be 



332 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

widely and more or less continuously distributed on New 
Guinea but may vary from locality to locality nevertheless. I 
am prepared to make subspecies in all of these cases, if that 
seems the best way to make the situation clear. In the first two 
kinds of situations listed above, the subspecies are isolated popu- 
lations and are probably often incipient species. In the third 
case, in which geographical variation occurs in presumably con- 
tinuous populations, recognition of subspecies is at least a useful 
technique of second-stage taxonomy, which helps put a whole 
fauna in understandable order. However, I have made sub- 
species very sparingly, and only when I think they really do 
clarify situations. 

Variation. During the writing of this paper I have become 
increasingly impressed by the amount of variation shown by 
many species. Variation of certain characters among Agonini 
has been discussed in Part II (pp. 94ff.). In the present part, 
cases of individual variation are noted that affect supposedly 
specific, generic, or even tribal characters. In the genus Clivina, 
for example, the number of elytral striae that are free at base 
has been supposed to be characteristic of whole species-groups, 
but several species in New Guinea vary individually in this re- 
spect, with either 3 or 4 free striae in different individuals (see 
description of C. kulti, notes under oiroi, and descriptions of 
puncticeps, dealata, rufula, erugata, and subfusa). The number 
of so-called "fixed" punctures on the 3rd elytral interval varies 
individually too in some species of Clivina (see description of 
C. kulti, notes under erugatclla) . In Tacliys truncatus I have 
found what seems to be dimorphism in presence or absence of 
conspicuous foveae on the mentum, a character heretofore sup- 
posed to be constant within species and usually within species- 
groups. Among certain Pterostichini I have found cases of 
individual variation in presence or absence of the basal elytral 
margin (usually considered a generic character, but it varies 
individually in Platycoclus archboldi) and in presence or ab- 
sence of interruptions of the elytral margin (usually considered 
a tribal character of Pterostichini, but individually variable in 
Paraloma for lis). Paraloma, fortis exhibits striking asexual di- 
morphism in form of prothorax. And Microschemus quadri- 
maculatus (tribe Panagaeini) is apparently dimorphic in colora- 
tion. Much other, minor variation is noted in various cases in 
the following pages, but I am sure that what I have detected 
is only a small part of all the variation that really occurs in the 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 333 

species in question. Third-stage taxonomists should discover 
much more of it. A number of cases of wing atrophy and five 
of apparent geographical or individual wing dimorphism (Cli- 
vina dealata and erugatella, Lesticns politus, Platycoelus depres- 
ses, Loxandrus latus) are noted now, but full discussion of 
variation of wings of Carabidae in New Guinea will be post- 
poned to Part IV. 

TAXONOMIC SECTION 
Subfamily CICINDELINAE 

The tiger beetles are treated as a subfamily of Carabidae in 
the Junk-Schenkling Col copter orum Catalogus, and I have in- 
cluded them here for the sake of completeness and because of 
their general interest and zoogeographic importance. However, 
they do not fall within my usual range of study, and I have 
therefore treated them in less detail than the other carabids. 

Seven genera of "tigers" are represented in New Guinea. 
Three of them (Tricondyla, Prothyma, Therates) have probably 
extended to New Guinea from the Orient, more or less recently. 
Two (Megacephala, Distipsidera) have evidently extended from 
Australia to (southern) New Guinea. One (Caledonomorpha, 
with 2 species) is endemic to New Guinea and is in fact known 
only from the eastern part of the island. And the final genus 
(Cicindela) is world-wide and is represented in New Guinea by 
about 8 stocks with various geographical relations (p. 341). 
Tricondyla is flightless. All other tiger beetles in New Guinea 
are winged and able to fly, so far as I know. 

Our present understanding of the classification and general 
distribution of tiger beetles is the result of a life-time of study 
by the late Dr. Walther Horn, who coordinated the works of many 
earlier authors and added immensely to them. Of Horn's many 
publications, the outstanding ones that cover the Cicindelinae 
as a whole are Fascicule 82 of Wytsman's Genera Insectorum 
(1908-1915) and Pars 86 of the Junk-Schenkling Coleoptero- 
rum Catalogus (1926 — in 1959 it was still available from Uit- 
geverij Dr. W. Junk, Van Stolkweg 13, The Hague, Holland). 
In treating the older species of the subfamily I usually refer to 
this Coleop(terorum) Cat(alogus) for synonymy and references 
rather than repeat them in full. On the other hand, I shall not 
take space to cite the Catalogus when it adds nothing to knowl- 
edge of particular species in New Guinea. There is an out-of-date 



334 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

but still useful revision of the "Cicindelidae" of Australia, by 
T. G. Sloane, in the Proceedings of the Linnean Society of New 
South Wales, Vol. 31, pp. 309-360,' pis. XXV-XXXI, 1906. 

Key to Tribes and Genera of Cicindelinae of New Guinea 

1. Metepisterna very narrow, deeply longitudinally channeled ("Aloko- 

sternale Phyle"); 4th tarsal segments asymmetrical (tribe Colly- 
rini) ; elytra soldered together, inner wings minute, vestigial; form 
antlike, 20-25 mm (in New Guinea) (p. 334) Tricondyla 

— Metepisterna wider, not deeply channeled ("Platysternale Phyle"); 

elytra not soldered, inner wings fully developed (in New Guinea) 2 

2. Pronotum with anterior lateral angles (seen from sides) prominent, 

projecting farther forward than anterior margin of prosternum; 
head wide but eyes smaller and less prominent than usual (tribe 

Megacephalini) (p. 335) Megacephala 

Pronotum with anterior lateral angles not prominent (tribe Cicinde- 
lini) 3 

3. Fourth segments of all tarsi very short, wide, densely pubescent below; 

color usually dark purplish or greenish above, often boldly banded 
with yellow (p. 338) ... Therates 

— Fourth tarsal segments slender, not densely pubescent below 4 

4. At least part of lower surface of body with decumbent white pubes- 

cence (p. 340) Cicindcla 

— Lower surface without decumbent pubescence 5 

5. Labrum not toothed; elytra spined (p. 336) Caledonomorpha 

— ■ Labrum toothed ; elytra not spined 6 

6. Clypeus without setae (p. 336) Prothyma 

— Clypeus with a pair of conspicuous setae (p. 337) Distipsidera 

Tribe COLLYRINI 

Genus TRICONDYLA Latreille 

Latreille 1882, in Latreille and Dejean, Hist. Nat. et Iconographie Coleop. 

d 'Europe 1, p. 65. 
Horn 1926, Coleop. Cat., Cicindelinae, p. 22 (see for additional references 

etc.). 

Tricondyla aptera (Olivier) 

Olivier 1790, Entomologie 2, no. 33, p. 7, PI. 1, fig. 1 (Cicindela). 
Horn 1926, Coleop. Cat., Cicindelinae, p. 27 (see for synonymy and addi- 
tional references), 
van Nidek 1959, Nova Guinea (new series) 10, pp. 177, 181. 

Notes. Tricondyla aptera is a well known, antlike, big-eyed, 
flightless tiger beetle, 20-25 mm. long, which ranges (with some 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 335 

variation) from New Guinea (and the Cape York Peninsula 
of Australia) to the Moluccas and Philippines, Timor, and the 
Solomons and New Hebrides. It is a very common species, 
"represented in every consignment from New Guinea and 
surrounding islands" (van Nidek). It runs on tree trunks and 
fallen trees in rain forest by day. The genus is primarily 
Oriental; T. aptera evidently represents a rather recent exten- 
sion of a flightless Oriental stock into the Australian Kegion. 

Tribe MEGACEPHALINI 

Genus MeGACEPHALA Latreille 

Latreille 1802, Hist. Nat. Crustaces et Inseetes 3, p. 79. 
Sloane 1906, Proc. Linn. Soe, New South Wales 31, 317-327. 
Horn 1926, Goleop. Cat., Cicindelinae, p. 61 (see for synonymy and addi- 
tional references). 

Notes. Megacephala {sensu lato) is discontinuously distrib- 
uted, occurring in Africa (not Madagascar) and part of the 
Mediterranean region and southwestern Asia, in Australia and 
southern New Guinea, and in tropical and warm temperate 
America. There are about 20 Australian species, rather diverse 
in form and color, some fully winged, others flightless. They are 
usually found running on the ground near water at night, and 
some of them come to light. The genus is represented in New 
Guinea only by 2 of the winged Australian species that are re- 
corded from Merauke in the south-coastal region of Netherlands 
New Guinea. 

Both the species that reach New Guinea are dark with metallic 
reflections, with the outer margins of the elytra yellow or reddish, 
the pale color extending onto the disc of each elytron about 
Y 3 from base. The following key to the species is based on 
Sloane (loc. cit.) ; I have seen no specimens of this genus from 
New Guinea. 

Key to Species of Megacephala Recorded from New Guinea 

1. Prothorax with lateral margins almost entire, cariniform behind (as 
well as in front of) posterior transverse impression of pronotum 

(p. 336) australasiae hwmeralis 

— Prothorax with lateral margins obliterated posteriorly, not cariniform 
behind posterior transverse impression (p. 336) bostocki 



336 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Megacephala australasiae humeralis Macleay 

Macleay 1863, Trans. Ent. Soc, New South Wales 1, p. 9. 
Horn 1913, Nova Guinea 9, p. 409. 

— — 1926, Coleop. Cat., Cieindelinae, p. 70 (see for synonymy and addi- 
tional references). 
Notes. This species is widely distributed in the northern 
half of Australia and is recorded also from Merauke, southern 
Neth. N.G., by Horn (1913). 



Megacephala bostocki Castelnau 

Castelnau 1867, Trans. R. Soc. Victoria 8, p. 36. 
Horn 1913, Nova Guinea 9, p. 409. 

1926, Coleop. Cat., Cieindelinae, p. 70 (see for synonymy and addi- 
tional references). 
Notes. Like the preceding, this northern Australian species 
is recorded from Merauke, southern Neth. N.G., by Horn (1913). 
It is curious that these two very similar species should be 
known from New Guinea only from a single source, and I sus- 
pect some mistake, although I have no other evidence of it. 



Tribe CICINDELINI 

Genus PrOTHYMA Hope 

Hope 1838, Coleopterist 's Manual 2, pp. 12, 27. 

Horn 1926, Coleop. Cat., Cieindelinae, p. 96 (see for synonymy and addi- 
tional references). 

Prothyma papua Darlington 

Darlington 1947, Psyche 54, p. 242, fig. 2. 

Notes. This species is known only from the north side of 
Milne Bay, Papua, at the eastern tip of New Guinea. Its habits 
are unknown. It represents a primarily African-Oriental genus 
which occurs also in Madagascar, etc., temperate China, and 
Mexico, and which is represented by aberrant subgenera in 
Celebes, northwestern Australia, New Caledonia, Fiji (an un- 
described species) and Samoa. 

Genus CALEDONOMORPHA Horn 

Horn 1897, Ann. Mus. Civ. Genova (Genoa) 37, p. 270. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 337 

Caledonomorpha jordani Horn 

Horn 1897, Ann. Mus. Civ. Genova (Genoa) 37, p. 270. 

1910, in Wytsman, Genera Insectorum, Fase. 82, p. 179, pi. 11, fig. 12. 

1932 Eee. S. Australian Mus. 4, p. 551 (Caledonica) . 

Darlington 1947, Psyche 54, p. 242 [type restriction]. 

Notes. Known localities for jordani are the Astrolabe Mts., 
Mt. Lamington, Dobodnra, and Fergusson Is., all of which are 
in Papua east of 47 °E., on or near what might be called the 
Bird's Tail of New Guinea. Specimens that I collected near 
Dobodnra were running and flying by day on stones and low 
foliage along small, rapid brooks in rain forests in foothills 
of the Owen Stanley Range. 



Caledonomorpha milneana Darlington 

Darlington 1947, Psyche 54, p. 241, fig. 1. 

Notes. This second species of Caledonomorpha is known only 
from the north side of Milne Bay, Papua, at the eastern tip 
of New Guinea, collected Dec. 1943, by myself. 



Genus DISTIPSIDERA Westwood 

Westwood 1837, Mag. Zool. Bot. 1, p. 251. 

Horn 1926, Coleop. Cat., Cicindelinae, p. 105 (see for additional references 
etc. ) . 
Notes. Distipsidera occurs in the northern half of the eastern 
coastal region of Australia and in southern New Guinea. In 
Australia the larger species are usually found hunting (by day) 
on the trunks of Eucalyptus trees in open forest; some of the 
smaller species, on tree trunks in rain forest. One large and 
one small species occur in New Guinea, but their habits there are 
unknown. 



Distipsidera papuana Gestro 

Gestro 1879, Ann. Mus. Civ. Genova (Genoa) 14, p. 556. 

Notes. The types of papuana came from the Katau (Binaturi) 
and Fly R., southern New Guinea (southern Papua). The 
species is described as small, about 12 mm. long, purplish black 
with elytral apices greenish and a pale humeral spot and median 
fascia (not reaching suture) on each elytron. 



338 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

DlSTIPSIDERA THIERFELDERI Horn 

Horn 1925, Ent. Mitteilungen 14, p. 179. 

Notes. Horn's single specimen of this species was from the 
south coast of Neth. N.G. The species is much larger than 
papuana, 19 mm. without the labrum, and is described as black 
with slight purplish or greenish reflections, with pale maculae 
on the elytra. 



Genus THEEATES Latreille 

Latreille 1817, in Cuvier, Begne Animal 3, p. 179. 

Horn 1926, Coleop. Cat., Cicindelinae, p. 110 (see for synonymy and addi- 
tional references). 
Notes. This is a primarily Oriental genus which extends to 
New Guinea and the Solomons but not to Australia. The species 
occur on low foliage in the undergrowth of rain forest. 

Key to Species of Therates Recorded from New Guinea 

1. Color above (except labrum) greenish or bluish without pale mark- 

ings 2 

Conspicuously bicolored 3 

2. Larger, c. 16-21 mm. (p. 338) . . labiatns 

(See also caligatus, p. 340.) 

— Smaller, c. 7% mm. (p. 339) cyaneus 

3. Metallic greenish or bluish, with a conspicuous yellow or reddish band 

across base of elytra (sometimes an additional pale blotch near 
middle of each elytron) 4 

— Not metallic, more extensively pale 5 

4. Larger, c. 13-15 mm. (p. 339) basalis 

— Smaller, c. 9 mm. (p. 339) festivus 

5. Head (except labrum) and prothorax piceous, elytra testaceous with 

a variable, sometimes divided piceous band or blotch behind middle; 
length c. 10-12 mm.; (occurrence in New Guinea doubtful) (p. 340) 

(fasciatus) 

— Color testaceous with front of head and post-median elytral band 

dark; c. 8 mm. (p. 340) chaudoiri 



Therates labiatus (Fabricius) 

Fabricius 1801, Systema Eleutheratorum 1, p. 232 (Cioindela). 
Horn 1926, Coleop. Cat., Cicindelinae, p. 110 (see for synonymy and addi- 
tional references), 
van Nidek 1959, Nova Guinea (new series) 10, pp. 177, 181. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 339 

Notes. T. labiatus is widely distributed and common in New 
Guinea and occurs also on the Aru and Kei Is., and it extends 
west and north to the Moluccas, Celebes, and the Philippines, 
and east to the Solomons. 



Therates basalis Dejean 

Dejean 1826, Species Coleop. 2, p. 437. 

Horn 1926, Coleop. Cat., Cicindelinae, p. Ill (see for synonymy and addi- 
tional references), 
van Nidek 1959, Nova Guinea (new series) 10, pp. 179, 182. 

Notes. This is the only species of the genus besides labiatus 
that is common and widely distributed in New Guinea. It ex- 
tends to several small, close-lying islands including Waigeu and 
Misol (but not farther west), and to the Solomons. Van Nidek 
discusses variation in dentition of the labrum. 



Therates festivus Boisduval 

Boisduval 1835, Voyage Astrolabe, Faune Ent. 2, p. 13. 
Horn 1926, Coleop. Cat., Cicindelinae, p. Ill (see for synonymy and addi- 
tional references). 
[rothschildi Horn 1896, Deutsche Ent. Zeitschrift 1896, p. 150. 
van Nidek 1959, Nova Guinea (new series) 10, pp. 179, 182. 

Notes. Horn (Joe. tit.) records typical festivus from New 
Guinea and Waigeu, and from Misol in the Moluccas ; van 
Xidek, from Misol, and from Soroiig and Japen Is., Neth. N.G. 

The form rothschildi Horn, with a spot near the middle of 
each elytron as well as the basal fascia pale, was described from 
Humboldt Bay, Neth. N.G., and later (1926) listed also from 
Japen Is. ("Jobi") ; van Nidek records another specimen from 
Japen Is. and there is one in the M.C.Z. from lower Busu River, 
Huon Peninsula, N-E. N.G., collected May 4, 1955, in lowland 
rain forest, by E. 0. Wilson. I doubt if rothschildi is even a 
good (geographical) subspecies. 



Therates cyaneus Chaudoir 

Chaudoir 1861, Bull. Soc. Nat. Moscou 34, Part 2, No. 4, p. 357. 
Horn 1926, Coleop. Cat., Cicindelinae, p. 111. 

Notes. New Guinea, Misol, and Celebes is the distribution 
given by Horn (loc. tit.) for cyaneus. 



340 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

TlIERATES CHAUDOIRI Schaum 

Schaum 1860, Berliner Ent, Zeitschrift 4, p. 185, pi. 3, fig. 1. 
Horn 1926, Coleop. Cat., Cicindelinae, p. Ill (see for synonymy and addi- 
tional references). 
Notes. Horn records chaudoiri only from northwestern 
(Neth.) New Guinea and Celebes. 

(Therates pasciatus [Fabricius]) 

Fabricius 1801, Systema Eleutheratorum 1, p. 244 (Cicindela). 
Horn 1926, Coleop. Cat., Cicindelinae, p. 112 (see for synonymy and addi- 
tional references). 
T. fascial us has been recorded from New Guinea but Horn 
questions its occurrence there. It does occur in the Philippines, 
Celebes, and Moluccas east at least to Halmahera. Whether it 
extends to (western?) New Guinea remains to be discovered. 

(Therates dimidiatus Dejean) 

Dejean 1825, Species Coleop. 1, p. 159. 
Horn 1926, Coleop. Cat., Cicindelinae, p. 113. 

T. dimidiatus is another species of which Horn questions old 
records from New Guinea. It inhabits the Malay Peninsula, 
Sumatra, Java, Borneo, and small islands in that vicinity. Its 
occurrence in New Guinea is therefore unlikely. It is a small 
species, resembling festivus but with spined elytra. 

Therates caligatus Bates 

Bates 1872, Ent. Month. Mag. 7, p. 285. 

van Nidek 1959, Nova Guinea (new series) 10, p. 178, fig. 1 (left). 

Notes. This species was described as from the Philippines but 
the locality may be doubtful. Van Nidek records it from the 
islands of Misol and Waigeu just west of the western tip of 
New Guinea; it has not been recorded from the mainland of 
New Guinea itself but may occur there. According to van Nidek 
it resembles labiatus but is smaller, with black tibiae and tarsi, 
and sutural angles of elytra more produced than in labiatus. 

Genus ClClNDELA Linnaeus 

Linnaeus 1758, Systema Naturae, ed. 10, 1, p. 407. 
Horn 1915, in Wytsman, Genera Insectorum Ease. 82, p. 239. 
1926, Coleop. Cat., Cicindelinae, p. 127 (see for synonymy and addi- 
tional references). 



DARLINGTON : CARABTD BEETLES OP NEW GUINEA 341 

Notes. In the usual, broad sense this genus is world-wide 
(excepting some cold places and some remote islands) and in- 
cludes about half of all existing species of tiger beetles — almost 
three-fourths of those of New Guinea. The species of the Austral- 
ian Region, including New Guinea, are separately listed by Horn 
(1915, pp. 311-321; 1926, pp. 193-203), and I have followed his 
arrangement, putting his group-headings in parentheses, and 
interpolating recently described species. 

According to Horn's (second) diagram of the phylogeny and 
geographical history of Cicindela (1915, pi. 23), the species of 
the genus in New Guinea represent about 8 original stocks, listed 
below roughly in order of age, with the oldest first. 

1. The tctrachoides and latreillei groups represent a phylo- 
genetically isolated (old?) stock that is confined to the 
Australian Region and has produced endemic groups in 
Australia as well as on New Guinea. 

2. Cicindela ancorifera, on New Guinea, represents a stock, 
probably originally derived from the Orient, that has di- 
versified especially on New Zealand (sic) and that is 
represented by endemic species-groups on Australia too. 

3. The very small (usually considerably less than 10 mm.) 
species of the placida-funerata, variolosa, boisduvali, and 
giiinecnsis groups have radiated primarily on New Guinea. 
Horn's diagram does not make clear whether or not they 
are derived from the same ancestral stock as No. 2. 

4-5. C. maino and denticollis are endemic species derived from 
Oriental stocks. 

6-8. C. decemguttata, discrcta, and semicincta are widely dis- 
tributed species shared with (and probably recently derived 
from) the Orient. 

There are so many species of this genus on New Guinea and 
I know so few of them that I have not tried to make a key to 
them or even to give recognition characters. Horn characterizes 
the groups in Genera Insectorum (pp. 313-321) and summarizes 
the distinguishing characters of a number of the very small 
species (of the placida-funerata group, etc.) in Deutsche Ent. 
Zeits. for 1904, p. 428. 

Most species of Cicindela hunt (by day) on the ground in 
open places, but a few, especially some small species, occur on 
low foliage of undergrowth of rain forest, often along brooks 
or partly cleared tracks where some sunlight comes through the 
forest canopy. 



342 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

(Group tetrachoides) 

ClCINDELA TETRACHOIDES GestrO 

Gestro 1876, Aim. Mus. Civ. Genova (Genoa) 8, p. 514. 
Horn 1926, Coleop. Cat., Cicindelinae, p. 194 (see for synonymy and addi- 
tional references), 
van Nidek 1953, Psyche 60, p. 155. 

1959, Nova Guinea (new series) 10, pp. 179, 182. 

Notes. Horn lists this species from New Guinea and eastern 
Ceram. In New Guinea, it is common and widely distributed 
at low altitudes, and I found it on the Bismarck Range at 5,000 
ft. or higher. It occurs on the ground in open places and is, 
I think, partly nocturnal. 

Cicindela inaequidens van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 179. 

Notes, van Nidek distinguishes this species (from the pre- 
ceding one) by its more evenly punctulate elytra; sides of elytra 
near apex strongly bent inwards ; sutural angles of elytra dentic- 
ulate in female. The types of this and of the preceding species 
are from Dilo, south coast of Papua, but this as well as the 
preceding species, is widely distributed in New Guinea, and 
this occurs also on Morotai Is., Moluccas. It would be inter- 
esting to know whether their habits differ. 

(Group latreillei) 

Cicindela latreillei Guerin 

Guerin 1830, in Duperrey, Voyage Coquille, Zool. 2, Part 2, First Div., 

p. 57 (latrcillii), Atlas, Ins. pi. 1, fig. 5, a, b. 
Horn 1926, Coleop. Cat., Cicindelinae, p. 194 (see for synonymy and addi- 
tional references), 
van Nidek 1959, Nova Guinea (new series) 10, p. 182. 

Notes. The latreillei group is confined to New Guinea includ- 
ing Japen Is. The species latreillei is recorded from Dorey in 
western Neth. N.G. ; Japen Is.; lower slopes of the Snow Mts. 
at 800-1500 m. (c. 2,600-4,875 ft.) ; and the Fly R,, Papua. 

Cicindela latreillei viridithoracica van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 180. 

Notes. Described from Klamono (Vogelkop) and Geelvink 
Bay, Neth. N. G. The status of this form is not yet clear. It 
can hardly be a geographical subspecies. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 343 

Cicindela velutina van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 182. 

Notes. A species of the latreillei group, described from 10 
specimens collected around Sigi Camp and Lower Mist Camp, 
Snow Mts., Neth. N. G. The altitudes are between 1350 and 
1700 m. (a little below or above 5000 ft.). 

Cicindela viridimicans van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 183. 

Notes. Another new species of the latreillei group, described 
from 15 specimens from Araucaria Camp, Rattan Camp, and 
Lower Mist Camp, on or near the lower slopes of the Snow Mts., 
Neth. N. G. Altitudes are from 800 to 1700 m. (c. 2,600-5,525 
ft.). 

Cicindela alticola van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 183. 

Notes. Still another species of the latreillei group, described 
from Top Cam]), 2100 m. (c. 6,825 ft.) in the Snow Mts., Neth. 
N. G. 

Cicindela rudolf-bennigseni Horn 

Horn 1912, Ent, Mitteilungen 1, p. 306. 

van Nidek 1959, Nova Guinea (new series) 10, p. 184. 

Notes. The type locality is Sattelberg, Huon Peninsula, N-E. 
N. G. ; van Nidek records the species from mountain slope 
above Bernhard Camp, 750 m. (c. 2,400 ft.), and Araucaria 
Camp, 800 m. (c. 2,600 ft.), Neth. N. G. 

Cicindela nigrivestis van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 184. 

Notes. The fourth new species of the latreillei group described 
by van Nidek in the paper cited. It is known from 4 specimens 
from mountain slope above Bernhard Camp, near Araucaria 
Camp, and Rattan Camp. Altitudes are 750-1200 m. (c. 2,400- 
3,900 ft.). 

Cicindela darlingtoni van Nidek 

van Nidek 1953, Psyche 60, p. 155, figs. 6, 7. 

Notes. The types are from Dobodura, Papua. 



344 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

(Group decemguttata) 

ClCINDELA DECEMGUTTATA URVILLEI Dejean 

Dejean 1831, Species Coleop. 5, p. 225 (dtirvillei) . 

Horn 1926, Coleop. Cat., Cicindelinae, p. 196 (see for additional references 

etc.). 
van Nidek 1953, Psyche 60, p. 156, figs. 4, 5. 
■ 1957, Treubia 24, pp. 1-2, map. 

1959, Nova Guinea (new series) 10, pp. 180, 184. 

Notes. According to Horn (loc. cit.), typical decemguttata 
occurs in Celebes, the Moluccas, Kei Is., etc. ; subspecies urvil- 
lei, in the Moluccas, New Guinea, Bismarck Archipelago etc. ; 
other subspecies, in the Solomons. Van Nidek (1957) discusses 
and maps the boundary between decemguttata scnsu stricto and 
subspecies urvillei in the Moluccas. The species does not reach 
Australia. It (represented by urvillei) is common and widely 
distributed in New Guinea, on the ground in open places. It 
is a rather large (c. 13-14 mm.) Cicindela, dark dull gray with, 
typically, 5 white marks on each elytron, but the marks vary 
somewhat (van Nidek 1953). 

(Group discreta) 
Cicindela discreta Schaum 

Schaum 1863, J. Ent. 2, p. 59. 

Horn 1926, Coleop. Cat., Cicindelinae, p. 196 (see for synonymy and addi- 
tional references), 
van Nidek 1953, Psyche 60, p. 156. 

1959, Nova Guinea (new series) 10, p. 184. 

Notes. C. discreta occurs in Sumatra, Java, Borneo, etc., the 
Philippines, Celebes, the Moluccas, New Guinea, New Britain, 
and North Queensland, Australia. It is a rather small species, 
found on the ground in open places. 

( Group semicincta-moseri) 
Cicindela semicincta Brulle 

Brulle 1834, in Silbermann, Rev. Ent. 2, p. 100. 

Horn 1926, Coleop. Cat., Cicindelinae, p. 197 (see for synonymy and addi- 
tional references). 

1932, Ree. S. Australian Mus. 4, p. 551. 

van Nidek 1953, Psyche 60, p. 158. 

Notes. Horn gives the range of semicincta as from New 
Guinea, etc. (including the Kei Is.) and the Bismarck Archi- 
pelago to the New Hebrides, Loyalty Is., New Caledonia. 



DARLINGTON : CARABID BEETLES OP NEW GUINEA 345 

and much of northern and eastern Australia. It is another 
rather small species which occurs on the ground in open places. 
The 2 other species of the group occur on Timor and the Tanim- 
bar Is. 

(Group placida-funerata) 
(Subgroup A) 

ClCINDELA EXCTSILABRIS Horn 

Horn 1905, Deutsche Ent. Zeits. 1905, p. 160. 

Notes. The type was from "Neu Guinea" without more exact 
locality. 

Cicindela pupilligera Chaudoir 

Chaudoir 186.1, Cat. Coll. Cicindeletes, p. 59. 
Horn 1904, Deutsche Ent. Zeits. 1904, p. 428. 

Notes. Chaudoir 's specimen was "Envoyee par M. Wallace, 
comme trouvee a Nouvelle-Guinee, ' ' and presumably came from 
Dorey, Neth. N. G. 

Cicindela io Horn 

Horn 1900, Deutsche Ent. Zeits. 1900, p. 203. 

1904, Deutsche Ent. Zeits. 1904, p. 428. 

—1915, in Wytsman, Genera Insectormn, Ease. 82, p. 317, pi. 18, fig. 8. 



van Nidek 1953, Psyche 60, p. 158. 

Notes. The types were from Milne Bay, Papua. I found the 
species at the same locality in December, 1943 (van Nidek det). 

Cicindela io micro-gem mea Horn 

Horn 1932, Eec. S. Australian Mus. 4, p. 551, fig. 2, a, b. 
van Nidek 1953, Psyche 60, p. 158. 

Notes. Mt. Lamington, Papua, is the type locality of this 
form. I found it at Dobodura, Mar.-July, 1944 (van Nidek det.). 

Cicindela delicata Bates 

Bates 1874, Ent. Month. Mag. 10, p. 265. 
Horn 1904, Deutsche Ent. Zeits. 1904, p. 428. 

1913, Arch. f. Naturg. 79, Abt, A, Heft 11, p. 31 (as innocens angus- 

tiformis). 

1926, Coleop. Cat., Cicindelinae, p. 198. 



346 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

Notes. Bates described delicata from "New Guinea (Wal- 
lace)," which presumably means Dorey, Neth. N. G. Horn's 
"innocens angustiformis," from Roon Is. (Geelvink Bay), 
Neth. N. G., is placed as a form of delicata in the Coleopterorum 
Catalogus. I do not know whether it is a recognizable subspecies. 

Cicindela placida Schaum 

Schaum 1863, J. Ent. 2, p. 60. 

Horn 1904, Deutsche Ent. Zeits. 1904, p. 428. 

■ 1915, in Wytsman, Genera Insectorum, Fase. 82, pi. 21, fig. 252. 

1926, Coleop. Cat., Cieindelinae, p. 198. 

Notes. Schaum 's type(s) came from "Mysol (D. Wallace)' 
in the Moluccas, and Horn (1926) lists the species also from 
New Guinea. 

Cicindela innocens Horn 

Horn 1893, Deutsche Ent, Zeits. 1893, p. 199. 

— 1913, Nova Guinea 9, Zool. 3, p. 410. 

Notes. The type was a 9 from "Nova Guinea," but Horn 
(1913) later recorded a 6 from " Heuvel-Biwak (Lorentz: XI 
1909, 750 m.)," which is apparently in the region of the upper 
Lorentz R., Neth. N. G. 

(Group placida-funerata, cont'd) 

(Subgroup B) 
Cicindela pupillata Schaum 

Schaum 1863, J. Ent, 2, p. 60. 

Horn 1904, Deutsche Ent. Zeits. 1904, p. 428. 

1926, Coleop. Cat., Cieindelinae, p. 198. 

Notes. This species too (like placida) was described from 
"Mysol (D. Wallace)'' in the Moluccas but is listed also from 
New Guinea by Horn (1926). 

Cicindela aruana Dokhtouroff 

Doklitouroff 1887, Ann. Soc. Ent. Belgique 31, p. 155. 
Horn 1904, Deutsche Ent, Zeits. 1904, p. 428. 

Notes. So far as I know, this species has been recorded only 
from the Aru Is., which belong with New Guinea, faunistically. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 347 

ClCINDELA LORIAE Horn 

Horn 1897, Ann. Mua. Civ. Genova (Genoa) (2) 17, p. 272. 

1904, Deutsche Ent. Zeits. 1904, p. 428. 

1926, Coleop. Cat., Cicindelinae, p. 198 (loriai). 

Notes. C. loriae is, I think, known only from the type locality, 
"Panmomn Riv. (Nonvelle Guinee britannique [= Papua])." 

ClCINDELA INNOCENTIOR Hom 

Horn 1904, Deutsche Ent. Zeits. 1904, p. 427. 

Notes. Sattelberg, on the Iluon Peninsula, N-E. N. G., is the 
type locality. 

ClCINDELA PSEUDO-PUPILLATA Horn 

Horn 1938, Ent. Beihefte Berlin-Dalilem 5, p. 12, pi. 59, fig. 2(5. 

Notes. Horn's single specimen was from 900 m. (e. 2,925 ft.) 
altitude in the Torricelli Mts., N-E. N. G. 

ClCINDELA DENUDATA Horn 

Horn 1935, Nova Guinea 17, p. 301. 

Notes. Kokoda, Papua, 1,200 ft. altitude, is the type locality. 

ClCINDELA CHEESMANAE Vail Nidek 

van Nidek 1954, Ann. Mag. Nat. Hist. (12) 7, p. 391, fig. 1 and pi. 9 
(chcesmannae). 
Notes. The types are from Camp Nok, Waigeu Is., Neth. N. G., 
2,500 ft. altitude. 

Cicindela klynstrai van Nidek 

van Nidek 1954, Ann. Mag. Nat. Hist. (12) 7, p. 393, fig. 2 and pi. 9. 

Notes. The types are from Japen ("Japan") Is., 500 ft. 
altitude, Neth. N. G. 

(Group placida-funerata, cont'd) 
(Subgroup C) 

ClCINDELA BENNIGSENIA Horn 

Horn 1901, Deutsche Ent. Zeits. 1901, p. 357. 

1904, Deutsche Ent, Zeits. 1904, p. 428. 

1932, Bee, S. Australian Mus. 4, p. 551, fig. 1, a, b. 

van Nidek 1953, Psyche 60, p. 158. 



348 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

Notes. Known localities for bennigsenia are Hercules R. 
( ?Hercules Bay, eastern N-E. N. G.) (type locality) and Mt. 
Lamington and Dobodura, Papua. 

ClCINDELA FUNERATA Boisduval 

Boisduval 1835, "Voyage Astrolabe, Faune Ent. 2, p. 4, pi. 6, fig. 1. 
Horn 1904, Deutsche Ent. Zeits. 1904, p. 428. 

1926, Coleop. Cat., Cicindelinae, p. 199 (see for synonymy and addi- 
tional references). 

1932, Eec. S. Australian Mus. 4, p. 551. 

van Nidek 1953, Psyche 60, p. 159 (funerata barbata). 

1959, Nova Guinea (new series) 10, pp. 184 (funerata), 185 (bar- 
bata). 

Notes. According to Horn (1926), this species occurs on 
New Guinea and adjacent islands and west to Batjan and Burn 
in the Moluccas, and subspecies barbata Horn occurs on the 
Bismarck Archipelago and perhaps the Solomons ; but van 
Nidek assigns specimens from eastern New Guinea to barbata. 
I do not know whether barbata is really a recognizable, geo- 
graphic subspecies. I found the species common at Dobodura 
and took it also at Milne Bay, Papua, and near Nadzab, N-E. 
N. G., and there is a long series in the M.C.Z. from Surprise 
Creek on the Morobe Plateau (also N-E. N. G.) (Stevens). 

Van Nidek (1959) now considers that funerata and barbata 
are distinct species. He may be right, but I cannot now un- 
scramble the old citations and records, so I shall let the two 
forms stand as one species for the time being. 

(Group placida-funerata, cont'd) 

(Subgroup doubtful) 
Cicindela toxopeusi van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 185. 

Notes. According to van Nidek, this should be a rather iso- 
lated species of the placida-funerata group, characterized by a 
pattern of elytral spots different from all other species of the 
group. It is described from 22 specimens from Bernhard Camp, 
the mountain slope above it, Araucaria Camp, and Rattan 
Camp, Neth. N. G. Altitudes are from 50 to 1200 m. (c. 150 
to 3,900 ft.). 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 349 

Cicindela olthofi van Nidek 

van Nidek 1959, Nova Guinea (new series) 10, p. 186. 

Notes. Kelated to the preceding species, and described from 
5 specimens from Bernhard Camp and the mountain slope above 
it, Neth. N. G., 50-750 m. (c. 150-2,400 ft.). 

(Group variolosa) 
Cicindela variolosa Blanchard 

Blanchard 1853, Voyage au Pole Sud — L 'Astrolabe et La Zelee, Zool. 4, 

p. 6, Atlas Ins. pi. 1, fig. 4. 
Horn 1926, Coleop. Cat., Cicindelinae, p. 199. 

Notes. This species was described as from the south coast 
of New Guinea. Horn questions its occurrence in New Guinea 
(I do not know why) and records it from Batjan in the Moluc- 
cas. 

(Group boisduvali) 
Cicindela boisduvali Horn 

Horn 1896, Deutsche Ent. Zeits. 1896, p. 152. 

1904, Deutsche Ent. Zeits. 1904, p. 428. 

1913, Nova Guinea 9, Zool. 3, p. 410. 

van Nidek 1959, Nova Guinea (new series) 10, p. 186. 

Notes. The types were from Humboldt Bay; Horn later 
(1913) recorded specimens from Alkmaar, on the upper Lorentz 
R., and van Nidek, from Hollandia and Bernhard Camp at low 
altitudes ; all these localities are in Neth. N. G. 

Cicindela kampeni Horn 

Horn 1913, Tijd. v. Ent. 56, p. 310. 

Notes. Described from 1 S from Hollandia, Neth. N. G. 

(Group guineensis) 
Cicindela guineensis Horn 

Horn 1892, Deutsche Ent. Zeits. 1892, p. 77. 

1904, Deutsche Ent. Zeits. 1904, p. 428. 

1932, Eec. S. Australian Mus. 4, p. 252, fig. 4. 

van Nidek 1959, Nova Guinea (new series) 10, p. 186. 

Notes. Horn's type came from " Neu-Guinea. " I do not think 
typical guineensis was recorded from a more exact locality until 
van Nidek reported 3 from Bernhard Camp, Neth. N. G. 



350 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

ClCINDELA GUINEENSIS UMBROSA Horn 

Horn 1932, Kee. S. Australian Mus. 4, p. 553, fig. 3, a, b. 
van Nidek 1953, Psyche 60, p. 159. 

Notes. The types of umbrosa were from Mt. Lamington, 
Papua, and I found it at Dobodura (van Nidek det.). 

( Group ancorif era-parry i-tuberculata) 

ClCINDELA ANCORIFERA Horn 

Horn 1897, Ann. Mus. Civ. Genova (Genoa) (2) 17, p. 271. 

— 1926, Coleop. Cat,, Cicindelinae, p. 199. 
van Nidek 1953, Psyche 60, p. 159. 

1959, Nova Guinea (new series) 10, p. 186. 

Notes. The type locality is Hatam, in the Arfak Mts. of 
Western Neth. N. G. Van Nidek (1953) records specimens from 
the Bismarck Range at 5,000-7,500 ft, altitude (taken by me) 
and Mt. Misim at 6,400 ft. in the Morobe District (Stevens), 
and Prof. E. O. Wilson took specimens at Tumnang, 1,400-1,600 
m. (c. 4,300-4,900 ft,), and between Nganduo and Yunzain, 
1,000-1,500 m. (c. 3100-4,625 ft.) on the Mongi watershed in the 
mountains of the Huon Peninsula ; all these localities are in 
N-E. N. G. Van Nidek later (1959) notes specimens from Rat- 
tan, Sigi, Lower Mist, and Mist Camps in the Snow Mts., Neth. 
N. G., at altitudes of 1200-1800 m. (c. 3,900-5,825 ft.). The 
species is apparently widely distributed in New Guinea at mid- 
dle altitudes. The great interest of ancorif era is in its appar- 
ent geographical relationships : all the other species of its 
group are confined to New Zealand ! Further study is needed 
to show whether this is a real relationship or a result of con- 
vergence. C. ancorif era lives on the ground in open places. 

(Group maino) 
Cicindela maino Macleay 

Macleay 1876, Proc. Linn. Soc. New South Wales 1, p. 165. 
Horn 1915, in Wytsman, Genera Inseetoruni, Fasc. 82, p. 320, pi. 19, fig. 1. 
Notes. Macleay 's specimens were collected on the sea beach 
at the mouth of the Katow (Binaturi) R., and Horn lists the 
species also from Redscar Bay. Both localities are on the south 
coast of Papua. It is a striking, fusiform, very long-legged 
species with complex markings. Horn (op. cit., p. 312) says 
it is derived from an Oriental stock. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 351 

(Group denticoUis) 

ClCINDELA DENTICOLLIS Horn 

Horn 1895, Deutsche Ent. Zeits. 1895, p. 88. 
— 1915, in Wytsman, Genera Insectorum, Fasc. 82, p. 321. 
Notes. Although the types were from "Nov. -Guinea" without 
further locally, Horn later gives the species' range as north- 
western (Neth. ) N. G. and the Aru Is. This species too is de- 
rived from a (different) Oriental stock (Horn, op. cit., p. 
321). 

Subfamily CARABINAE 

Although this is an artificial or composite subfamily, it is 
recognized in the Cole opt erorum Catalogns, and it is temporar- 
ily useful as a device for arranging the elements of a complex 
family of which the phylogeny is not yet understood. 

Tribe OZAENLNI 

Ozaenini Auct., including Banninger 1927, Deutsche Ent. Zeits. 1927, p. 177. 
Ozaenidae Auct., including Jeannel 1946, Coleop. Carabiques de la Region 
Malgache, Part 1, p. 46. 
Notes. This apparently primitive tribe is almost pantropical 
in distribution. A single genus and species of it reach New 
Guinea, from the Orient. 

Genus PsEUDOZAENA Castelnau 

Castelnau 1834, Etude Ent. 1, p. 55. 

Banninger 1927, Deutsche Ent. Zeits. 1927, p. 192. 

Jeannel 1946, Coleop. Carabiques de la Region Malgache, Part 1, p. 48. 

Diagnosis. None required here. This is the only ozaenine 
genus in New Guinea. 

Description. See references given above. 

Genotype. P. megacephala Castelnau = Ozaena orientalis 
Klug. 

Generic distribution. As limited by Banninger, the genus 
is confined to the Malay Peninsula, Formosa, and the Indo- 
Australian Archipelago east to the Philippines, New Guinea, 
the Admiralties, the Solomons, and (introduced?) the Palau 
Is. In the broader sense of Jeannel, it occurs also in eastern 
tropical Asia, Africa, and Madagascar. 



352 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

PSEUDOZAENA ORIENTALIS OPACA (Chaildoir) 

Picrus opacus Chaudoir 1868, Ann. Soc. Ent. Belgique 11, p. 46. 
Pseudozaena opaca Andrewes 1924, Ann. Mag. Nat. Hist. (9) 14, p. 585. 
Pseudozaena tenebrosa Sloane 1890, Kec. Australian Mus. 1, p. 102. 
Pseudozaena tricostata tenebrosa Banninger 1927, Deutsche Ent. Zeits. 

1927, p. 192. 
Pseudosaena tricostata opaca Gressitt 1953, Bull. Bishop Mus. No. 212, 
p. 95, fig. 45a. 

Description. None required here, except to note that the 
species is winged. See figure 1. 

Types. Of opacus Chaudoir, apparently from Ceram and 
Ternate in the Moluccas (Andrewes 1924), now presumably in 
the Oberthiir Collection at the Paris Mus. ; of tenebrosa Sloane, 
from British New Guinea (Papua), now probably in the Sloane 
collection at Canberra, Australia. 

Occurrence in New Guinea. Papua : Dobodura, Mar. -July 
1944 (Darlington) 1 ; Kokoda, 1,200 ft., (Cheesman) 1 ; Port 
Moresby area (L. Jones, British Mus.) ; Brown R., May 22, 
1956 (E. J. Ford, Jr., Bishop Mus.) ; Daru, mouth of Fly 
R., July 1941 (R. G. Wind, California Acad.) ; Kiunga, 
Fly R.,' July 23-25, 1958 (W. W. Brandt, Bishop Mus.) ; 
Kikori, Feb. 27, 1920 (J. T. Zimmer, Chicago Mus.). N-E. 
N. G. : Aitape, Aug. 1944 (Darlington) ; Stephansort, Astro- 
labe Bay (Biro, Hungarian National Mus.) ; Torricelli Mts., 
Mokai Village, 750 m. (c. 2450 ft.), Dec. 8-15, 1958 (W. W. 
Brandt, Bishop Mus.). Neth. N. G. : Humboldt Bay region, 
including Hollandia and Cyclops and Bewani Mts. up to 
1,200 ft. (various sources) ; mountain slope above Bernbard 
Camp, 100 m. (about 325 ft.), April 1939 (Toxopeus) ; Arau- 
caria Camp, Snow Mts., 800 m. (about 2,600 ft.), Mar. 1929 
(Toxopeus) ; Upper Setekwa R., Snow Mts. (t. Banninger) ; 
route of the Kaiserin-Augustafluss Expedition (t. Banninger) ; 
Geelvink Bay, 1878 (Raffray & Maindron, Paris Mus.) ; Arfak 
Mts. (t. Banninger) ; Wasian, Sept. 1939 (R. G. Wind, M.C.Z.). 
Several additional specimens seen without exact localities or 
from localities I have not been able to find. The records suggest 
that this insect occurs throughout New Guinea from sea level 
into the foothills of the mountains. As to its habits, I have 
taken it in flood debris and have seen specimens collected "at 



i For disposition of material and for abbreviations used see present part of 
this work, p. 324, and part 2 (1952), pp. 90-91. 



DARLINGTON : CARABID BEETLES OP NEW GUINEA 353 

light" and (from the Philippines) "in sawdust at mill." Gres- 
sitt (loc. tit.) says that, in the Palau Is., it is "a moderately 
common predaceous beetle in logs and trunks where Oryctes is 
found. . . . One adult kept in the laboratory fed for five weeks 
on Oryctes eggs alone, and another lived for three weeks on eggs 
and young larvae only." 

Notes. After examination of new material, including 39 speci- 
mens from New Guinea originally assembled for study at the 
M.C.Z. (more seen later), and after comparison of the $ 
copulatory organs, I am prepared to go even further than Ban- 
ninger (1927) in reducing the number of forms in this genus, 
and to recognize in Pseudozaena sensn stricto only one species 
with three geographical subspecies: typical orient alis (Klug) 
of the Malay Peninsula, Sumatra, Java, and Borneo ; sub- 
species opaca (Chaudoir) of Formosa, the Philippines, the 
Moluccas, New Guinea, and the Palau Is. (if native there) ; 
and subspecies tricostato Montrousier of New Britain and the 
Solomons. A specimen from the Kei Is. (H. C. Siebers, British 
Mus.) has stood in the Andrewes Collection under orientalis, 
but is actually opaca. It is probably the basis of Andrewes' Kei 
Is. record of orientalis; the Kei Is. should be deleted from the 
range of this form. Specimens from both New Guinea and the 
Philippines vary so much in sculpture of elytra that I cannot 
distinguish the New Guinean population as a separate sub- 
species. A specimen from Waigeu Is., west of New Guinea 
(Cheesman), and 3 from the Admiralty Is. (U.S.N.M., Bishop 
Mus.) are like tricostato, but should not be referred to that 
subspecies without examination of additional material. These 
specimens from coastal islands may be relicts of a tricostate 
population that may formerly have occurred on New Guinea 
but that (if it occurred) has changed or been replaced. 

Tribe PAUSSINI 

Paussidae Auct. (in part). 

Paussini Darlington 1950, Trans. Amer. Ent. Soe. (Philadelphia), 76, p. 90. 

Although often treated as a separate family, the paussids 
are derived from the Ozaenini and are best placed after them in 
the Carabidae (Darlington op. tit.). 

Only two species of paussids have been recorded from New 
Guinea, one representing an Australian genus the other an 
Oriental one, but the occurrence of the latter in New Guinea is 
doubtful. 



354 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Key to Genera of Paussini Recorded from New Guinea 

1. Antennae with 10 free segments, the 9 flagellar ones expanded and 

flattened Arthropterus 

— Antennae appearing 2-segmented, the flagellum fused, with 2 conspicu- 
ous processes posteriorly Euplatyrhopahis 



Genus ARTHROPTERUS Macleay 

Macleay 1838, in A. Smith, Illustrations Zool. S. Africa, Invertebratae, 

p. 75. 
Gestro 1910, Junk-Sehenkling Coleop. Cat., Paussidae, p. 7 (see for addi- 
tional references etc.). 
Darlington 1950, op. tit., pp. 94, 95, 106. 

Diagnosis. See Darlington, op. cit. 

Description. None needed here. 

Genotype. Cerapterus macleayii Donovan, of Australia. 

Generic distribution. Living only in Australia (many species) 
and New Guinea (1 species) ; supposedly fossil in the Baltic 
amber in Europe (several species). 

Arthropterus novellus Kolbe 

Kolbe 1924, Ent. Mitteilungen 13, p. 72. 

Description. None needed here. 

Types. Two specimens labeled only "New Guinea" (Stau- 
dinger), in Stettin and Berlin Museums. 

Occurrence in New Guinea. Known only from the types. 

Notes. The state of the wings in this species is unknown. In 
Australia, the wings are fully developed in some Arthropterus, 
vestigial in others. 

Genus EUPLATYRHOPALUS Desneux 

Desneux 1905, in Wytsman's Genera Insectorum, 35me fasc, Paussidae, 

p. 18. 
Darlington 1950, Trans. American Ent. Soc. (Philadelphia) 76, pp. 98, 107. 

Diagnosis. See Darlington, op. cit. 

Description. None needed here. 

Genotype. Platyrhopcdus aplustrifer Westwood, of India. 

Generic distribution. India and Burma to Sumatra and Java ; 
and perhaps New Guinea. (No paussid of any sort is yet known 
from Celebes or the Moluccas.) 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 355 

Euplatyrhopalus wasmanni van Emden 

van Emden 1927, Ent. Blatter 23, p. 127. 

Description. None required here. The species is probably 
fully winged. 

Type. From New Guinea? According to van Emden, it may 
really be from Borneo; type should be in Dresden Mus. 

Occurrence in New Guinea. Known only from the type -- if it 
came from New Guinea. 

Tribe SCAEITINI 

Scaritini Auct., including Andrewes 1929, Fauna British India etc., Coleop., 

Carabidae 1, p. 208. 
Scaritidae Auct., including Jeanne! 1946, Coleop. Carabiques de la Region 
Malgache, Part 1, p. 212. 
New Guinea has relatively few genera of this tribe, only 3, 
against more than 15 in the Orient and about 20 in Australia. 
The dominant, nearly cosmopolitan genus, Scarites, which in- 
cludes many Oriental species, extends east to Celebes and Timor 
but does not reach New Guinea or Australia. And the dominant 
Australian "carenums, " with about a dozen genera and hun- 
dreds of species in Australia, are still unknown in New Guinea, 
although one or two species of them may yet turn up in south- 
ern New Guinea, in the extensive, open Eucalyptus country 
there. 

Key to Genera of Scaritini Known from New Guinea 

1. First segment of antenna received in deep groove under eye; antenna 

with first 4 segments glabrous (except for fixed tactile setae) ; 1 

supraocular seta over each eye (subtribe Scaritina) Geoscaptus 

— First segment of antenna not received in deep groove; antenna with first 
2 segments glabrous; 2 supraocular setae ever each eye (subtribe 
Clivinina) 2 

2. Elytra strongly dentate at humeri; antennal segment 2 attached very 

eceentricly to segment 1; length (in New Guinea) c. 3% mm 

Syleter 

— ■ Elytra not dentate at humeri; first 2 segments of antenna normal; 

length (in New Guinea) c. 4—18 mm Clivina 

Genus (rEOSCAPTUS Chaudoir 

Cliaudoir 1855, Bull. Soc. Nat. Moscou 28, Part 1, No. 1, p. 5. 
Sloane 1905, Proc. Linn. Soc. New South Wales 30 pp. 103-108. 



356 BULLETIN : MUSEUM OF COMPARATIVE, ZOOLOGY 

Banninger 1937, Deutsche Ent. Zeits. 1937, pp. 118, 133-137. (Selected 
references only) 

Diagnosis. Subeylindrical, shining black, Scarites-like cara- 
bids, with the maxillae broadly rounded apically, not curved 
in and not pointed or toothed as in Scarites etc. 

Description. None required here. 

Genotype. Geoscaptus laevissimus Chaudoir of Australia. 

Generic distribution. Eastern and northern Australia, New 
Guinea. 



Geoscaptus cacus (Macleay) 

Scarites cacus Macleay 1863, Trans. Ent. Soc. New South Wales 1, p. 67. 
Geoscaptus cacus Banninger 1937, Deutsche Ent. Zeits. 1937, pp. 135, 136. 
(Selected synonymy and references only.) 

Description. See generic description and figure 2. Within the 
genus, this species is notably convex (subeylindrical) and rela- 
tively small, 16-24 mm. (Banninger). Specimens from New 
Guinea are near the minimum size for the species, c. 16-20 mm. 

Type. From "Port Denison" (near Bowen, Queensland, Aus- 
tralia) ; should be in Macleay Mus., Sydney. 

Occurrence in New Guinea. Papua: 1, Port Moresby area, 
May 1947 (L. Jones, British Mus.) ; 2, Dobodura, Mar.- July 
1944 (Darlington). N-E. N. G.: 1, vie. Nabzab, July 1944 
(Darlington). Neth. N. G.: 1, Hollandia, Apr. 1945 (Malkin, 
U.S.N.M.). The habitat in New Guinea is not recorded, but 
in Australia the genus occurs (by day) under cover near water. 

Notes. Widely distributed also in northern and eastern Au- 
stralia. The species (like others of the genus) is winged and 
probably a good flier. 



Genus SYLETER An dr ewes 

Andrewes 1941, Ann. Mag. Nat. Hist. (11) 7, p. 317. 
Psilus Putzeys 1877, Ann. Soc. Ent. Belgique 20, Compt. Kend., p. 46. 
Andrewes 1929, Fauna British India etc., Coleop., Carabidae 1, pp. 344, 
386-389. 

Diagnosis. See preceding references and key to genera. 

Description. See Andrewes 1929, pp. 386-387. 

Genotype. Ardistomis paradoxa Putzeys of Siam, etc. 

Generic distribution. Burma and Indochina to the Philip- 
pines, New Guinea, and the tip of Cape York, Australia ; Africa. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 357 

SYLETER PAPUA 11. Sp. 

Description. Form (fig. 3) of Ardistoniis (rather like stout 
Dyschirius) ; reddish piceous, elytra darker, appendages dark 
reddish ; surface moderately shining, reticulate microsculpture 
approximately isodiametric on front of head, pronotum, and 
striae of elytra, absent on elytral intervals, which polished and 
shining. Head .59 and .59 width prothorax; eyes prominent (nor- 
mal), genae short and oblique; antennae normal for genus, outer 
segments slightly longer than wide; labrum 7-setose (single setae 
sometimes missing) ; clypeus truncate with angles rounded, cly- 
peal suture obliterated ; frontal sulci deep, subparallel, but ir- 
regular and slightly curved outward in front and behind ; a 
strong ridge along outer edge of each sulcus above eye, separated 
from eye by a channel ; front otherwise almost evenly convex 
except often with a faint longitudinal impression at middle, im- 
punctate ; neck impressed only at sides, with a row of punctures 
on each side but widely interrupted at middle. Prothorax broad, 
rounded; width/length (including peduncle) 1.05 and 1.07; 
sides broadly rounded anteriorly, more strongly posteriorly; 
anterior angles blunt, only slightly prominent, posterior ones 
broadly rounded but marked by faint teeth ; apex broadly emar- 
ginate, truncate at middle ; lateral margins entire, each with 
usual 2 setae, at basal angle and about y 3 from apex ; disc 
convex, with deep transverse impression anteriorly and shallower 
longitudinal median line, and with a small cluster of vague 
punctures on each side a little behind middle, but otherwise im- 
punctate. Elytra 1.18 and 1.19 width prothorax; base emargin- 
ate between ends of 4th striae, with tubercles at anterior ends of 
2nd and 3rd intervals ; base strongly margined on each side from 
4th striae to humeri ; latter strongly toothed, and margins be- 
hind them crenulate; sides slightly diverging behind humeri, 
then broadly rounded to apices ; disc strongly convex ; striae 
deep and entire, reticulate rather than punctate ; intervals mod- 
erately convex, impunctate, except 3rd interval 3-punctate, with 
anterior puncture near 3rd striae and other two near 2nd one. 
Inner wings fully developed. Lower surface microreticulate and 
in part roughened but not distinctly punctate. Legs: front 
tibia with a long, curved apical process and 2 strong teeth ex- 
ternally; middle tibia without spur. Measurements: length (in 
normal position) c. 3.3-3.8; width c. 1.1-1.3 mm. 

Types. Holotype 6 (M.C.Z. No. 30,152) and 22 paratypes 



358 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

from Dobodura, Papua, Mar. -July 1944 (Darlington). Addi- 
tional paratypes as follows: Papua: 13, Oro Bay, Dec. 
1943-Jan. 1944 (Darlington). Neth. N. G. : 10, Hollandia, July- 
Sept. 1944 (Darlington) ; 52, Maffin Bay, Aug. 1944 (Darling- 
ton) and 12, same locality, Aug. 1944 (E. S. Ross, California 
Acad.). My specimens were all taken in very wet places in and 
around shaded swamps. 

Measured specimens. The S holotype and 1 9 paratype from 
Dobodura. 

Notes. This species occurs also in northern Cape York, Au- 
stralia. It is similar to and probably a representative of Syleter 
paradoxus (Pntzeys) of southeastern Asia, Sumatra, and Bor- 
neo, but as compared with paradoxus in the Andrewes Collection 
the present new species is a little broader, with relatively broader 
prothorax, and with the line of punctures across the neck more 
widely interrupted at middle. 

Genus ClJVINA Latreille 

Latreille 1802, Hist. Nat. Crustaces et Inseetes 3, p. 96. 
Sloane 1896, Proc. Linn. Soc. New South Wales 21, pp. 143-257, 275-280 
(the Australian species). 

• 1904, Proc, Linn. Soc. New South Wales 29, pp. 710-733 (Australian 

species). 
Andrewes 1929, Fauna British India etc., Coleop., Carabidae 1, pp. 344, 

351-381. 
Kult 1947, Acta Soc. Ent. Czechoslovakia 44, p. 32 (subgenera). 

1951, Acta Soc. Ent. Czechoslovakia 48, pp. 16-32 (the Oriental 

species). 
(Selected references only) 

Diagnosis and description. See preceding references and key 
to genera. 

Genotype. Tenebrio fossor Linnaeus, of Europe. 

Generic distribution. Almost cosmopolitan; many species on 
all continents, but few or none in cold places where (in the 
north) the genus tends to be replaced by Dyschirius. 

Notes. Both Oriental and Australian groups of this genus 
are represented in New Guinea, but the exact relationships of 
some of the species are doubtful. 

I have made a special comparison of the Clivina of Cape 
York (where I collected long series in 1958) with those of 
New Guinea. A few species (zebi, basalis, sellata, ferruginea, 
inopaca) in the two places seem to be the same, but most are 
different. Some of the species of this genus, especially those in 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 359 

the last few couplets of the following- key, are difficult to separate 
from each other and may now be in process of evolutionary radi- 
ation. 

Most Clivina live in the ground in wet places, but each prefers 
a special niche within this general habitat (damp soil in rain 
forest, or mud by standing water, or sandy river banks, etc.). In 
Australia some species of the genus have entered deserts, and in 
the Philippines one occurs in decaying logs. Most species are 
winged and many fly to light, but the wings of a few have 
atrophied. Wing atrophy has in fact occurred in three separate 
stocks of the genus in New Guinea (see toxopei, dealata, and 
crugatella). 

Species of Clivina previously described from New Guinea but 
not recognized from description 

Clivina guineensis Kult 

Kult 1951, Acta Soc. Ent. Czechoslovakia 48, pp. 29, 30. 

Type. From Astrolabe Bav, N-E. N. G. (from Staudinger, in 
Kult Coll.). 

Notes. A rather small red species which may (or may not) be 
similar to rufulus (below) ; the brief description suggests sig- 
nificant differences. 

Clivina schaubergeri Kult 

Kult 1951, Acta Soc. Ent. Czechoslovakia 48, pp. 29, 30. 

Type. From "Fly River," Mew Guinea (Kult Coll.). 

Notes. A black, shining, 8.5 mm. member of the ephippiata 
group characterized otherwise only by "head without neck con- 
striction ; intervals moderately convex, striae finely punctate, 
third stria with 4 little distinct pores" and especially by "anten- 
na! joints 1.5X longer than wide." I do not think this species 
is represented in the material before me. 

These 2 species are not included in the following key. 

Key to Known Species of Clivina of New Guinea 

1. Anal (last ventral) segment with the 2 seta-bearing punctures on each 

side close together (fig. 18) 2 

— Anal segment with the 2 seta-bearing punctures on each side widely 
separated (fig. 19) 3 

2. Middle tibia with very short spur (fig. 20) ( tranquebarica group) 

(p. 362) zebi 



360 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

— ■ Middle tibia with longer spur; (anterior puncture of 3rd elytral in- 
terval displaced, near 2nd stria) (p. 362) . . (castanea) 

3. Very large (c. 14-18 mm.) ; (whole front coarsely, irregularly wrin- 

kled) (p. 363) toxopei 

— Smaller (4-10 mm.) 4 

4. Cylindrical, prothorax very elongate (L/W 1.23) ; legs and antennae 

very short and stout, intermediate antennal segments transverse 
(p. 364) brevicornis 

— Form only normally convex, or depressed -5 

5. Labrum 6-setose (median seta lacking) 6 

— Labrum 5- or 7-setose (median seta present) 7 

6. Third elytral interval 4-punctate; (rather depressed; piceous or red- 

dish; c. 5-6 mm.) (p. 365) (wallacei) 

— Third elytral interval 3-punctate; (black, appendages ferruginous; 5.3 

mm.) (Icomdrcki Kult, if labrum 6-setose; see also couple 9) 

TcomdreM 

7. Labrum 5-setose 8 

— Labrum 7-setose 10 

8. Large (c. 8.5-10 mm.); 3rd elytral interval 4-punctate (p. 366) kulti 

— Smaller (c. 6 mm.) ; 3rd elytral interval 3-punctate 9 

9. Frontal (preocular) plates with outer margins normal, oblique an- 

teriorly (fig. 10) ; neck constriction shallow or interrupted at middle; 
anterior transverse groove of pronotum normal (p. 368) birol 

— Frontal plates broader, rounded (fig. 11); neck constriction entire, 

deeper, more sharply defined ; anterior transverse impression of pro- 
notum very deep (p. 369) TcomdreJci 

10. Clypeus with wings separated from median part by at least slight 

notches (figs. 13, 14) ; elytron usually with 4 striae free at base, 

except in dedlata 11 

Clypeus with wings not separated from median part by notches (figs. 
15-17) (but wings in some cases advanced and forming obtuse angles 
with median part) ; (elytron usually with 3 striae free at base, but 
4th stria sometimes free in rufula, tripuncta, erugata, subfusa, etc.) 

14 

11. Spur of middle tibia (fig. 24) very long, longer than tibial width; 

(anterior trochanter with a small acute tooth at apex, on lower edge 
of leg; front with rather coarse, scattered punctures at least an- 
teriorly) 12 

— Spur of middle tibia not longer than tibial width 13 

12. Fully winged; elytron usually with 4 striae free at base; larger (5.8- 

6.5 mm.) ; usually black (p. 370) puncticeps 

— ■ Inner wings full or vestigial; elytron with 3 or 4 striae free at base; 
smaller (4.5-5.4 mm.) ; brown or partly brown dedlata 

(a) Fully winged (Fly River I (p. 373) (subsp. antecessor) 

(b) Wings vestigial, vestiges about % length of elytra; (Papua) 
(p. 372) (dedlata scnsu strict o ) 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 361 

(c) Wings vestigial, vestiges shorter, scarcely reaching beyond pos 
terior edge of metathorax; (Neth. N. G., N-E. N. G.) (p. 373) 

(subsp. bradhyptera) 

13. Eyes large, genae very short (fig. 14) ; anterior transverse groove of 

pronotum normally impressed (p. 374) vigil 

— Eyes smaller; genae (measured obliquely from sides of neck to pos- 

terior edges of eyes) nearly as long as eyes; anterior pronotal groove 
obsolete (its place taken by an irregular line of dark pigment under 
the surface of the pronotum) (p. 375; deleta 

14. Third intervals of elytra 3-punctate 15 

— Third intervals 4-punctate (australasiae-ephippiata group) 16 

15. Rather depressed; rufous; front not impressed and not punctate 

(p. 377) rufula 

— Less depressed; usually blackish with elytral margin ± pale; front 

vaguely impressed, usually punctate (p. 378) tripuncta 

16. Supraocular convexities (frontal earinae) smoothly continuous with 

swollen preocular plates (but if form is parallel-sided and ratio 
width elytra/prothorax less than 1.10 and abdomen conspicuously 
punctate, see gressitti, couplet 19) 17 

— Supraocular convexities ± interrupted (impressed or abruptly nar- 

rowed) near anterior supraocular setae 18 

17. Smaller (c. 5-6 mm.), broader, slightly flatter (p. 380) erugatella 

— Larger (c. 6-8 mm.), narrower, slightly more convex (p. 382) . erugata 

18. Spur of middle tibia (figs. 32, 36) short, very near apex of tibia. . . .19 
Spur of middle tibia always longer and usually not so near tibial 

apex 20 

19. Subfusif orm ; ratio width elytra/prothorax c. 1.25; length over 6.5 

mm. (p. 384) . subfusa 

— Very parallel sided; width elytra/prothorax less than 1.10; length 

under 6 mm. (p. 385; gressitti 

20. Bicolored, black with anterior part of elytra red (p. 383) basalts 

— Xot thus bicolored 21 

21. Very small (4 mm.) ; (brown) (p. 387) sellata 

— ■ Larger 22 

22. Anterior trochanter with a small acute tooth at apex, on lower side of 

leg, and color brown, and length under 6 mm. (p. 387) . ferruginea 

Anterior trochanter usually not toothed, or if toothed, other characters 

not as above 23 

23. Front femur (fig. 38) less stout; spur of middle tibia (fig. 33) near 

tibial apex (p. 388) fessa 

— Front femur stouter (fig. 39) ; spur of middle tibia (figs. 34, 35) not 

so near apex 24 

24. Last ventral segment punctate (p. 389) sansapor 

— Last ventral segment with reticulate mieroseulpture but usually not 

punctate .25 

25. Elytral disc not dull .26 

— Elytral disc dull (with reticulate mieroseulpture) 28 



362 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

26. More slender, prothorax considerably longer than wide (L/W 1.09) 

(p. 390) csikii 

— Less slender, prothorax about as long as wide (between L/W 1.04 & 

W/L 1.03) 27 

27. Middle of front conspicuously (but variably) punctate (p. 393) . 

brandti 

— Front not or only slightly punctate (p. 392) . inopaca 

28. Black; elytral striae more distinctly punctate; intervals more convex, 

3rd with the 4 dorsal punctures less distinct (p. 395) szekessyi 

— More brownish, antennae and legs more reddish ; elytral striae less dis- 

tinctly punctate; intervals less convex, 3rd with the 4 discal punc- 
tures more distinct (p. 396) netolitzlcyi 

Clivina zebi Kult 

Kult 1951, Acta Soc. Ent. Czechoslovakia 48, pp. 22, 24. 

Description (recognition characters only). This is the only 
member of the Oriental Clivina tranquebarica group known to 
reach New Guinea. It is distinguished from other New Guinean 
Clivina by characters given in the key. 

Type. The type is from Borneo, in Dr. Kult's collection. 

Occurrence in New Guinea. Papua: 13, Dobodura, Mar. -July 
1944 (Darlington). Neth. N. G. : 10, Hollandia, July-Sept. 1944 
(Darlington) ; 1, same locality, May 1945 (H. Hoogstraal, 
M.C.Z.) ; 2, Maffin Bay, Aug. 1944 (Darlington). The species 
occurs chiefly, I think, in shaded swamps. 

Notes. Clivina zebi ranges from Sumatra to the Philippines 
and New Guinea ; I have seen 4 specimens also from Cape 
Gloucester, New Britain ; and in 1958 I found it common at 
several localities on the Cape York Peninsula, Australia. I do 
not think it has received a name in Australia. 

(Clivina castanea Westwood) 

Westwood 1837, Proe, Zool. Soc. London 1837, p. 128. 

Andrewcs 1929, Fauna British India etc., Coleop., Carabidae 1, pp. 355, 374, 

fig. 54. 
Clivina parryi Putzeys 1863, Mem. Soc. R. Sci. Liege 18, p. 60. 
(Selected references and synonymy only) 

Description. A rather large, black Clivina, characterized in 
the key (above). 

Type. The type of castanea, from Manila in the Philippines, is 
now in British Mus. The types of parryi were supposed to be 
from New Guinea, but it is doubtful if they really came from 
there. They should be in Brussels Mus. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 363 

Occurrence in New Guinea. I know no recent records; the 
old ones are doubtful. 

Notes. Although this Oriental species is commonly said to 
extend to New Guinea, the supposed New Guinean specimens 
are old and of unstated history. They may have been collected 
by Wallace ; if so, their real locality is doubtful. Clivina cas- 
tanea is usually a very common species where it occurs. Its 
absence from all recent collections seen from New Guinea sug- 
gests that it does not really occur there. 

Clivina toxopei n. sp. 

Description. Form as figured (fig. 4) ; very large (in 
genus) ; black, appendages dark ; moderately shining, pronotum 
and elytra with fine reticulate microsculpture distinct in type 
but less distinct or partly absent in paratype. Head (fig. 8) 
.72 and .68 width prothorax (in type and paratype) ; eyes small 
but convex, enclosed behind by genae ; antennae short, median 
segments slightly wider than long, normally pubescent; man- 
dibles short and stout ; labrum 7-setose ; clypeus subtruncate at 
middle, wings continuous with median part but more advanced, 
separated from preocular lobes by notches ; facial carinae short 
and poorly defined ; frontal foveae very deep, irregular ; whole 
front of head coarsely and irregularly wrinkled, not punctate 
but sometimes vaguely punctulate ; neck constriction entire or 
nearly so, not punctate. Protliorax exactly as long as wide by 
standard measurement but appearing slightly longer (because 
anterior angles are more advanced than front of prothorax at 
middle), widest near basal angles, strongly narrowed anter- 
iorly ; apex broadly emarginate ; anterior angles very narrowly 
rounded, almost right; posterior angles obtuse-rounded, not dis- 
tinctly dentate; sides almost straight (slightly arcuate), finely 
margined, each with usual 2 setae ; disc with median line and 
anterior transverse impression entire, impressed, not punctate ; 
surface of disc variably wrinkled or strigulose, not punctate but 
inconspicuously punctulate, with an irregular longitudinal im- 
pressed line on each side distinct in type but vague in paratype. 
Elytra elongate-oval, slightly wider than prothorax (B/P 1.1 
in both specimens); base slightly emarginate; humeri slightly 
prominent anteriorly but broadly rounded into sides, not den- 
tate ; each elytron with 3 striae free at base, the 4th turning 
out and joining or almost joining outer striae at humerus; 



364 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

striae entire, deeply impressed, punctate in type, not in para- 
type; intervals convex, 3rd 4-punctate on outer side. Inner 
wings atrophied ; elytra locked together or connate. Lower sur- 
face : prosternal process wide and flat before coxae ; proepisterna 
shining, only partly and lightly microreticulate ; lateral cavities 
of peduncle reticulate and more or less wrinkled; metepisterna 
shortened, about % longer than wide (judged by eye — I do 
not want to set standards of measurement) ; abdomen with fine 
reticulate microsculpture, impunctate ; apical segment with 2 
seta-bearing punctures on each side widely separated. Legs nor- 
mal, not stouter than usual ; front tibia 3-dentate, the 4th 
(upper) tooth reduced to an inconspicuous angulation; middle 
tibia (fig. 21) with spur on outer side near apex about as long 
as width of tibia. Measurements (type and paratype) : length c. 
18 and 14.5: width c. 4.8 and 4.1 mm. 

Types. Holotype 9 (Leiden Mus.) from Sigi Camp, Snow 
Mountains, Neth. N. G., 1,500 m. (c. 4,650 ft.), Feb. 1939 (L. J. 
Toxopeus) ; and 1 9 paratype (M.C.Z. No. 30,153) from Arau- 
caria Camp, also in the Snow Mountains, 800 m. (c. 2,480 ft.), 
March 1939 (Toxopeus). 

Measured specimens. The types. 

Occurrence in New Guinea. Known only from the types. 

Notes. This very large Clivina has no known close relatives in 
New Guinea. It resembles and may be related to certain Au- 
stralian species of Sloane's procera group, but differs from the 
Australian ones known to me in the strong wrinkling of the 
front of the head. The occurrence of this specialized (flightless) 
species in the Snow Mts. suggests that other species derived from 
the same (Australian) ancestral stock will eventually be found 
elsewhere in the mountains of New Guinea. 

Clivina brevicornis n. sp. 

Description. Cylindrical ; brownish piceous, appendages 
paler; moderately shining, pronotum and elytra without (or 
with indistinct) reticulate microsculpture. Head (fig. 12) .80 
width prothorax ; eyes small but rather prominent ; genae short, 
almost forming right angles with neck ; antennae very short, 
median segments transverse, normally pubescent ; mandibles 
normal, short, curved ; labrum 7-setose ; clypeus broadly emar- 
ginate, with wings continuous with median part, separated from 
preocular lobes by moderate notches; clypeal suture irregularly 
impressed ; facial carinae short ; front irregularly wrinkled and 



DARLINGTON : CARABID BEETLES OP NEW GUINEA 365 

with median fovea, slightly and irregularly punctate ; neck wide, 
not impressed above. Prothorax much longer than wide (L/W 
1.23), widest near base, but not much narrowed anteriorly; 
sides sinuate or broadly emarginate in outline before middle ; 
anterior angles rounded, not produced; posterior angles broadly 
rounded, not dentate ; lateral margins reaching basal margin ; 
disc with median line and anterior transverse line entire, nor- 
mally impressed; surface scarcely strigulose, almost impunctate 
except for an inconspicuous linear group of punctures on each 
side near base, but surface of disc abraded anteriorly or pos- 
sibly with some reticulate microsculpture. Elytra cylindrical, 
scarcely wider than prothorax (E/P 1.05) ; base nearly trun- 
cate ; humeri rather narrowly rounded, not dentate ; sides sub- 
parallel, faintly, broadly sinuate about % from base; striae 
impressed, punctate, the first 4 free at base ; intervals moder- 
ately convex, 8th carinate at base, 3rd inconspicuously 4-pune- 
tate, the punctures almost lost against the punctate 3rd stria. 
Inner wings fully developed. Lower surface: prosternal process 
narrow before anterior coxae ; proepisterna reticulate but not dis- 
tinctly punctate ; abdomen reticulate basally especially at sides, 
shining apically, impunctate ; last ventral with 2 setae on each 
side widely separated. Legs short ; front tibia 3-dentate, the 4th 
(upper) tooth missing; middle tibia (fig. 22) with spur about 
Y 3 from apex and about as long as width of tibia. Measurements: 
length c. 4.3; width between 1.0 and 1.1 mm. 

Type. Holotype (M.C.Z. Type No. 30,154), sex not deter- 
mined, from Dobodura. Papua, New Guinea, Mar.-July 1944 
(Darlington). 

Occurrence in, Neiv Guinea. Known only from the type. 

Measured specimen. The type. 

Notes. This new species is similar to and probably related to 
Clivina bullata Andrewes, of which I have seen the type, from 
Timor, in the British Museum. However, the present new species 
is smaller, with sides of prothorax more sinuate before middle, 
and with front of head much less strongly sculptured than in 
bullata, though on the same pattern. 

(Clivina wallacei Putzeys) 

Clivina castanea Putzeys 1863, Mem. Soe. E. Sci. Liege 18, p. 35 (part) 

(not castanea Westw.) 
Clivina westwoodi Putzeys 1866, Ann. Soc. Ent. Belgique 10, p. 109 (part) 
Andrewes 1926, Ann. Mag. Nat. Hist. (9) 17, p. 373. 



366 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Clivina wallacei Putzeys 1866, Ann. Soc. Ent. Belgique 10, p. 127. 
Andrewes 1926, Ann. Mag. Nat. Hist, (9) 17, pp. 373, 374. 

1929, Ann. Mag. Nat. Hist, (10) 4, p. 355. 

(Selected references only) 

Description. Sufficiently characterized in preceding key to 
species of Clivina. 

Types. The (supposedly) new Guinean cotype of castanea 
Putzeys (westwoodi Putzeys) is in the Putzeys Collection, Brus- 
sels Mus., where Andrewes examined it. Of wallacei, Andrewes 
(1929, p. 355) records seeing 2 specimens, "including the type," 
also in the Putzeys Collection, Brussels; he gives the type 
locality as New Guinea, but Putzeys says the specimens are 
from Celebes, which I think is probably correct (see below). 

Occurrence in New Guinea. The only records are old and 
doubtful (see below). 

Notes. Putzeys originally described his castanea as from Cey- 
lon and New Guinea, and renamed it westwoodi when he found 
that castanea was preoccupied. Andrewes, finding that Putzeys' 
original specimens represented two species, applied the name 
westwoodi to the one from Ceylon. In the meantime Putzeys had 
described wallacei (without noting its resemblance to his west- 
woodi) from specimens from "Makassar (Celebes) " and "Dorey 
(Celebes)," and Andrewes, after examination of Putzeys' types, 
has applied the name to the present species. "Dorey (Celebes) ' 
is a jumbled locality, for Dorey is really in New Guinea. How- 
ever, the "Dorey" label is always to be doubted (see introduc- 
tion, p. 331). I know of no recent material of this species from 
New Guinea, and I suspect that all the old New Guinea records 
are based on mislabeled specimens collected by "Wallace probably 
on Celebes. 

Clivina kulti n. sp. 

Description. Form as figured (fig. 5) ; large; a little broader 
than usual in genus, but slightly narrowed anteriorly, slightly 
flatter above than usual; black, appendages brownish; shining, 
most of upper surface without reticulate microsculpture. Head 
(fig. 9) .69 and .70 width prothorax (in measured specimens) ; 
eyes prominent, genae short, almost forming right angles with 
sides of neck ; antennae normal, median segments about as long 
as wide, normally pubescent; mandibles short, curved; labrum 
5-setose (in all specimens) ; clypeus truncate at middle, clypeal 



DARLINGTON : CARABID BEETLES OP NEW GUINEA 367 

wings continuous with median part but strongly advanced, sep- 
arated from preocular lobes by slight notches; clypeal suture 
not visible; supraocular convexities continuous with preocular 
lobes, separated from front by deep sulci which extend from 
neck constriction to base of clypeus, becoming deeper and sinu- 
ous anteriorly ; front slightly, almost evenly convex, impunctate ; 
neck constriction interrupted at middle, slightly punctate at 
sides. Prothorax slightly Avider than long (W/L 1.10 and 1.13), 
widest not far before posterior angles, rather strongly narrowed 
anteriorly; sides nearly straight and converging in about an- 
terior half ; apex emarginate ; anterior angles narrowly rounded, 
subrectangular ; posterior angles obtuse-rounded, not dentate; 
disc rather flat, with lightly impressed middle line and anterior 
transverse impression ; surface of disc slightly strigulose, vir- 
tually impunctate. Elytra 1.14 and 1.12 width prothorax; base 
broadly emarginate; humeri rounded, not dentate; each elytron 
with usually 4 striae free at base, rarely only 3 (base of 4th 
stria sometimes turned out, rarely joining base of 5th — holo- 
type has only 3 striae free on left elytron, 4 on right) ; striae 
moderately impressed, entire, usually faintly punctulate ; inter- 
vals slightly convex, 3rd 4-punctate (on both elytra of all indi- 
viduals except right elytron of type only 3-punctate). Inner 
wings fully developed. Lower surface: prosternal process rather 
wide before coxae, weakly longitudinally impressed; proepis- 
terna roughened but rather shining; abdomen microreticulate, 
almost impunctate except apical segment rugose-punctate at 
sides; 2 seta-bearing punctures on each side apical segment far 
apart, Legs rather slender (in genus); front tibia 3-dentate; 
middle tibia (fig. 25) with spur on outer side near apex not 
quite as long as width of tibia. Measurements: length c. 8.5-10.4; 
width c. 2.6-3.0 mm. 

Types. Holotype $ (M.C.Z. No. 30,155) and 7 paratypes all 
from Aitape, N-E. N. G., Aug. 1944 (Darlington), taken in flood 
debris in forest or recently forested areas. 

Occurrence in New Guinea. Known only from the type local- 
ity. 

Measured specimens. The $ holotype and 1 (sex not det.) 
paratype. 

Notes. This well defined species is sufficiently characterized 
in the key (above). 



368 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

Clivina biroi Kult 

Kult 1951, Acta Soe. Ent. Czechoslovakia 48, pp. 28, 30. 

Description. Medium sized, moderately broad, subparallel, 
moderately convex ; black, appendages irregularly dark brown, 
antennae paler ; shining, upper surface almost entirely without 
reticulate microsculpture. Head (fig. 10) .71, .71, .71 width 
prothorax (in measured specimens) ; eyes prominent, genae 
short, almost forming right angles with sides of neck; antennae 
normal, median segments as wide as or slightly wider than long ; 
mandibles short; labrum 5-setose (in all specimens); clypeus 
broadly emarginate, with wings continuous with median part 
and somewhat advanced, separated from preocular lobes by 
notches ; preocular lobes with outer edges oblique, nearly straight 
and strongly converging in about anterior % ; clypeal suture in- 
distinct ; supraocular convexities nearly continuous with swollen 
preocular lobes, separated from front by subparallel, irregular 
sulci, which become wider and strongly sinuous anteriorly ; front 
slightly, almost evenly convex or with a slight V-shaped impres- 
sion anteriorly, with a group of punctures near middle anter- 
iorly; neck constriction shallow, narrowly interrupted at middle, 
slightly punctate toward sides. Prothorax about as long as wide 
(L/W 1.02, 1.03, 1.03), nearly quadrate, slightly narrowed an- 
teriorly ; sides vaguely sinuate about anterior % ; apex almost 
truncate, faintly emarginate ; anterior angles almost right, nar- 
rowly rounded, not advanced ; posterior angles obtuse, not or 
vaguely dentate ; lateral margins entire, each with usual 2 setae, 
near basal angle and about % °f prothoracic length from apex ; 
disc rather convex, with usual impressed middle line and anter- 
ior transverse impression, with surface slightly, irregularly 
strigulose, not distinctly punctate. Elytra slightly wider than 
prothorax (E/P 1.08, 1.04, 1.05), rather short (in genus) ; base 
almost truncate ; humeri rounded, not dentate ; each elytron with 
3 or 4 striae free at base ; striae rather deeply impressed, entire, 
finely punctate ; intervals rather convex, 7th and 8th joined near 
base where narrow and convex but scarcely carinate, 3rd 3- 
punctate in all specimens. Inner wings fully developed. Lower 
surface: prosternal process average (neither very broad nor 
very narrow) before coxae; proepisterna with light reticulate 
microsculpture externally, shining and vaguely punctate in- 
ternally ; abdomen with reticulate microsculpture only at base, 
shining in last 3 or 4 segments, punctate especially near sides 
of last 4 segments, and with last segment wholly punctate in 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 369 

some individuals; 2 seta-bearing punctures each side last ven- 
tral segment far apart. Legs rather stout ; front tibia 3-dentate ; 
middle tibia (fig. 23) with spur near apex nearly as long as 
width of tibia. Measurements: length 5.4-6.4; width 1.5-1.8 mm. 
(Knit gives length as 5.9-6.8 mm.). 

Types. Holotype 9 from Sattelberg, N-E. N. G. (Biro, Hun- 
garian National Mus.) and 1 9 paratype from Madang, N-E. 
N. G. (Kult collection). I am indebted to Dr. Z. Kaszab for an 
opportunity to examine the type. 

Occurrence in New Guinea. Papua: 19, Dobodura, Mar.- 
July 1944 (Darlington) ; 1, Milne Bay, Dec. 1943 (Darlington). 
N-E. N. G.: 13, Aitape, Aug. 1944 (Darlington); 1, Nadzab, 
July 1944 (Darlington) ; 3, Chimbu Valley, Bismarck Range, 
5000-7500 ft, Oct. 1944 (Darlington); 1, Adelbert Mts. : Wa- 
numa, 800-1000 m. (c. 2600-3250 ft.), Oct. 23, 1958 (J. L. Gres- 
sitt, Bishop Mus.) taken in light trap. Neth. N. G. : 1, Maffin 
Bay, Aug. 1944 (Darlington). The species usually occurs in 
shaded swamps. 

Measured specimens. The 9 type and a pair ( $ 9 ) from 
Dobodura, listed in this order. 

Notes. Sufficiently compared with other species in the key to 
species of Clivina. 

The type of biroi has the whole last ventral segment closely 
punctate, while most individuals of the species have the last 
ventral nearly smooth at middle, but at least one specimen from 
Dobodura has it closely punctate, showing that the character 
varies individually. The type has only 3 striae free at base of 
each elytron, and this is the case also in the Nadzab, Maffin 
Bay, and most Aitape individuals, but one from Aitape has the 
4th striae more or less free too, as it is in specimens from Chim- 
bu Valley, Dobodura, and Milne Bay. This is an example of 
partly individual and partly geographical variation in a charac- 
ter sometimes considered very important in classification. 

Clivina komareki Kult 

Kult 1951 Acta Soc. Ent. Czechoslovakia 48, pp. 18, 31. 

Description (significant characters only). A medium-sized 
black Clivina, characterized by unusually wide, strongly arcuate 
frontal plates (fig. 11) and unusually deep anterior transverse 
impression of pronotum, and by other characters given by Kult 
and in the key to species of Clivina. 



370 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Type. From "Gulf of Papua," Papua, in Dr. Knit's collec- 
tion. 

Occurrence in New Guinea. Papua-, the type. Neth. N. G. : 1, 
Hollandia, Nov. 1944 (II. Iloogstraal, M.C.Z.). 

Notes. My single specimen of this strongly defined species 
fits the description of komdreJci reasonably well in most ways, 
except that the labrum, described as 6-setose in komdreJci by 
Kult, is only 5-setose. My specimen is in good condition, and 
the 5 labral setae are clearly visible and symmetrically placed. 
This difference (if in fact there is a difference) is not necessarily 
important ; the number of labral setae varies in some other spe- 
cies. The proportions of the Hollandia specimen are: head .77 
width prothorax; prothoracic width/length 1.06; width elytra/- 
prothorax 1.12. Measurements: length c. 5.8, width c. 1.7 mm. 
These figures are close to those given by Kult for the type. 
Kult recognized no close relatives of this species. However, it 
is probably related to and possibly derived from Clivina oiroi 
(above), which has the same general form and group characters 
without the more striking special characters of komdreki. 

Clivina puncticeps n. sp. 

Description. Form as figured (fig. 6) ; rather slender, sub- 
parallel, moderately convex ; black or dark brown, margins of 
elytra sometimes paler, appendages dark brown or rufous; shin- 
ing, upper surface (except at sides) almost without reticulate 
microsculpture. Head (fig. 13) .75 and .75 width prothorax (in 
measured specimens); eyes prominent but smaller than usual; 
genae oblique or rounded-oblique, as long as or not much shorter 
than eyes ; antennae normal, intermediate segments about as 
wide or slightly wider than long ; mandibles short ; labrum usu- 
ally 7-setose, individually 6-setose (e.g. an example from Aitape 
has an intermediate seta missing on the right side) ; clypeus 
truncate or slightly emarginate, separated from wings by mod- 
erate notches; clypeal wings prominent, narrowly rounded, sep- 
arated from preocular lobes by deep notches ; clypeal suture ob- 
solete ; supraocular convexities narrow, almost cariniform, sep- 
arated from swollen preocular lobes; frontal sulci widely sepa- 
rated, slightly diverging anteriorly and posteriorly ; clypeus with 
transverse swollen area or transversely wrinkled ; front irregu- 
larly convex, slightly impressed at middle, irregularly in part 
rather coarsely punctate ; neck constriction slightly impressed 



DARLINGTON: C'AHABID BEETLES OF NEW GUINEA 371 

and punctate at sides, usually (not always) interrupted at mid- 
dle. Prothorax subquadrate, slightly longer than wide, length/ 
width 1.06 and 1.09, only slightly narrowed anteriorly; sides 
usually subsinuate before middle, slightly arcuate in front of 
and behind the sinuation ; lateral margins entire, each with 
usual 2 setae ; anterior margin broadly emarginate ; anterior 
angles rather narrowly rounded, scarcely advanced; posterior 
angles weakly dentate ; pronotum with usual anterior transverse 
impression and longitudinal median line, with a few irregular 
transverse strigae, and finely, irregularly, inconspicuously punc- 
tulate. Elytra slightly Avider than prothorax (measurement of 
width impossible because left elytron of all specimens raised to 
show inner wings), long, subparallel ; base subtruncate-emargi- 
nate, with small tubercles at front ends of 2nd and usually 3rd 
intervals ; humeri rather narrowly rounded ; 3 or 4 inner striae 
of each elytron free at base (individual variation) ; striae 
entire or nearly so, well impressed, distinctly punctate; in- 
tervals moderately convex, 8th finely carinate at base, 3rd 4- 
punctate on outer edge, intervals otherwise scarcely visibly 
punctate. Inner wings fully developed in all specimens. Lower 
surface: proepisterna rugose; abdomen rugose or punctate at 
sides and across almost whole apical segment; 2 apical setae on 
each side widely separated. Legs: anterior tibia strongly 4-den- 
tate (but upper tooth sometimes reduced) ; anterior trochanter 
(of all specimens) with a small acute tooth at apex, on lower 
edge of leg; middle tibia (fig. 24) with a long spur (longer than 
width of tibia) on outer side about one third from apex. Meas- 
urements: length 5.8-6.5; width c. 1.5 mm. (width not measured 
exactly because of raising of elytra). 

Types. Holotype & (M.C.Z. Type No. 30,156) and 13 para- 
types from vicinity of Ilollandia, Neth. N. G., July-Sept. 1944 
(Darlington), and 1 additional paratype from same locality, 
Apr. 1945 (Malkin, U.S.N.M.). Additional paratypes : 2, Aitape, 
N-E. N. G., Aug. 1944 (Darlington), and 1, Idenburg R., Neth. 
N. G., 400 m. (about 1,300 ft.) July 15-Sept. 15, 1938 (J. Olthof, 
Leiden Mus.). My specimens were taken from damp soil but 
not in very wet places. 

Other material. One additional specimen, not a type, from 
Hollandia (Darlington), with front of head abnormally flat- 
tened and wrinkled. Also 1 from Dobodura. Papua, Mar. -July 
1944 (Darlington) that I refer here with doubt: the front is 
flatter than in typical punt ice ps and seems at first impunctate, 



372 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

but vague shallow punctures can be seen on careful examination, 
and the insect is browner (less black) and perhaps slightly more 
depressed than typical panticeps, but otherwise nearly the same. 
I do not know whether it is an aberrant punticeps or a different, 
perhaps geographical form. 

Measured specimens. The holotype and one paratype from 
Hollandia. 

Notes. A very distinct species, placed in relation to others 
in the key to species of Clivina. 

Clivina dealata n. sp. 

Description. Form slender, subparallel, moderately convex ; 
brownish or reddish with disc of elytra sometimes darker ; shin- 
ing, upper surface (except at sides) with reticulate microsculp- 
ture faint (on front) or absent, (except in subsp. antecessor, 
q. v.), but head in part coarsely and pronotum finely punctate. 
Head .74 and .75 width prothorax ; eyes rather small, about as 
long as (oblique) genae ; antennae normal, intermediate seg- 
ments about as wide as long; mandibles short; labrum 7-setose; 
clypeus truncate, middle part usually (not always) separated 
from wings by slight notches, with wings narrowly rounded, 
separated from preocular plates by deeper notches ; clypeal suture 
obsolete ; supraocular convexities almost cariniform, separated 
from swollen preocular plates ; frontal sulci widely separated, 
subparallel, somewhat irregular ; clypeus transversely swol- 
len; front convex, not or irregularly and faintly impressed, 
sometimes with a slight impression at middle, coarsely but vari- 
ably punctate; neck constriction not much impressed, punctate 
especially at sides, usually interrupted at middle. Prothorax 
subquadrate, about as long as wide, length/width .97 and .99, 
slightly narrowed anteriorly; sides nearly straight, slightly con- 
verging anteriorly with entire margins and usual setae ; anterior 
margin slightly emarginate ; anterior angles narrowly rounded, 
scarcely prominent ; posterior angles with distinct blunt teeth ; 
pronotum with usual impressed lines and finely, inconspicuously 
punctate, with a more impressed line of coarser punctures on 
each side behind middle. Elytra slightly wider than prothorax, 
slightly shorter and more oval than in preceding species but 
otherwise similar, with 3 or 4 striae free at base. Inner wings 
strongly reduced, about half as long as elytra (but sometimes 
crumpled). Lower surface rugose and punctate about as in pre- 
ceding species, with similar widely spaced apical ventral setae. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 373 

Legs as in preceding species, with similar acute teeth at tips of 
anterior trochanters. Measurements: length 4.3-5.2; width c. 
1.3-1.4 mm. 

Types. Holotype <J (M.C.Z. No. 30,157) and 15 paratypes 
from Dobodura, Papua, Mar. -July 1944 (Darlington), taken in 
damp soil but not in very wet places. 

Other material. Known only from the types, but the follow- 
ing subspecies represent the species elsewhere in New Guinea. 

Measured specimens. The $ holotype and 1 9 paratype. 

Notes. This new species is evidently related to and perhaps 
derived from the preceding one (punticeps) as shown by many 
characters including the toothed anterior trochanters. I think, 
however, that it is a distinct species. Besides having reduced 
wings, it is smaller and (as shown by the prothoracic propor- 
tions) relatively shorter than puncticeps and paler in color. A 
subspecies of dealata (below) occurs with puncticeps at Hol- 
landia without intergrading. 

Clivina dealata braciiyptera n. subsp. 

Description. Form as figured (fig. 7) ; nearly the same as typi- 
cal dealata but with uniformly shorter wing vestiges, which 
scarcely extend beyond the posterior edge of the metasternum 
and which are only a small fraction as long as the elytra. Pro- 
portions of measured specimens are head .74 and .71 width pro- 
thorax ; length/width prothorax 1.03 and 1.03; relative width 
elytra not measured because left elytron of all specimens raised 
to show inner wings. Measurements: Length 4.5-5.4; width c. 
1.4 mm. 

Types. Holotype $ (M.C.Z. 30,158) and 21 paratypes from 
Hollandia, Neth. N. G., July-Sept. 1944 (Darlington). 

Other material. Six specimens from Aitape, N-E. N. G., Aug. 
1944 (Darlington), and 38 from Maffin Bay, Neth. N. G., Aug. 
1944 (Darlington and E. S. Ross, California Acad.) are referred 
to this subspecies, but not as types. 

Measured specimens. The $ holotype and 1 paratype. 

Notes. See under preceding and following subspecies. 

Clivina dealata antecessor n. subsp. 

Description. Essentially the same as typical dealata and sub- 
species brachyptera but with inner wings fully developed, eyes 
a little larger, and elytra with reticulate microsculpture (with 



374 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

meshes either slightly longitudinal or imperfect). Color of all 
specimens piceous with suture and sides of elytra as well as ap- 
pendages reddish brown. Proportions : head .72 and .75 width 
prothorax ; prothoracic width/length 1.05 and 1.03 ; width ely- 
tra/prothorax c. 1.17 and 1.21. Measurements: length 5.0-5.5; 
width 1.5-1.6 mm. 

Types. Holotype (sex not determined) (Bishop Mus.) and 
3 paratypes all from Kiunga, Fly Ii. Papua, Sept. 24-25 (holo- 
type) and Aug. 1-3, 8-10, 11-13, 1957 (W. W. Brandt). One 
paratype now in M.C.Z. (No. 30,306). 

Measured specimens. The holotype and 1 paratype. 

Notes. These specimens were received after the descriptions 
of dealata and brachyptcra had been drawn. They appear to 
represent the winged population from which the short-winged 
subspecies have been derived. 

Clivina vigil n. sp. 

Description. Subparallel, rather depressed (in genus) ; ru- 
fous, shining, head and discs of pronotum and elytra without 
reticulate microsculpture (which, however, is present laterally). 
Head (fig. 14) .78 and .81 width prothorax; eyes large and 
prominent (in genus), genae very short, forming (blunt) right 
angles with neck ; antennae normal, intermediate segments about 
as long as wide ; mandibles short, normal ; labrum 7-setose ; cly- 
peus typically subtruncate but with angles advanced and denti- 
form in Nadzab example, middle part separated from wings by 
shallow, obtuse notches (possibly variable) ; clypeal wings small, 
arcuate or subangulate, separated from preocular lobes by 
notches; clypeal suture obsolete; supraocular convexities carini- 
form, sharply separated from swollen preocular lobes ; frontal 
sulci rather short, arcuate; front convex (sometimes irregularly 
so), slightly impressed at middle, rather finely and irregularly 
punctate ; neck constriction typically impressed and punctate 
only at sides, widely interrupted at middle (but not interrupted 
in Nadzab example). Prothorax subquadrate, slightly wider 
than long (width/length 1.09 and 1.10), slightly narrowed an- 
teriorly ; sides slightly and almost evenly arcuate, with entire 
margins and usual 2 setae ; front edge slightly emarginate ; an- 
terior angles rounded, not advanced; posterior angles with dis- 
tinct but blunt teeth ; basal marginal gutter wider and more 
rugose than usual ; disc with usual anterior transverse line and 
median longitudinal line rather lightly impressed, and with 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 375 

surface finely, sparsely, inconspicuously punctate. Elytra about 
i/4 wider than prothorax (E/P 1.26 and 1.29), subtruncate at 
base with rounded humeri; sides almost straight (faintly arcu- 
ate) to behind middle, margins crenulate behind humeri; each 
elytron with 4 striae free at base ; striae entire or nearly so, 
moderately impressed, vaguely or not punctate ; intervals slight- 
ly convex, 8th not carinate at humeri, 3rd 4-punctate, intervals 
otherwise not distinctly punctate. Inner wings fully developed. 
Lower surface: proepisterna roughened (deeply microreticulate) 
externally, more or less punctate internally; abdomen impunc- 
tate but with close reticulate microsculpture ; last ventral seg- 
ment with 2 setae on each side wide apart. Legs: front tibiae 
3-dentate ; middle tibiae (fig. 26) with spur on outer side near 
apex about as long as tibial width. Measurements: length c. 
5.7-6.1 ; width e. 1.7-1.8 mm. 

Types. Holotype 9 (M.C.Z. No. 30,159) and 1 paratype from 
Dobodura, Papua, Mar. -July 1944 (Darlington). I do not know 
the habitat of this species. 

OtJier material. One, Nadzab, Markham Valley, N-E. N. G., 
Aug. 1944 (K. V. Krombein, U.S.N.M.), differing from the types 
as indicated in the preceding description. 

Measured specimens: the holotype and paratype from Dobo- 
dura. 

Notes. This species is sufficiently distinguished from other 
New Guinean species in the key above. It resembles and is prob- 
ably related to clenticollis Sloane of Australia (described from 
(NW?) Western Australia and represented in the M.C.Z. by 
specimens from the Burdekin River near Charters Towers, 
Queensland), but differs in detail, notably in the shallower 
clypeal notches and wider basal pronotal gutter of vigil. 

Clivina deleta n. sp. 

Description. Rather broad (in genus), depressed; rufous ; 
shining, reticulate microsculpture faint on head, virtually absent 
on pronotum and elytra. Head .78 and .76 width prothorax; 
eyes smaller and less convex than in preceding species, genae 
oblique ; antennae normal, median segments about as wide as 
long ; mandibles short ; labrum 7-setose ; clypeus subtruncate or 
broadly emarginate, middle part usually (not always) separated 
from wings by slight notches ; clypeal wings small, separated 
from preocular plates by deeper notches ; clypeal suture usually 
not impressed, but front usually with one or two transverse 



376 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

impressed lines which simulate a clypeal suture ; supraocular 
convexities sharply separated from swollen preocular plates; 
frontal sulci deep, slightly diverging and almost reaching neck 
constriction posteriorly ; front moderately convex but usually 
with irregular impressions or transverse impressed lines as noted 
above, not distinctly punctate ; neck constriction shallow, punc- 
tate, sometimes interrupted at middle. Prothorax subquadrate, 
usually slightly narrowed anteriorly, width/length 1.00 and 
1.00; sides very broadly arcuate, almost straight at middle, with 
usual entire margins, and each with usual 2 setae but with 
posterior one farther in from margin than usual ; front of 
prothorax subtruncate or broadly emarginate, with anterior 
angles very narrowly rounded and only slightly advanced ; 
posterior angles distinctly but obtusely dentate ; pronotum with 
well impressed median line but with anterior transverse im- 
pression obsolete at least at middle (an irregular line of black 
pigment under the surface marks the usual position of the 
transverse impression so that it is necessary to look carefully 
to see that the impression itself is absent) ; surface of pronotum 
finely, irregularly, inconspicuously punctate. Elytra about % 
wider than prothorax (E/P 1.31 and 1.31), broadly emarginate 
in front with tubercles at ends of second and third intervals ; 
humeri rounded, margins behind them almost straight to behind 
middle, slightly crenulate behind humeri; 4 striae on each 
elytron free at base ; striae entire, vaguely punctate ; intervals 
slightly convex, 7th briefly and inconspicuously carinate at 
base, 3rd 4-punctate on outer side, intervals vaguely or not 
punctulate. Inner wings fully developed. Lower surface: pro- 
episterna rugose and sides of body posteriorly also more or less 
rugose or subpunctate ; anal segment with usual 2 setae on each 
side widely separated. Legs: front tibia 4-dentate 1 middle tibia 
(fig. 27) with spur on outer side near apex not longer than tibial 
width. Measurements: length 4.5-5.5; width ±1.5 mm. 

Types. Holotype $ (M.C.Z. No. 30,160) and 5 paratypes 
from Dobodura, Papua, Mar.-July 1944 (Darlington). I did 
not distinguish this species in the field and do not know its 
habits (see under following species). 

Measured specimens. The $ holotype and 1 9 paratype. 

Notes. Superficially this species resembles several other small. 



i In one example the right front tibia is only 3 -dentate, without trace of upper 
tooth, although the left front tibia is normally 4-dentate, with upper tooth small 
but distinct and with an apical seta. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 377 

depressed, reddish species found in New Guinea, but it differs 
from the others in (among other things) the partial deletion of 
the anterior transverse impressed line of the pronotum. 

Clivina rufula n. sp. 

Description. Subparallel, slightly depressed ; rufous ; shining, 
almost entire upper surface without reticulate microsculpture 
(but latter sometimes faintly visible on head). Head (fig. 15) .75 
and .74 width prothorax ; eyes rather prominent (but less so than 
in vigil), genae short; antennae normal, median segments about 
as long as wide ; mandibles short ; labrum 7-setose ; clypeus sub- 
truncate, very broadly emarginate, middle part continuous (or 
nearly so) with wings, which are rounded, separated from pre- 
ocular lobes by usually obtuse notches ; clypeal suture absent ; 
supraocular convexities continuous with swollen preocular lobes ; 
frontal sulci well impressed, irregular, diverging anteriorly and 
posteriorly ; front almost evenly convex, sometimes vaguely im- 
pressed at middle, slightly or not punctate; neck constriction 
almost absent, slightly punctate at sides, widely interrupted at 
middle. Prothorax subquadrate, more or less narrowed anter- 
iorly, width/length 1.02 and 1.06; sides very broadly and 
slightly arcuate, with entire margins each with usual 2 setae; 
prothorax anteriorly subtruncate, broadly emarginate ; anterior 
angles very narrowly rounded, not or only slightly prominent; 
posterior angles subdentate; disc with usual impressed lines, 
surface finely and inconspicuously punctate. Elytra nearly 14 
wider than prothorax (E/P 1.24 and 1.22) ; emarginate an- 
teriorly, each with a tubercle at base of 2nd interval; humeri 
rounded, sides behind them very broadly arcuate, almost straight 
at middle, slightly crenulate behind humeri; 3 or 4 striae free 
at base on each elytron ; striae entire, moderately punctate ; 
intervals moderately convex, 7th briefly and inconspicuously 
carinate at humeri, 3rd 3-punctate on outer edge; intervals 
faintly or not distinctly punctulate. Inner wings fully de- 
veloped. Lower surface: sides of body slightly roughened or 
microreticulate, with sides of some ventral segments shallowly 
punctate; last ventral with 2 apical setae on each side wide 
apart. Legs: front tibia 3-dentate on outer side; middle tibia 
(fig. 28) with moderate spur on outer side near apex. Meas- 
urements: length 4.2-4.8; width c. 1.3-1.5 mm. 

Types. Holotype $ (M.C.Z. No. 30,161) and 69 paratypes 



378 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

all from Dobodura, Papua, Mar. -July 1944 (Darlington). Ac- 
cording to my notes, small, depressed, rufous species of Clivina, 
including the present one, were found under stones and other 
cover by rivers, among grass roots in sand, and at light. 

Measured specimens. The S holotype and 1 2 paratype. 

Notes. In form of clypeus and front this species approaches 
the australasiae group, but it differs from typical members of 
the group in having the 3rd elytral interval with 3 rather than 
4 punctures. 

Clivina tripuncta n. sp. 

Description. Rather stout (in genus), normally convex; black 
or dark reddish brown, margins of elytra usually (narrowly or 
broadly) paler, and appendages paler brown; shining, micro- 
sculpture faint or absent on head, virtually absent on pronotum 
except at extreme base and on elytra except at sides and apex. 
Head .71 and .71 width prothorax; eyes prominent but not very 
large, genae oblique, antennae normal, median segments slightly 
wider than long; mandibles short; clypeus subtruncate, broadly 
emarginate, middle part not separated from wings, latter sep- 
arated from preocular lobes by distinct notches ; labrum 7-setose ; 
supraocular convexities slightly or not separated from swollen 
preocular lobes; frontal sulci subparallel, irregular, slightly 
diverging anteriorly and posteriorly; clypeal suture not im- 
pressed; front convex, sometimes irregularly impressed, usually 
more or less punctate at middle; neck constriction impressed 
(and punctate) only at sides, usually widely interrupted at 
middle. Prothorax rather wide, slightly narrowed anteriorly, 
W/L 1.10 and 1.09; sides broadly, slightly arcuate except often 
subsinuate near middle, with entire margins each with usual 2 
setae; prothorax broadly emarginate anteriorly, anterior angles 
very narrowly rounded but scarcely advanced ; posterior angles 
subdentate; disc with usual impressed lines and also some 
scattered punctation of mixed (moderate and fine) punctures. 
Elytra rather short (in genus), about % wider than prothorax 
(E/P 1.21 and 1.20), slightly emarginate anteriorly, with tuber- 
cles at bases of 2nd and usually 3rd intervals ; humeri rounded, 
sides behind them broadly arcuate except almost straight before 
middle, slightly crenulate at and behind humeri; each elytron 
with 3 or 4 striae free at base ; striae well impressed, entire, 
rather finely punctate; intervals moderately convex, 7th not 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 379 

distinctly carinate at humeri, 3rd 3-punctate on outer edge, inter- 
vals not distinctly punctate. Inner wings fully developed. 
Lower surface: proepisterna rugose-punctate; sides of abdomen 
finely rugose near base, subpunctate at sides of posterior seg- 
ments, which are shining, almost without surface sculpture at 
middle ; last ventral with 2 setae on each side wide apart. 
Legs: front tibia 4-dentate but upper tooth minute; middle tibia 
(fig. 29) with a short spur on outer side less than i/i from apex. 
Measurements (Dobodura series only) : length 4.1-5.2; width c. 
1.4-1.7 mm. (specimens from some other localities run larger). 

Types. Holotype S (M.C.Z. No. 30,162) and 37 paratypes 
from Dobodura, Papua, Mar. -July 1944 (Darlington). 

Additional material. Papua: 14, Milne Bay, Aug. 1944 (Dar- 
lington) ; 2, Fly R. 5 mi. below Palmer R., May 1936, and 1, 
Palmer R. at Black R., June 1936 (Archbold Exp., A.M.N.H.). 
N-E. N. G.: 2, Torricelli Mts., Mobitei, 750 m. (c. 2450 ft.), 
Feb. 28-Mar. 4 and Mar. 5-15, 1959 ("W. W. Brandt, Bishop 
Mus.); 1, Torricelli Mts., Mokai Village, 750 m. (c. 2450 ft,), 
Jan. 1-23, 1959 (W. W. Brandt, Bishop Mus.) ; 1, Adelbert Mts., 
Wanuma, 800-1000 m. (c. 2600-3250 ft.), Oct. 24, 1958 (J. L. 
Gressitt, Bishop Mus.) ; 1, Sambeang, 400 m. (about 1300 ft.), 
Mongi Watershed Huon Peninsula, Apr. 21, 1955 (E. O. Wil- 
son, M.C.Z.) ; 21, Aitape, Aug. 1944 (Darlington). Neth. N. G. : 
3, Hollandia, July-Sept. 1944 (Darlington), 3, same locality, 
May 1945 (H. Hoogstraal, M.C.Z.), and 1, same locality, Feb. 
12, 1945 (Hoogstraal, Chicago Mus.) ; 1, Sabron, Cyclops Mts., 
Camp 1, 1,200 ft,, May 15, and 1, same locality, Camp 2, 2.000 
ft,, July 1936 (Cheesman) ; 4, Maffin Bay, Aug. 1945 (Darling- 
ton), and 1, same locality, Oct. 1944 (K. V. Krombein, 
U.S.N.M.) ; 1, Fac Fac, June 1939 (Wind, M.C.Z.) ; 2, "Neth. 
New Guinea," Sept, 1944 (T. Aarons, California Acad.) ; 1, 
Camp 1, Mt. Nok, Waigeu Is., 2,500 ft., May 1938 (Cheesman). 
Of these, 1 of my Hollandia specimens and the single specimens 
from Fac Fac and Waigeu Is. have the 3rd intervals with 4 
(not 3) punctures; 1 of my Hollandia specimens has the left 
elytron 3- and the right only 2-punctate ; and the specimens 
from the Fly and Palmer R. and Sambeang, though 3-punctate, 
are larger than usual, about 6 mm. or a little longer. I refer 
these specimens to this species with some doubt. The species 
apparently occurs throughout New Guinea at low altitudes, usu- 
ally, I think, in shaded swamps. I have a typical specimen also 
from Cape Gloucester, New Britain. 



380 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Measured specimens. The $, holotype and 1 9 paratype. 

Notes. This somewhat variable species is further discussed 
and compared in "Notes" under the following species, eruga- 
tella. 

Clivixa erugatella n. sp. 

Description. Kather stout, slightly depressed; black or pice- 
ous, sometimes with sides of elytra paler, legs rather dark brown, 
antennae paler brown; shining, reticulate microsculpture faint 
or absent on front, present at sides and base but indistinct on 
disc of pronotum, and usually absent or indistinct on main part 
of elytral disc but present on sides and apex of elytra and some- 
times extending to part of disc. Head .63 and .67 width pro- 
thorax ; eyes not large but rather prominent ; genae rather 
short ; antennae normal, median segments about as long as wide ; 
mandibles short, labrum 7-setose ; clypeus subtruncate except 
wings slightly advanced, middle part continuous with wings, 
latter separated from preocular lobes by shallow notches; cly- 
peal suture absent ; supraocular convexities smoothly continuous 
with swollen preocular lobes ; frontal sulci subparallel, sinuous 
anteriorly ; front varying from smoothly convex and continuous 
with clypeus to slightly irregularly impressed, sometimes with 
median impression, usually finely and inconspicuously punctulate, 
sometimes with a few coarser punctures ; neck slightly impressed 
at sides, not at middle. Prothorax rather large, with pronotum 
rather depressed; width/length 1.09 and 1.13; somewhat nar- 
rowed anteriorly ; broadly emarginate in front, anterior angles 
very narrowly rounded, almost angulate, not produced; pos- 
terior angles obtusely subangulate, subdentate ; sides broadly and 
slightly arcuate, each with entire margin and usual 2 setae ; disc 
with usual impressed lines, surface rather finely and incon- 
spicuously punctulate. Elytra rather short (in genus), slightly 
wider than prothorax (E/P 1.14 and 1.16) ; slightly emarginate 
anteriorly, with tubercles at bases of first three intervals ; each 
elytron with 3 striae free at base (in all specimens) ; striae 
entire, well impressed, finely punctate or subpunctate ; intervals 
moderately convex, 7th briefly or not distinctly cariniform at 
base, third usually 4-punctate (see notes), surface of intervals 
not or indistinctly punctulate. Inner wings fully developed in 
type series, but reduced in some other specimens (see notes). 
Lower surface: proepisterna with isodiametric microsculpture 
(but not punctate) externally, rugose or punctate internally; 



DARLINGTON: CARABID BEETLES OF NEW GUINEA 381 

sides of posterior ventral segments not or lightly punctate ; retic- 
ulate microsculpture visible but lightly impressed at middle of 
abdomen ; apical ventral segment with usual 2 setae on each side 
wide apart. Legs: anterior tibia weakly 3-dentate ; middle tibia 
(fig. 30) with a moderate spur on outer side near apex. Measure- 
ments: length 5.0-6.6; width 1.6-2.1 mm. 

Types. Holotype 6 (M.C.Z. No. 30,163) and 32 paratypes 
from Hollandia, Neth. N. G., July-Sept. 1944 (Darlington). 

Other material N-E. N. G.: 36, Aitape, Aug. 1944 (Darling- 
ton). Neth. N. G.: 8, Maffin Bay, Aug. 1944 (Darlington), 
and 2, same locality, Aug. and Sept, 1944 (E. S. Ross, California 
Acad.). 

Measured specimens. The S holotype and 1 9 paratype. 

Notes. Most individuals of this species have 4 punctures on 
the third interval of each elytron, but there are a few excep- 
tions: one from Hollandia has only 3 punctures on each side; 2 
specimens from Aitape have 5 punctures on one side, 4 on the 
other, and one has 4 on one side, 3 on the other ; and one speci- 
men from Maffin Bay has 4 on one side, 3 on the other. All 
specimens from Hollandia and Maffin Bay have the inner wings 
fully developed, but of the 36 from Aitape only 2 have fully 
developed wings and the other 34 have the wings reduced to 
about half the length of the elytra. 

This species is very similar to the preceding one, tripuncta, 
but I feel sure they are different species, although no single 
character will distinguish every specimen. The numbr of punc- 
tures on the third elytral interval will distinguish most speci- 
mens. On direct comparison the present species, erugatella, has 
slightly smaller eyes than tripuncta, a slightly different clypeal 
outline (more truncate at middle with wings more distinctly but 
still slightly advanced), usually less punctate head, relatively 
slightly larger and flatter prothorax, and abdomen less punc- 
tate but with more distinct reticulate microsculpture, and at Ait- 
ape erugatella usually has reduced wings, tripuncta always fully 
developed ones. In tripuncta the elytra have 3 or 4 striae free 
at base ; in erugatella, always 3. Moreover, tripuncta apparently 
occurs throughout New Guinea, while erugatella, though com- 
mon where it occurs, is known only from three localities all near 
the middle part of the north coast. I did not clearly distinguish 
these two species in the field. They probably both occur in wet 
places, but I do not know whether they actually occur together. 



382 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Clivina erugata n. sp. 

Description. About average form and convexity; black or 
piceous, appendages reddish or brownish ; reticulate microsculp- 
ture faint or absent on front of head and on middle of pronotum 
and elytra, but present laterally. Head (fig. 17) .68 and .67 
width prothorax ; eyes normally convex, genae short and oblique ; 
antennae normal, median segments about as long as or slightly 
longer than wide ; mandibles short ; labrum 7-setose ; clypeus 
broadly emarginate, with wings not separated from middle but 
slightly advanced, separated from preocular lobes by distinct 
notches ; clypeal suture absent ; supraocular convexities continu- 
ous with or only slightly separated from swollen preocular lobes ; 
frontal sulci subparallel, sinuous anteriorly ; front almost evenly 
convex or slightly and irregularly impressed, slightly punctulate, 
sometimes with a few coarser punctures near middle ; neck con- 
striction slightly impressed and punctate at sides, usually inter- 
rupted at middle. Prothorax about as long as wide (L/W 1.05 
and .96), narrowed anteriorly (B/A 1.52 and 1.57) ; subtruncate 
or broadly emarginate anteriorly ; anterior angles narrowly 
rounded, not advanced ; posterior angles obtusely angulate, 
bluntly dentate ; lateral margins entire, each with usual 2 setae ; 
disc normally convex, with usual impressed lines, and faintly 
punctulate. Elytra slightly wider than prothorax (E/P 1.21 
and 1.25) ; base slightly emarginate, with tubercles at bases of 
2nd and 3rd intervals; humeri rounded, sides behind humeri 
nearly straight to or beyond middle ; each elytron with usually 
3 (sometimes 4) striae free at base; striae entire, well impressed, 
punctulate ; intervals moderately convex, 3rd 4-punctate, 7th 
and 8th united but scarcely (or briefly) carinate at humerus, 
intervals not distinctly punctulate. Inner wings fully developed. 
Lower surface: proepisterna with reticulate microsculpture and 
some wrinkles externally, lightly rugose internally ; abdomen 
microreticulate, vaguely punctate laterally ; last ventral with 2 
setae each side far apart. Legs: front tibia 3-dentate; middle 
tibia (fig. 31) with a moderate spur on outer side near apex. 
Measurements: length 6.5-8.0; width 2.0-2.5 mm. 

Types. Holotype S (M.C.Z. No. 30,164) and 28 paratypes 
from Dobodura, Papua, Mar. -July 1944 (Darlington). 

Other material. Papua: 10, Oro Bay, Dec. 1943 (Darlington). 
N-E. N. G. : 3, Aitape, Aug. 1944 (Darlington). Neth. N. G. : 
8, Hollandia, July- Sept. 1944 (Darlington), and 1, same local- 
ity, Apr. 1945 (B. Malkin, U.S.N.M.) ; 2, Hollandia area, W. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 383 

Sentani, Cyclops Mts., 50-100 m. (c. 150-325 ft.), June 22-24, 
1959 (J. L. Gressitt, Bishop Mus.) in light trap; 13, Maffin Bay, 
Aug. 1944 (Darlington), and 3, same locality, June and Aug. 
1944 (E. S. Ross, California Acad.); and 1, "Neth. New 
Guinea," Oct. 20, 1944 (T. Aarons, California Acad.). I have 
this species also from Cape Gloucester, New Britain, Jan.-Feb. 
1944 (Darlington), and Muda P. T. Area, New Georgia Is., 
British Solomons, Nov. 20, 1943 (J. G. Franclemont, Cornell 
U. Coll.). 

Measured specimens. The S holotype and 1 $ paratype. 

Notes. This is the only species I know from New Guinea that 
comes close to answering the short description of Clivina schau- 
bergeri Kult, but schaubcrgcri is evidently slightly larger, with 
more slender antennae than the present species. 

Clivina basalis Chaudoir 

Chaudoir 1843, Bull. Soc. Nat. Moseou 16, No. 4, p. 733. 

Sloane 1896, Proc. Linn. Soc. New South Wales 21, pp. 212 (key), 213. 

1905, Proe. Linn. Soc. New South Wales 29, p. 721 (key). 

Csiki 1927, Coleop. Cat., Carabidae, Carabinae, p. 498 (see for synonymy 

and additional references). 
Clivina ephippiata Putzeys (new synonym). 
Putzeys 1846, Mem. Soc. E, Sci. Liege 2, p. 602. 

Sloane 1920, Proe. Linn. Soc. New South Wales 45, p. 320 (see for Aus- 
tralian synonyms ) . 
Csiki 1927, Coleop. Cat., Carabidae, Carabinae, p. 502 (see for additional 
references). 

Description. See key to species of Clivina, and following notes. 

I 1 !ipes. Of basalis, from "Nouvelle Hollande" (= Australia), 
presumably now in the Oberthiir collection, Paris Mus. ; of 
ephippiata, from Java, in the Chevrolat collection, Oxford Uni- 
versity Mus. 

Occurrence in New Guinea. Papua: 5, Dobodura, Mar.-July, 
1944 (Darlington), taken at light; 1, Oro Bay, July 12, 1944 
(A. H. Mallery, Bishop Mus.). Also 1 from Koitaki, N. G., 
1,500 ft., Oct. -Nov. 1928. (Pemberton, Hawaiian Sugar Planters' 
Association) ; I have been unable to locate this locality. 

Notes. This species varies in color both individually and geo- 
graphically. Specimens from temperate southeastern Australia 
{e.g. Sydney and Brisbane) and from New Guinea and Celebes 
usually have the elytra bi colored, red anteriorly and black pos- 
teriorly, with the black color often reaching the 9th intervals 



384 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

and margins laterally (basalis). Specimens from tropical Au- 
stralia and Java usually have black area of the elytra reduced to 
a large, but variable, post-median macula (ephippiata) . There 
is some individual variation too, and entirely pale (immature?) 
individuals sometimes occur with the bicolored ones. This widely 
distributed species (if it is all one species) deserves more study 
than I can give it now. For the present I can see no real dif- 
ference except color to separate the geographical populations, and 
the color forms are distributed so irregularly (discontinuously) 
that I do not care to treat them as subspecies. Sloane (1920) 
has already indicated the apparent identity of the tropical Aus- 
tralian and Javan forms. 

Clivina subfusa n. sp. 

Description. A little broader than usual in genus ; normally 
convex ; reddish piceous, disc of elytra often darker ; reticulate 
microsculpture faint or absent on front of head and discs of 
pronotum and elytra, present on posterior declivity of pronotum 
(isodiametric or slightly transverse) and sides and apices of 
elytra (slightly longitudinal). Head .68 and .60 width pro- 
thorax ; eyes moderately prominent but enclosed behind by short 
genae which form obtuse ( nearly right ) angles with neck ; 
antennae normal, middle segments slightly longer than wide ; 
mandibles short ; labrum 7-setose ; clypeus broadly emarginate, 
wings not separated from middle part but separated from pre- 
ocular lobes by moderate notches ; clypeal suture absent ; supra- 
ocular convexities separated from preocular lobes by impres- 
sions; frontal sulci curving outward posteriorly, almost straight 
between eyes, sinuous anteriorly ; front irregularly punctate ; 
neck constriction scarcely impressed but marked at sides by 
punctate areas which are usually (not always) separated by an 
impunctate median space. Prothorax slightly wider than long, 
W/L 1.07 and 1.08; base/apex 1.45 and 1.43; sides slightly 
arcuate ; anterior margin broadly emarginate, but anterior an- 
gles scarcely advanced ; posterior angles obtuse, dentate ; median 
longitudinal and anterior transverse impressed lines normal ; 
disc finely punctate. Elytra 1.30 and 1.23 width prothorax; 
humeri normally rounded; each elytron with usually 3 (rarely 
4) striae free at base ; striae entire, well impressed, punctulate ; 
intervals moderately convex, inner ones tuberculate or sub- 
tuberculate at extreme base, 7th subcarinate at base, 3rd with 
4 discal punctures (sometimes very inconspicuous) near outer 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 385 

edge ; surface of intervals vaguely or not punctulate. Inner 
wings fully developed. Lower surface: proepisterna strongly ru- 
gulose, and also lightly microreticulate externally; ventral seg- 
ments microreticulate, and with punctate-rugulose areas later- 
ally, these areas being near the anterior margins of the last three 
segments and entirely across the base of the last segment ; last 
ventral segment with 2 setae on each side widely separated. 
Legs: anterior tibiae 3-dentate; middle tibiae (fig. 32) with 
spur short, near apex. Measurements: length 6.6-7.3; width 2.1- 
2.3 mm. 

Types. Holotype (sex not det.) (M.C.Z. No. 30,165) and 2 
paratypes from Chimbu Valley, Bismarck Range, N-E. N. G., 
5,000-7,500 ft., Oct. 1944 (Darlington), taken in open country, 
presumably in wet places. Additional paratypes as follows. 
N-E. N. G. : 1, Baindoang, Salawaket Range, 1800 m. (c. 5850 
ft.), Sept. 15, 1956 (E. J. Ford Jr., Bishop Mus.). Neth. N. G. : 
1, Baliem Camp, Snow Mts., 1,600 m. (about 5,200 ft,), Dec. 
1938 (Toxopeus) ; 4, Ibele (Iebele) Camp, Snow Mts., 2,250 m. 
(about 7,325 ft.), Nov.-Dec. 1938 (Toxopeus); 1, Fac Fac, S. 
coast of Bomberai, 100-700 m. (c. 325-2300 ft,), June 9, 1959 
(J. L. Gressitt, Bishop Mus.) in light trap. 

Other material. One, Dobodura, Papua, Mar. -July 1944 
(Darlington). This specimen is not typical; it has the 3rd in- 
tervals only 3-punctate ; more material is needed to show whether 
it is really subfusa or whether it may be a lowland form related 
to (mountain-living) subfusa. 

Measured specimens. The holotype and one paratype from 
Chimbu Valley. 

Notes. This species is distinguished from others in New 
Guinea by characters given in the key, above. It is similar to 
trnncata Putzeys as identified by Andrewes, but, as compared 
with Andrewes' specimen of truncata, the new species has small- 
er eyes and differs slightly in other ways : e.g. the rugose-punc- 
tate line across the neck is almost entire in truncata, usually 
(but not always) widely interrupted at middle in subfusa. 

Clivina gressitti n. sp. 

Description. Slightly more parallel and more convex than 
average (incipiently subcylindrical, but not strongly so) ; ru- 
fous; rather shining, reticulate microsculpture almost absent 
above. Head .70 and .67 width prothorax ; eyes of only moderate 
size but prominent, genae forming c. right angles with sides of 



386 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

neck; antennae normal, rather short, intermediate segments as 
wide as or slightly wider than long ; mandibles short ; labrum 
7-setose; elypeus subtnmcate with wings slightly advanced, 
middle part not separated from wings; wings separated from 
preocular lobes by shallow notches ; clypeal suture absent ; su- 
praocular convexities separated or very nearly separated from 
preocular lobes by slight impressions; frontal sulci long, some- 
what diverging posteriorly, sinuous anteriorly ; front with or 
without median fovea but always with a few scattered median 
punctures and sparse, scattered punctules; neck constriction 
weak, punctate only at sides, interrupted at middle. Prothorax 
slightly longer than wide (L/W 1.05 and 1.02), moderately 
narrowed in front, slightly so behind (base/apex 1.29 and 1.38) ; 
sides nearly straight and converging anteriorly for much of 
length, faintly sinuate before middle ; apex subtruncate or very 
broadly emarginate ; posterior angles obtuse-rounded, finely 
bluntly denticulate; disc with usual impressed lines, more dis- 
tinctly punctate than usual, but punctures rather widely spaced 
and variable in size; no distinct (or at most an indefinite) line 
of coarser punctures each side basally. Elytra only slightly 
wider than prothorax (E/P 1.09 and 1.04) ; humeri rounded- 
prominent ; margins slightly crenulate behind humeri ; each ely- 
tron with 3 striae free at base ; striae entire (slightly abbreviated 
apically as usual), moderately impressed, punctulate; intervals 
convex, not distinctly punctulate, 3rd very inconspicuously 4- 
punctate, 8th rather weakly carinate or subcarinate at base. 
Inner wings fully developed. Lower surface: proepisterna and 
sides and apex of abdomen rather coarsely punctate ; last ven- 
tral with 2 apical setae on each side wide apart. Legs: front 
femur very stout ; front tibia 3 -dentate with 4th tooth indicated ; 
front trochanter not prominent at apex; middle tibia (fig. 36) 
with spur short (about y 2 long as width of tibia), less than V4 
from apex of tibia. Measurements: length 5.3-5.5; width 1.4-1.5 
mm. 

Types. Holotype (sex not determined) (Bishop Mus.) and 
3 paratypes (1 in M.C.Z., No. 30,307) all from Kiunga, Fly R., 
Papua, various dates in July, Aug., and Oct., 1957 (W. W. 
Brandt). 

Measured specimens. The type and 1 paratype. 

Notes. This species has many of the same technical characters 
as subfusa, including the short tibial spur, but differs notably 
in form, being more slender and parallel-sided. The difference is 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 387 

well shown by the difference in proportions of prothorax and in 
ratio of elytra/prothorax. 

Clivina sellata Putzeys 

Putzeys 18(36, Ent. Zeitung (Stettin) 27, p. 40. 

Sloane 1905, Proc. Linn. Soc. New South Wales 29, p. 719 (in key). 

Clivina inconspicua Sloane (new synonym). 

Sloane 1890, Proc. Linn. Soc. New South Wales 21, p. 277. 

— 1905, Proc. Linn. Soc. New South Wales 29, p. 719 (in key). 
(Unimportant references omitted.) 

Description. See key. C. sellata is a small, rather convex 
(subcylindrical) species, typically (in Australia) dark brown, 
with elytra paler with a post-median discal dark spot ; the 
(color form?) inconspicua. is entirely testaceous. 

Types. Of sellata, from Australia, in the Chaudoir collec- 
tion now presumably with the Oberthur collection, Paris Mus. ; 
of inconspicua, from King's Sound, Australia, in Macleay Mus., 
Sydney. 

Occurrence in New Guinea. Papua: 1, Dobodura, Mar.-July, 
1944 (Darlington). 

Notes. C. sellata is widely distributed in eastern and north- 
ern Australia. Testaceous individuals are to some extent geo- 
graphically segregated, but they do not seem to form a clearly 
defined geographical subspecies. The single example from New 
Guinea is testaceous and is less convex than usual in sellata, but 
I find no positive characters to separate it. C. sellata and related 
species in Australia are difficult to distinguish and in need of 
study. 

In Australia, sellata occurs in sandy banks of streams. 

Clivina ferruginea Putzeys 

Putzeys 1868, Ann. Soc. Ent, Belgique 11, p. 14. 

Sloane 1896, Proc. Linn. Soc. New South Wales 21, pp. 198, 199, 275. 

1905, Proc. Linn. Soc. New South Wales 29, p. 720 (key). 

(Unimportant references omitted.) 

Description. See key; ferruginea is a plain brown species of 
the difficult australasiae group, with 4-dentate anterior tibiae. 

Type. Prom Eockhampton, Australia, in the Castelnau col- 
lection. The collections and parts of collections made by Castel- 
nau are widely scattered. I do not know where the type of fer- 
ruginea now is. I did not find it at Melbourne in 1957. 



388 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

Occurrence in New Guinea. Papua: 2, Port Moresby, Feb.- 
May 1943 (W. B. Jones, A. M. N. H.) ; 1, same locality, May 
1947 (L. Jones, British Mus.). 

Notes. This species is common and widely distributed in 
northern Australia, in wet ground near water. That it extends 
to southern New Guinea is not surprising. The country around 
Port Moresby is much like parts of northern Australia. 

Clivina fessa n. sp. 

Description. Slightly broader and more depressed than us- 
ual ; reddish-piceous, appendages paler ; moderately shining, but 
elytra with at least traces of reticulate microsculpture extend- 
ing onto disc. Head (fig. 16) .76 and .73 width prothorax; eyes 
large (in genus) ; genae short, not entirely enclosing eyes behind, 
forming right angles with sides of neck ; antennae normal, inter- 
mediate segments about as long as wide; mandibles short; lab- 
rum 7-setose; clypeus subtruncate, very broadly emarginate, 
with middle part not separated from wings but latter separated 
from preocular lobes by moderate notches; clypeal suture ab- 
sent ; supraocular convexities separated from preocular lobes by 
variable (sometimes slight) impressions; frontal sulci straight 
and diverging posteriorly, irregular and curving outward an- 
teriorly; front irregularly convex, finely and sparsely punctate 
or with coarser punctures anteriorly: neck scarcely impressed, 
punctate at sides, zone of punctures irregularly interrupted at 
middle. Prothorax slightly wider than long (W/L 1.10 and 
1.08); base/apex 1.33 and 1.29; sides slightly arcuate; front 
margin truncate-emarginate, with anterior angles scarcely ad- 
vanced; posterior angles rounded-obtuse but marked by blunt 
teeth ; median longitudinal line and anterior transverse line well 
impressed; disc punctulate and transversely strigose but with- 
out reticulate microsculpture except at sides and base (but retic- 
ulations sometimes extend to parts of disc). Elytra somewhat 
wider than prothorax (E/P 1.34 and 1.30) ; humeri rounded but 
moderately prominent ; sides subparallel, subcrenulate behind hu- 
meri ; each elytron with 3 striae free at base; striae entire, mod- 
erately impressed, faintly punctulate; intervals slightly convex, 
3rd 4-punctate on outer edge, neither 7th nor 8th distinctly 
carinate at base; intervals with reticulate microsculpture very 
lightly impressed on disc and sometimes absent in an anterior- 
median area. Inner wings fully developed. Lower surface: 
proepisterna rather strongly rugose or punctate-rugose ; ventral 



DARLINGTON: CARABID BEETLES OF NEW GUINEA 389 

segments of abdomen rugose or punctate laterally, the rugose- 
punctate areas almost meeting across the base of the apical seg- 
ment, which is otherwise lightly microreticulate ; two apical 
setae on each side wide apart. Legs: anterior femora (fig. 38) 
more slender than usual; anterior tibia strongly 3-dentate with 
4th (upper) tooth barely indicated; middle tibia (fig. 33) with 
spur about as long as width of tibia, and less than *4 tibial 
length from apex. Measurements (types only): length 6.6-6.9; 
width 2.0-2.1 mm. 

Types. Holotype (M.C.Z. No. 30,166) (sex not clet.) from 
Hollandia, Neth. N. G., July-Sept. 1944 (Darlington) ; 1 para- 
type, same locality, May 1945 (Hoogstraal, M.C.Z.) ; 1 para- 
type, Fac Fac, Neth. N. G., June 1939 (R. G. Wind, M.C.Z.). 

Other material. One, Camp Nok, Waigeu Is., 2,500 ft., April 
1938 (Cheesman) ; this specimen is smaller (c. 5.6 mm.) than 
the types, with elytral margins more strongly crenulate. Also 1, 
Guadalcanal, Solomons, July 15, 1943 (P. W. Oman, U.S.N.M.), 
which has the characters of fessa except that the 7th elytral in- 
terval is almost carinate at base. 

Measured specimens. The type and paratype from Hollandia. 

Notes. Although this species is superficially rather similar 
to several others that occur in New Guinea, it is well character- 
ized by the relatively slender anterior femora, the position of 
the spur of the middle tibia, and (usually) the lack of a distinct 
carina at the base of the 7th or 8th elytral intervals. I do not 
know the relationships of the species. 

Clivina sansapor n. sp. 

Description. Subparallel, about average convexity; reddish 
piceous, appendages paler; moderately shining, most of upper 
surface without reticulate microsculpture but latter visible at 
extreme margins and basal declivity of pronotum and sides and 
apex of elytra. Head .73 and .71 width prothorax; eyes mod- 
erately prominent but almost enclosed behind by short genae, 
which form almost right angles with neck; antennae normal, 
middle segments about as long as wide ; mandibles short ; lab- 
rum 7-setose; clypeus subtruncate, or very broadly emarginate, 
with wings not separated from middle part but separated from 
preocular lobes by moderate notches; clypeal suture absent; 
supraocular convexity sharply separated from swollen preocular 
lobes ; frontal sulci subparallel between eyes, sinuous anteriorly ; 
front slightly convex with a few punctures near middle : neck 



390 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

constriction only a little impressed but marked by lateral punc- 
tate areas which meet or almost meet at middle. Prothorax as 
long as or slightly longer than wide (L/W 1.04 and 1.00) ; base/ 
apex 1.33 and 1.36; sides almost straight (actually faintly 
subsinuate) in outline, slightly converging anteriorly; apex 
subtruncate, very broadly emarginate ; anterior angles not ad- 
vanced ; posterior angles obtuse, marked by distinct blunt denti- 
cles ; median longitudinal line and anterior transverse line nor- 
mally impressed; a well marked longitudinal group of punc- 
tures on disc on each side behind middle slightly nearer margin 
than median line ; disc otherwise at most faintly punctulate, with 
a few irregular transverse strigae. Elytra slightly wider than 
prothorax (E/P 1.14 and 1.13) ; humeri rounded but prominent; 
each elytron with three striae free at base ; striae well impressed 
basally, very light toward apex, punctulate; intervals slightly 
convex, 3rd 4-punctate on outer edge, 8th carinate at base ; sur- 
face of intervals not distinctly punctulate. Inner wings fully 
developed. Lower surface: proepisterna rugulose-punctate in- 
ternally, microreticulate externally; abdomen microreticulate, 
more or less punctate laterally; last ventral segment punctate 
especially laterally but with some punctures scattered over most 
of surface, with 2 apical setae on each side wide apart. Legs 
rather stout; anterior tibiae strongly 3-dentate, with 4th (up- 
per) tooth indicated by a slight obtuse angle; middle tibiae (fig. 
34) with spur long, about Y 3 from apex. Measurements: length 
6.1-6.6 ; width 1.6-1.7 mm. 

Types. Holotype £ (M.C.Z. No. 30,167) and 3 paratypes all 
from Sansapor, Neth. N. G., Aug. 1944 (Darlington). 

Other material. Three, Kiunga, Fly R., July 23-25, Aug. 5-7, 
14-17, 1957 (W. W. Brandt, Bishop Mus.). These specimens are 
doubtfully assigned here. They may be variants of brandti. 

Measured specimens. The holotype and 1 paratype. 

Notes. This species is distinguished from similar ones in New 
Guinea primarily by the ventral punctation. I do not know its 
relationships. 

Clivina csikii Kult 

Kult 1951, Acta Soc. Ent. Czechoslovakia 48, pp. 29, 30. 

Description (checked against the type). More slender and 
convex than average ; nearly black, appendages yellowish ; rather 
shining, reticulate microsculpture indistinct or absent on front 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 391 

and on discs of pronotnm and elytra, present at sides and base 
pronotum and sides and apex of elytra. Head .80 width pro- 
thorax; eyes moderately large and prominent, genae forming c. 
right angles with sides of neck; antennae normal, intermediate 
segments as wide as or slightly wider than long; mandibles 
short; labrum 7-setose; clypens subtrnncate, very slightly emar- 
ginate, middle part not separated from wings and wings not 
advanced; wings separated from preocular lobes by moderate 
notches; clypeal suture absent; supraocular convexities sep- 
arated from preocular lobes by well defined impressions ; frontal 
sulci straight and parallel between eyes, sinuous anteriorly; 
front with slight median impression, rather vaguely punctate ; 
neck impression rather weak, punctate at sides, interrupted at 
middle. Prothorax longer than wide (L/W 1.09), slightly nar- 
rowed anteriorly (base/apex 1.27) ; sides nearly straight; apex 
subtruncate (broadly emarginate at middle) ; posterior angles 
rounded except marked by very faint sinuations ; disc with usual 
impressed lines, finely and sparsely punctulate, with line of 
coarser punctures each side behind middle. Elytra slightly wider 
than prothorax (E/P 1.14); humeri prominent but rounded; 
each elytron with 3 striae free at base; striae entire (except nor- 
mally abbreviated at extreme apex), well impressed, punctulate; 
intervals convex, 3rd 4-punctate on outer edge, 8th forming a 
long, fine carina at base. Inner wincjs fully developed. Lower 
surface: proepisterna microreticulate or rugulose but not punc- 
tate; most of abdomen including most of last ventral segment 
closely microreticulate but not punctate; last ventral segment 
with 2 setae on each side wide apart. Legs: front tibiae strongly 
3-dentate with 4th (upper) tooth indicated; middle tibia with 
spur about 14 from apex and about as long as tibial width. 
Measurements: length c. 5.0 (given by Kult as 4.8, but the speci- 
men is not quite straight) ; width c. 1.3 mm. 

Type. From Madang (Friedrich-Wilh.-hafen), N-E. N. G., 
1901 (Biro, Hungarian National Mus.). I am indebted to Dr. 
Z. Kaszab for an opportunity of examining it. 

Measured specimen. The type. 

Notes. I have seen no other specimen of this species. It oc- 
curs within the geographical range of the following species, 
which is probably related but distinct. See notes under the fol- 
lowing species. 



392 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Clivina inopaca n. sp. 

Description. Form average, normally convex ; black or 
brownish; antennae, mouth parts, and legs (irregularly) brown; 
shining, microsculpture indistinct or absent on discs of prono- 
tum and elytra, present at sides and base of pronotum and sides 
and apex of elytra. Head. .74 and .75 width prothorax; eyes 
moderately large and prominent; genae short, forming c. right 
angles with neck ; antennae normal, intermediate segments about 
as long as wide ; mandibles short ; labrum 7-setose ; clypeus very 
broadly emarginate, middle part not separated from wings, lat- 
ter separated from preocular lobes by variable notches; clypeal 
suture absent; supraocular convexities separated from preocu 
lar lobes by impressions; frontal sulci straight and parallel be- 
tween eyes, sinuous anteriorly; front with median impression 
and a few scattered punctures ; neck only slightly impressed but 
with transverse punctate areas on each side, sometimes almost 
meeting at middle. Prothorax as wide as long (W/L 1.00 and 
1.03), moderately narrowed anteriorly (B/A 1.37 and 1.34); 
sides slightly arcuate, almost straight at middle, sometimes sub- 
sinuate, sometimes slightly crenulate ; apex subtruncate, slightly 
emarginate, with angles scarcely advanced; posterior angles 
broadly rounded, each marked by a slight, blunt tooth ; disc with 
usual impressed lines, with longitudinal group of punctures on 
each side behind middle slightly nearer side than middle, and 
with a little very fine, sparse, inconspicuous punctulation else- 
where. Elytra slightly wider than prothorax (E/P 1.20 and 
1.17), normally formed; humeri prominent but rounded; each 
elytron with three striae free at base; striae entire at apex (ex- 
cept on final declivity), well impressed, punctate or punctulate; 
intervals convex, 3rd 4-punctate on outer edge ; 8th carinate at 
base ; surface of intervals not distinctly punctulate. Inner wings 
fully developed. Lower surface: proepisterna rugulose but not 
punctate; sides of abdomen shagreened but not distinctly punc- 
tate; last ventral segment lightly shagreened (with close isodia- 
metric microsculpture) but not punctate, with two apical setae 
on each side wide apart. Legs: front femur stout (fig. 39) ; front 
tibiae strongly 3-dentate, with 4th (upper) tooth weakly devel- 
oped; middle tibiae (fig. 35) with spur about as long as tibial 
width on outer side at or above 14 from apex. Measurements 
(of types): length 6.3-6.9; width 1.7-1.9 mm. 

Types. Holotype 9 (M.C.Z. No. 30,168) and 4 paratypes 
from Dobodura, Papua, Mar. -July 1944 (Darlington), taken in 
wet places. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 393 

Other material. Papua: 1, Upper Fly R., Oroville, Aug. 10- 
12, 1936 (Archbold Exped., A. M. N. H.) ; 2, Kiunga, Fly R., 
Sept. 24-25, 1957 (W. W. Brandt, Bishop Mus.). N-E. N. G. : 1, 
Nadzab, July 1944 (Darlington) ; 12, Aitape, Aug. 1944 (Darl- 
ington). Neth. N. G. : 1, Hollandia, May 1945 (Hoogstraal, 
M.C.Z.) ; 1, Hollandia area, W. Sentani, Cyclops Mts., 150-250 
m. (c. 500-800 ft.), June 25, 1959 (T. C. Maa, Bishop Mus.) in 
M. V. light trap. Also 3, Cape Gloucester, New Britain, Jan.- 
Feb. 1944 (Darlington). 

Measured specimens. The 9 holotype and 1 9 paratype. 

Notes. Distinguishing characters of inopaca are given in the 
key to Clivina of New Guinea. The species is similar to some 
Australian species of the australasiae group and does itself occur 
in Australia (Cape York Peninsula) but does not seem to have 
been described there. It is also similar to csikii (above) but is 
larger, relatively broader, and less convex. 

The Aitape specimens vary so much that I tried to separate 
some of them as different species characterized by more convex 
form, shallower notches between clypeal wings and preocular 
lobes, and more coarsely punctate elytral striae, but these char- 
acters failed to hold even in the series from Aitape. Proportions 
and measurements of specimens from Aitape (H/P .72 and .71; 
prothoracic W/L 1.02 and 1.00 and B/A 1.38 and 1.37; and 
E/P 1.14 and 1.17; length 5.5-7.3, width 1.5-2.0 mm.) are not 
significantly different from those of the types. 

This species lives in damp ground in swamps and by standing 
water. 

Clivina brandti n. sp. 

Description. Form and convexity about average; black or 
reddish pieeous, suture not or not much paler, appendages 
browner; moderately shining, reticulate microsculpture faint or 
absent above except at sides and base of pronotum and sides 
and apex of elytra. Head .70 and .72 width prothorax; eyes 
moderately large and prominent, genae forming c. right angles 
with sides of neck; antennae normal, intermediate segments 
about as wide as long; mandibles short; labrum 7 -setose ; clypeus 
subtruncate, usually broadly (sometimes slightly irregularly) 
emarginate, middle part not separated from wings; wings not 
or not much advanced, separated from preocular lobes by 
notches; clypeal suture indistinct (at most indicated by rather 
poorly defined transverse impression) ; supraocular convexities 



394 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY 

separated from preocular lobes by well defined impressions ; 
frontal sulci long - , straight and parallel between eyes, sinuous 
anteriorly ; front irregularly but usually distinctly and some- 
what longitudinally impressed, with some coarse punctures at 
middle and finer scattered punctures (but somewhat variable 
in punctatioiO ; neck constriction moderate, rather coarsely 
punctate, usually not interrupted at middle. Protltorax slightly 
longer than wide (L/W 1.03 and 1.04), moderately narrowed 
anteriorly, slightly so posteriorly (base/apex 1.38 and 1.32) ; 
sides nearly straight for much of length except subsinuate be- 
fore middle ; apex subtruncate or slightly emarginate ; posterior 
angles very obtuse, bluntly denticulate ; disc with usual im- 
pressed lines, punctulate, with line of coarser confluent punc- 
tures each side near base. Elytra slightly wider than prothorax 
(E/P 1.08 and 1.12); humeri prominent but rounded; each 
elytron with 3 striae free at base; striae entire (except normally 
abbreviated at apex ) , well impressed, punctulate ; intervals 
convex, finely and inconspicuously punctulate, 3rd 4-punctate 
on outer edge, 8th long-carinate at base. Inner wings fully de- 
veloped. Lower surface: proepisterna microreticulate, longitud- 
inally rugose internally, but hardly punctate ; abdomen micro- 
reticulate but not punctate or at most with last segment faintly 
subpunctate ; last ventral with 2 apical setae on each side wide 
apart. Legs: front femora moderately stout ; front tibiae strongly 
4-dentate but upper tooth of course small ; front trochanters more 
or less acute and prominent at apex; middle tibiae (fig. 37) with 
spurs about Vi from apex longer than tibial width. Measure- 
ments: length 6.5-7.0; width 1.7-1.9 mm. 

Types. Holotype (sex not determined) (Bishop Mus.) and 
29 paratypes from Kiunga, Fly R., Papua, various dates in 
July, Aug., Sept., and Oct., 1957 (W. W. Brandt). Some para- 
types now in M.C.Z. (No. 30,308). 

Other material. With the types of brandti at Kiunga, Brandt 
collected also 3 specimens with sides and apex of abdomen punc- 
tate (sansapor), 3 with (partly decomposed) reticulate micro- 
sculpture on disc of elytra (szckessyi), and 2 larger, smoother 
individuals without discal microsculpture (inopaca). Whether 
these actually represent different species or whether they are 
extreme variations of one species I cannot be sure — this is one 
of those difficult cases in which a taxonomist can only make a 
tentative classification and hope that more material will solve 
the problem. In the meantime I have listed the specimens in 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 395 

question under the species indicated. 

Measured specimens. The holotype and 1 paratype. 

Notes. Althoug-h this species clearly falls in the australasiae 
group of Clivina (see couplet 14 of key to species of genus), the 
rather strong punctation of head and unusually long spur of 
middle tibia suggest a possible relationship with puncticeps too. 

Clivina szekessyi Kult 

Kult 1951, Acta Soc. Ent, Czechoslovakia 48, p. 30. 

Description (of type). About average for genus in form and 
convexity; nearly black, suture (anteriorly) and edges of elytra 
slightly reddish, and head and pronotum not quite black (slight- 
ly reddish) in strong light, appendages reddish brown; front 
shining, not distinctly microreticulate, discs of pronotum and 
elytra entirely microreticulate or at least covered with rather 
close-set minute impressed lines probably representing slightly 
decomposed microreticulation. Head .70 width prothorax; eyes 
moderately large and prominent, genae short, forming c. right 
angles with sides of neck; antennae normal, median segments 
about as wide as long (or just longer than wide) ; mandibles 
short; labrum 7-setose; clypeus very broadly emarginate, wings 
not separated from median part and not advanced but sepa- 
rated from preocular lobes by nearly rectangular notches; pre- 
ocular lobes with outer edges oblique for much of length ; clypeal 
suture indistinct ; preocular lobes separated from supraocular 
convexities by well defined impressions; frontal sulci parallel 
posteriorly, sinuous anteriorly; front slightly irregular, sub- 
foveate at middle, irregularly punctulate ; neck impression mod- 
erate, punctate at sides, rather narrowly interrupted at middle. 
Prothorax: width/length .99; base/apex 1.42; sides faintly sin- 
uate before middle, slightly and sparsely crenulate ; apex sub- 
truncate (broadly emarginate) ; posterior angles broadly 
rounded, subdentate ; disc with usual impressed lines, with some 
transverse strigulation, not distinctly punctate but with im- 
pressed line each side before base. Elytra slightly wider than 
prothorax (E/P 1.14), normally formed; humeri broadly 
rounded but prominent ; each elytron with 3 striae free at base ; 
striae well impressed, virtually entire, finely punctate ; intervals 
somewhat convex, 3rd with 4 moderately distinct dorsal punc- 
tures on outer edge, 8th carinate at base, surface of intervals 
with microsculpture as described but not otherwise punctate. 



396 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Inner ivings fully developed. Lower surface : proepisterna micro- 
reticulate externally, rather lightly punctate or rugulose intern- 
ally ; ventral segments microreticulate but not punctate ; last 
ventral with 2 setae each side wide apart. Legs: front femora 
moderately stout, with lower edges slightly arcuate but not 
strongly lobed ; front tibiae strongly 3-dentate externally with 
4th (upper) tooth indicated as a rectangular projection; middle 
tibia with spur longer than tibial width and about y± from 
apex. Measurements: length 7.0; width 2.0 mm. (almost exact). 

Types. Holotype 9 (Hungarian National Mus.) from Sim- 
bang, Huon Gulf, N-E. N. G., 1899 (Biro) ; and 1 9 paratype 
(Kult collection) from Fenichel (also apparently collected by 
Biro in New Guinea, but I have not found the locality). 

Measured specimen. The holotype. 

Occurrence in New Guinea. I tentatively assign here the fol- 
lowing specimens. Neth. N. G. : 1, Hollandia, July-Sept, 1944 
(Darlington) ; 7, same locality, Apr., May, and June 1945 (Mal- 
kin, U.S.N.M.) ; 1, same locality, May 1945 (Hoogstraal, 
M.C.Z.) ; 2, Waris, S. of Hollandia, 450-500 m. (c. 1450-1625 
ft.), Aug. 1-2, 1959 (T. C. Maa, Bishop Mus.) ; 3, Sabron, Cy- 
clops Mts., 930 and 1,200 ft., May and June 1936 (Cheesman). 
N-E. N. G.: 1, Nadzab, July 1944 (Darlington); 1, Torricelli 
Mts., Siaute, sea level, Sept. 9-17, 1958 (W. W. Brandt, Bishop 
Mus.). Papua: 35, Dobodura, Mar.-July 1944 (Darlington); 3. 
Kiunga, Fly R., July 11-14, 26-30, Aug. 1-3, 1957 (W. \V. 
Brandt, Bishop Mus.). 

Notes. The specimens listed above vary somewhat in color 
(black or brown) and other minor characters, but I cannot dis- 
tinguish more than one species. Most of the Dobodura specimens 
were taken in shady, grassy ground by standing water. I am 
indebted to Dr. Z. Kaszab for an opportunity of examining the 
type of this species. 

Clivina netolitzkyi Kult 

Kult 1951, Acta Soc. Ent, Czechoslovakia 48, p. 30. 

Description. Kult indicates that this species has the same 
characters as the preceding one except color brownish with 1 
external interval of each elytron piceous and antennae and legs 
reddish ; elytra with striae less distinctly punctate, intervals 
less convex, 3rd with 4 dorsal punctures more distinct ; and 
front tibia with upper tooth indistinct. Length 6.9 mm. 



DARLINGTON : CARABID BEETLES OP NEW GUINEA 397 

Type. Holotype $ (Knit collection) from Erima, Astrolabe 
Bay, N-E. N. G." 

Notes. I cannot judge, from the description, whether or not 
this species is really distinct from the preceding one. 

(Subfam. MOKMOLYCINAE) 

(Tribe MORMOLYCINI) 

(Genus MORMOLYCE Hagenbach) 

Hagenbach 1825, Mormolyce Novum Coleopterorum Genus, p. 3. 
Rousseau 1906, in Wytsman, Genera Inseetorum, Fasc. 40, p. 3. 
Csiki 1928, Coleop. Cat., Carabidae, Mormolycinae, p. 1. 
Andrewes 1930, Cat. Indian Carabidae, p. 222 (see for additional refer- 
ences). 

1941, Ann. Mag. Nat. Hist. (11) 7, p. 315. 

Notes. This genus of 5 or 6 species of very large, extraor- 
dinarily flattened and expanded Carabidae occurs in the Malay 
Peninsula and (southern?) Thailand, and Sumatra, Java, and 
Borneo. It has been doubtfully recorded from New Guinea by 
Rousseau on the authority of Ritsema, but the record is pre- 
sumably an error. 

Subfam. HARPALINAE 
(Tribe APOTOMINI) 

(Genus APOTOMUS Illiger) 

Illiger 1807, Magazin fiir Insektenkunde 6, p. 348. 

Csiki 1928, Coleop. Cat., Carabidae, Harpalinae 1, p. 5 (see for additional 

references). 
Andrewes 1930, Cat. Indian Carabidae, p. 32. 
— ■ — • 1935, Fauna British India etc., Coleop., Carabidae 2, p. 29. 
Jeannel 1946, Coleop. Carabique de la Region Malgache, Part 1, p. 316. 

Diagnosis. See Andrewes ' key to tribes (1935, pp. Iff.); 
Apotomus is the only genus of its tribe. The species are small 
(usually 3 to 4 mm.), black or brown, pubescent, with peduncu- 
late prothorax and fully developed inner wings. 

Description. See Andrewes (1935). 

Genotype. Scarites rufus Rossi (Mediterranean region). 

Generic distribution. The Mediterranean region, parts of 
Africa, Madagascar, tropical Asia and islands to Celebes and 
Philippines, and Australia, but perhaps not New Guinea. 



398 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

(Apotomus atripennis Motschulsky) 

Motschulsky 1858, Etude Ent. 7, p. 22. 

Andrewes 1935, Fauna British India etc., Coleop. Carabidae 2, p. 30 (see 
for synonymy and additional references). 

Description. None needed here; see Andrewes (loc. cit.). 

Type. From near Colombo, Ceylon ; in Moscow University 
Mus. (t. Andrewes). 

Occurrence in New Guinea. Doubtful. 

Notes. A. atripennis is widely distributed in tropical Asia 
and extends east to Celebes and the Philippines. It has been 
recorded from New Guinea, but I doubt its occurrence there : 
the only supposedly New Guinean specimens that I have seen 
are from Wallace's dubious "Dorey" collections (see p. 331). 
In Luzon, this species occurs in open, grassy country, on or in 
the surface of the ground. It is sometimes common in flood 
debris and at light. 

Tribe BEMBIDIINI 

Osiki 1928, Coleop. Cat., Carabidae, Harpalinae 1, p. 27 (see for synonyms 

and additional references). 
Sloane 1921, Proc. Linn. Soe. New South Wales 46, p. 192 (Australian 

genera). 
Andrewes 1935, Fauna British India etc., Coleop., Carabidae 2, p. 80. 
Bembidiitae Auct. inch Jeannel 1916, Coleop. Carabiques de la Begion Mal- 

gache, Part 1, p. 331. 
This is a large tribe of small Carabidae most (but not all) of 
which live on the surface of the ground, often (but not always) 
by water or in wet places. Most are nocturnal and hide by day 
under vegetation or in ground litter or in loose soil or sand, but 
a few species live on tree trunks or are arboreal, and a few live 
deep in the soil, and a few are diurnal rather than nocturnal. 
For some reason, very few species of this tribe occur in caves, 
although many Trechini do so. 

The two principal genera of the tribe are both almost world- 
wide in distribution, but they are complementary in their main 
areas of abundance. Bembidion is dominant in cool northern 
regions, with comparatively few species scattered in the tropics 
and the southern hemisphere. Tachys is dominant in the tropics 
and some south-temperate regions, with comparatively few 
species in the cool north. (It should be added that some special- 
ists, including Jeannel, split both these old genera into many 
smaller ones.) 



DARLINGTON: CARABID BEETLES OF NEW GUINEA 399 

Key to Genera of Bembidiini of New Guinea 

Scutellar striae present; front tibiae with apices normal (irregularly 

rounded); length (in New Guinea) c. 4 mm. (p. 399).. Bembidion 

Scutellar striae absent ; front tibiae with outer apical angles obliquely 

truncate-emarginate ; length (in New Guinea) less than 4 mm 2 

Upper surface glabrous except for fixed setae (in New Guinean spe- 
cies) ; apical recurved stride of elytron usually (not always) well 

developed (p. 340) Tachys 

Upper surface with short pubescence; apical strides absent or rudi- 
mentary (p. 484) Limnastis 



Genus BEMBIDION Latreille 

Latreille 1802, Hist. Nat. Crustaces et Insectes 3, p. 82. 

Sloane 1921, Proc. Linn. Soc. New South Wales 46, p. 193 (the Australian 

species). 
Csiki 1928, Coleop. Cat., Carabidae, Harpalinae 1, p. 32 (see for synonyms 

and additional references). 
Andrewes 1930, Cat. Indian Carabidae, p. 38. 

1935, Fauna British India etc., Coleop., Carabidae 2, p. 92. 

Darlington 1960, Pacific Insects 1, pp. 332 ff. 

Diagnosis. See key (above), and Andrewes 1935. 

Description. See Andrewes 1935. 

Genotype. Cicindela quaelrimaculata Linnaeus (Holarctic). 

Generic distribution. See under tribe Bembidiini, above. A 
few Asiatic stocks of Bembidion reach Sumatra, Java, Borneo, 
Celebes, and the Philippines (Darlington 1960), some at low 
altitudes and others on mountains, and several species of the 
genus occur in ( chiefly southern ) Australia ; in fact one Asiatic 
species, B. sobrinum, is represented in Australia. But only one 
species of the genus has been found in New Guinea, and it be- 
longs to the specialized, coastal subgenus Cillenus (see notes un- 
der the following species). The apparent absence of Bembidion 
on the mountains of New Guinea (together with other evidence) 
suggests that these mountains have not been part of a route by 
which temperate Carabidae have dispersed between Asia and 
Australia. 

Bembidion (Cillenus) albertisi Putzeys 

Putzeys 1875, Ann. Mus. Civ. Genova (Genoa) 7, p. 748. 
Andrewes 1938, Proc. B. Ent. Soc. London (B) 7, p. 192. 
Darlington 1953, Coleopterists ' Bull. 7, p. 16. 



400 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Description. Rather narrow, subdepressed, dull (greenish?) 
black, each elytron with a small post-humeral f errugineous spot ; 
length 4 mm. ; see Andrewes for further details. 

Type. From Sorong, Neth. N. G., in Genoa Civic Mus. (t. 
Andrewes) ; I have not seen it. 

Occurrence in New Guinea. Known only from the type. 

Notes. The subgenus (or genus) Cillenus includes a moderate 
number of species scattered from Europe and Japan to Aus- 
tralia and New Zealand. Most occur on the sea coast, in the 
inter -tidal zone. The habits of albertisi are unknown, but a re- 
lated species (alatum Darl.), on Morotai Is. in the Moluccas, 
was found in gravel bars beside running fresh water near the 
sea. 

Genus TACHYS Stephens 

Stephens 1828, Illustrations of British Ent., Mandibulata 2, pp. 2, 4. 

Sloane, 1921, Proe. Linn. Soc. New South Wales 46, pp. 194 ff . 

Csiki 1928, Coleop. Cat., Carabidae, Harpalinae 1, p. 165 (see for synonymy 

and additional references). 
Andrewes 1925, Ann. Mus. Civ. Genova (Genoa) 51, pp. 327 ff. 

1930, Cat. Indian Carabidae, p. 323. 

1935, Fauna British India etc., Coleop. Carabidae 2, p. 206. 

Jeannel 1946, Coleop. Carabique de la Region Malgache, Part 1, p. 334. 

Diagnosis. See key (above), and Andrewes 1935. 

Description. See Andrewes 1935. Technical characters of 
Tachys are: size small, palpi subulate (that is with last segment 
reduced to vestige ; this character separates Tachys etc. from all 
other more or less similar small Carabidae in New Guinea) ; 
outer angle front tibia obliquely truncate-emarginate ; scutellar 
strioles absent; apical recurved strioles of elytra usually (not 
always) present; and (in New Guinean species) dorsal pubes- 
cence (excepting fixed tactile setae) absent. Other characters, 
perhaps less important for practical purposes, include mandibles 
short, curved, without setae in scrobes; 2 supraocular and 2 
lateral prothoracic setae present on each side (in all New 
Guinean species) ; wings usually fully developed, but reduced 
or dimorphic in a few species (see serrula subspecies inales, and 
the species ambulatus, avius, and orachys in the following 
pages) ; dorsal reticulate microsculpture present or absent, when 
present usually isodiametric on head, transverse on pronotum 
and elytra ; lower surface usually with prosternum more or less 
impressed longitudinally and metasternum usually (not always) 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 401 

variably margined anteriorly between middle coxae (see An- 
drewes 1925, p. 335) ; $ anterior tarsi dilated or not dilated, if 
dilated with 2 segments of each tarsus widened (only 1 seg- 
ment widened in some Oriental species), the widened segments 
biseriately squamulose below ; and $ with 1,9 2 setae each side 
last ventral segment, the position of the inner setae in 9 differ- 
ing in different groups. 

Genotype. Tachys Scutellaria Stephens (Europe). 

Generic distribution. Almost world-wide (see under tribe 
Bembidiini, above). 

Notes. There are good structural characters by which Tacliys 
can be divided into subgenera or genera (whichever are more 
useful) and some authors, including Jeannel, have divided it. 
This is probably the right course in the end, but I am not ready 
to follow it with the New Guinean species. The Tacliys of New 
Guinea ought to be classified on the same system as those of 
adjacent areas. Classifications have been made of the Tachys of 
Australia (Sloane 1921) and the Orient (Andrewes 1925). How- 
ever, these two classifications are very different and not easily 
reconciled with each other. I have chosen to follow Andrewes' 
system because it is more comprehensive than Sloane 's and be- 
cause most New Guinean species fit into it well. Some species 
of Tachys are very wide-ranging (see following paragraph). 
This suggests that Tachys, or at least some Tachys, have dis- 
persed comparatively recently. Most species of the genus are 
fully winged, and their small size probably makes them espe- 
cially liable to wind dispersal. They are sometimes carried by 
man : I have before me 3 widely distributed Oriental-New Guin- 
ean species intercepted in plant material shipped to the United 
States (see under T. truncatus, ceylanicus, and fumicatus, be- 
low). 

I recognize 63 full species of Tachys in New Guinea (a few 
others are of doubtful occurrence) belonging to 10 species- 
groups, of which the distribution is outlined as follows. The 
fasciatus and politus groups are more or less world-wide, and 
each includes very wide-ranging species as well as many more- 
localized ones : T. fasciatus apparently occurs throughout the 
whole warmer part of the Old World, and T. ceylanicus {politus 
group) extends from tropical Asia at least to New Guinea. Both 
these groups include subgroups that seem to be in process of 
speciation on New Guinea. The species of these groups occur 
as a rule on the ground in a variety of wet places, although an 



402 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

occasional species has entered some special habitat : e.g. T. aeneus 
(politics group) occurs in rather dry places away from water. 
The quadrillion group is probably nearly world-wide in distrib- 
ution but lives especially in coastal or saline habitats although 
some species occur elsewhere. The species of this group are few 
but often wide-ranging. Two of them apparently extend from 
southern Asia to New Guinea or Australia. The nanus group 
is probably nearly world-wide too, in a different special habitat, 
on or under the bark of trees, although one very distinct New 
Guinean species (wallacci) has invaded the lower foliage of 
rain forest. Some species of this group too are wide-ranging : 
e.g. umbrosus extends from southern Asia to New Guinea and 
has apparent close relatives in temperate Eurasia and North 
America and in eastern Australia. The fumicatus group may 
consist of only three species (unless additional ones occur in 
Africa), but fumicatus itself extends from Africa to Japan to 
New Guinea, and the other two (probably closely related) spe- 
cies are in Europe and Australia. The haliploidcs group occurs 
at least from Europe to Australia ; the single species that surely 
occurs in New Guinea (latissimus) apparently ranges from 
southern Asia to Australia. The truncatus group is another 
widely distributed one. It consists of a few, small species, some 
widely distributed (truncatus extends from southern Asia to New 
Guinea and has apparent close relatives in Europe and Aus- 
tralia), some apparently localized. In contrast to the preceding 
ones, the serra group, of three distinct species plus additional 
subspecies, is almost confined to New Guinea, extending only 
(so far as known) to the Moluccas and New Britain (not Aus- 
tralia) ; it has probably evolved on New Guinea from an Oriental 
ancestor. The acaroides group consists of a few obscure species, 
too little known to be significant. Finally, the singularis group, 
also very poorly known, presents an extraordinary geographical 
problem. T. singularis is described from Celebes. The only other 
known species of the group, yunax, occurs in New Guinea and 
the West Indies (see p. 482). In summary, I should say that the 
geographical relationships of New Guinean Tachys are complex, 
but are more Oriental than Australian. New Guinea and Aus- 
tralia do share some species, and a few of these may be Aus- 
tralian in origin, but most of them seem to be Oriental species 
or members of Oriental groups that have reached New Guinea 
from Asia rather recently and have continued to Australia. 
There are a number of very peculiar species or species-groups 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 403 

of Tachys in Australia that are not represented in New Guinea. 
Collecting Tachys is a task for specialists. Even good general 
collectors, like Toxopeus and Miss Cheesman, get few of them. 
Biro, however, did get a long series of one obscure species as 
well as a few specimens of other species, probably by sifting. 
Different species of Tachys have to be looked for laboriously in 
a great variety of habitats: by "treading" in many different 
sorts of fresh and saline wet places (and a few occur in dry 
places, too), by washing the banks of large and small streams 
and of pools and ponds, in and out of shade, and by "drowning" 
or sifting various sorts of debris, loose soil, and leaf litter espe- 
cially from damp places including rain forest; and additional 
species occur on fallen logs and under bark in rain forest, per- 
haps on fungi, or (Tachys wallacei) on low foliage or moss-like 
epiphytes in rain forest. A few hours collecting Tachys along 
the edge of a good flood, as it rises or soon afterward, is likely 
to be worth days of ordinary collecting, although the exact habi- 
tats of flood-collected specimens are often doubtful ; and some 
Tachys fly to light. 

Key to Groups of Tachys of New Guinea 

1. Mentum with 2 conspicuous t'oveae at base (except in minute truncatus, 

p. 431) 2 

— Mentum without conspicuous f oveae 5 

2. Apical elytral stride lacking or, if present, not extending forward as 

far as posterior dorsal elytral punture (p. 430) truncatus group 

— Apical striole present, the posterior elytral puncture on or behind its 

hooked anterior end 3 

3. Elytral margins strongly serrate behind humeri; striation of elytra en- 

tire or nearly so (p. 404) . . . .srrra group 

— Elytral margins not serrate (in New Guinean species) ; striation usually 

reduced 4 

4. Posterior dorsal puncture of elytron inside apical striole behind (not 

attached to) its hooked tip (fig. 40); anterior dorsal puncture 
usually outside 3rd stria (p. 410) fasciatus group 

— Posterior dorsal puncture of elytron attached to hooked tip of apical 

striole (fig. 41) ; anterior puncture on (position of) 3rd stria (and 
farther back than in fasciatus group) (p. 427) . . . quadrillum group 

5. Anterior part of apical striole about half way between suture and 

margin of elytron; elytral margin not distinctly setulose or serrate (i 

— Apical striole close to margin, or obsolete; elytral margin setulose or 

serrate 8 

6. Elytron with 2 dorsal punctures 7 

— Elytron with 1 dorsal puncture; (basal sulcus of pronotum with 2 



404 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

foveae at middle; claws denticulate; form very compact and con- 
vex) (p. 473) haliploides group 

7. Frontal foveae not extending onto clypeus; labrum truncate or moder- 

ately emarginate (p. 435) politus (inel. cxaratus) group 

— Frontal foveae long, linear, deep, converging anteriorly, extending onto 

and nearly meeting on clypeus; labrum semi-circularly emarginate 
(p. 468) / it mica t us group 

8. Apical stride close to margin of elytron ; stria 8 present 9 

— Apical stride and stria 8 obsolete (p. 472) acaroides group 

9. Elytral margins setulose but not strongly serrate (p. 476) 

nanus group 

— Elytral margins strongly serrate (p. 481) singularis group 

Tachys serra Group 

Tachys serra and its relatives do not fit any of Andrewes' 
groups of Oriental Tachys. In Andrewes' key (1925, pp. 336 ff.) 
they fall between the recurvicollis and triangularis (=fasciatus) 
groups, combining (with conspicuous pores on the mentum) an- 
tennal segments 2 and 3 subequal with elytral margins strongly 
serrate, and they differ from both groups in having stria 8 of 
the elytra deeply impressed for its whole length. However they 
resemble T. dclicatus Andrewes, of Singapore, in having the 
elytra almost fully striate (but stria 8 is obsolete in front in 
delicatus) , in position of dorsal punctures of elytra, in serration 
of elytral margins, and in some other ways. The serra group 
may therefore be derived from a dclicatus-like, presumably 
Oriental member of the fasciatus group. The serra group con- 
sists of 3 full species, which occur together in eastern New 
Guinea (at Dobodura), and several distinct geographical forms 
in other parts of New Guinea which I am calling subspecies to 
emphasize their relationships. Outside New Guinea, the group 
is represented by a relative of serra on Morotai Island in the 
Moluccas, and serrula extends to New Britain. The group is not 
represented in Australia and has no close relatives there. The 
species of the group are much alike in structure and differ 
chiefly in size, proportions, and shape of prothorax. I shall 
therefore describe only one species (serra) in detail, and shall 
compare the other species with it. All the species are found 
in debris, loose soil, etc. on the ground in damp places in rain 
forest. They are not primarily associated with open water but 
sometimes occur in rotten stumps etc. standing in water in 
swampy places. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 405 

Key to New Guinean Species of Tachijs of serra Group 

1. Broad, prothoracic width/length c. 1.39-1.57; large, length 3.0-3.6 

(rarely only 2.8) mm serra 

(la) Base of prothorax relatively narrow (base of prothorax/width 

of head c. 1.40-1.45), and sides of prothorax broadly and 

rather strongly sinuate before base (p. 405) serra sensu strictn 

(lb) Base of prothorax very broad (base of prothorax/width of 

head c. 1.70-1.80), and sides of prothorax broadly but less 

strongly sinuate (p. 407) (subsp. latiserra) 

(lc) Base of prothorax/width of head c. 1.51-1.53; sides of pro- 
thorax briefly sinuate near base (p. 407) .... (subsp. breviserra) 
— ■ More slender and/or smaller 2 

2. Eelatively slender, prothoracic width/length c. 1.35-1.37, base of pro- 

thorax/width of head c. 1.26-1.31; length 2.5-2.7 mm. (p. 408) 

tenuiserra 
— Less slender but much smaller ; prothoracic width/length c. 1.37-1.46 ; 

length 1.9-2.4 mm serrula 

(-a) Base of prothorax wider (base of prothorax/width of head 

e. 1.39-1.40) ; inner wings fully developed (p. 408) 

serrula sensu stricto 

(-b) Base of prothorax narrower (base of prothorax/width of head 

c. 1.31-1.33) ; inner wings reduced (p. 409) .... (subsp. inales 

TACHYS SERRA 11. Sp. 

Description. Form of fasciatus group but more convex than 
usual; brown (or reddish or castaneous), appendages paler; 
slightly iridescent ; microsculpture distinct and isodiametric or 
slightly transverse on head, absent or faint (transverse where 
visible) on disc of pronotum, absent or faint (sometimes faint 
indications of very fine transverse lines) on elytra. Head rather 
small, .62, .61, and .59 width prothorax; eyes rather small (com- 
pared to fasciatus) but convex, with genae behind them short, 
oblique ; antennae normal, segments 2 and 3 subequal, median 
segments almost 3X long as wide; frontal grooves normal, 
irregularly subparallel; mentum toothed, with 2 conspicuous 
pores at base. Prothorax transverse-subcordate ; width/length 
1.39, 1.40, and 1.43; base somewhat narrower than widest part 
but much wider than apex, base/apex 1.40, 1.40, and 1.41, width 
of base/width of head 1.40, 1.44, and 1.45 ; sides rather strongly 
rounded anteriorly, broadly and rather strongly sinuate before 
base; apex broadly emarginate; base almost subtruncate (actu- 
ally broadly but slightly lobed at middle, trending slightly back- 
ward at sides) ; anterior angles rounded; posterior angles acute, 



406 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

not carinate ; lateral margins rather narrow anteriorly, merging 
with disc posteriorly but not elevated posteriorly, not forming 
baso-lateral foveae (in fasciatus etc. the prothoracic margins 
are more elevated posteriorly so that rather vague baso-lateral 
foveae are formed) ; disc with anterior transverse impression al- 
most obsolete, median line distinct, basal transverse sulcus deep, 
punctate but without special median foveae. Elytra broad, 
width elytra/width prothorax 1.39, 1.37, and 1.34; humeri prom- 
inent, almost (obtusely) angulate; basal margin ending opposite 
or inside of end of 4th stria ; margin conspicuously dentate at 
and behind humeri, the dentations becoming less conspicuous 
posteriorly; striation entire (except most striae more or less 
abbreviated at base and apex); striae punctulate; 8th entire, 
deep anteriorly, deep and sinuous posteriorly, less deep but 
still well impressed at middle ; apical striole deep, hooked anter- 
iorly between ends 3rd and 4th discal striae; intervals slightly 
convex, with a few faint, scattered punctules ; anterior dorsal 
puncture on inner edge 5th interval (just outside 4th stria) 
about 14 from base, posterior puncture inside apical striole just 
behind hook. Inner wings fully developed. Lower surface: pres- 
ternum longitudinally impressed ; metasternum narrowly mar- 
gined anteriorly (between middle coxae) ; lower surface mostly 
impunctate, not pubescent (except last ventral segment of 9 ). 
Legs normal, claws simple. Secondary sexual characters normal ; 
$ with 2 segments each front tarsus very widely dilated (much 
wider than in fasciatus) ; $ with 1, $ 2 setae each side last ven- 
tral, the inner pair in 9 far forward (distant from margin) ; 
and 9 with traces of pubescence on last ventral segment so short, 
sparse, and inconspicuous that it is easily overlooked even at 
100X. Measurements: length c. 3.2-3.6; width c. 1.5-1.7 mm. 

Types. Holotype $ (M.C.Z. No 30,169) and 14 paratypes 
from Dobodura, Papua, Mar. -July 1944 (Darlington) ; and 1 
additional $ paratype from Oro Bay (near Dobodura), Dec. 
1943-Jan. 1944 (Darlington) ; and 1, Brown R., Papua, May 25, 
1956 (E. J. Ford Jr., Bishop Mus.). 

Measured specimens. The $ holotype and 2 ( S 9 ) paratypes 
from Dobodura. 

Notes. This very distinct new species represents a new sub- 
group of Tachys characteristic of New Guinea and some ad- 
jacent islands. Its relationships to other Tachys are discussed 
above and indicated in preceding keys. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 407 

Tachys serra latiserra n. subsp. 

Description. Generally similar to typical serra but differing 
in form (fig. 42) and proportions, especially in relatively wider 
base and less sinuate sides of prothorax. Head .55, .56, .56, and 
.55 width prothorax. Prothorax: width/length 1.57, 1.56, 1.52, 
and 1.57 ; base/apex 1.63, 1.58, 1.63, 1.63 ; base of prothorax/ - 
width of head 1.80, 1.70, 1.74, 1.75. Elytra: width elytra/width 
prothorax 1.33,, 1.31, 1.33, 1.35. Measurements: length 2.8-3.4; 
width 1.3-1.5 mm. 

Types. Holotype $ (M.C.Z. No. 30,170) and 28 paratypes 
all from Maffin Bay, Neth. N. G., Aug. 1944 (Darlington). 

Other material. Sixteen, Hollandia, Neth. N. G. July-Sept. 
1944 (Darlington) ; 1, same locality, Nov. 1944 (H. Hoogstraal, 
M.C.Z.) ; 6, Aitape, N-E, N. G., Aug. 1944( Darlington). These 
localities are all in the central part of the north coast of New 
Guinea. I have also before me one specimen from Madang 
("Friedrich-Wilh.-hafen," 1896, Biro, Hungarian National 
Mus.) intermediate both geographically and structurally be- 
tween serra and latiserra. Width of base of prothorax/width of 
head of this specimen is 1.58. 

Measured specimens. The <5 holotype and 1 9 paratype from 
Maffin Bay, 1 $ from Hollandia, and 1 $ from Aitape ; pro- 
portions listed in this order in each case. 

Notes. See key and preceding description. 

Tachys serra breviserra n. subsp. 

Description. Generally similar to typical serra and subspecies 
latiserra, but differing in form and proportions especially of 
prothorax. Head .58 and .59 width prothorax. Prothorax: 
width/length 1.53 and 1.54; base/apex 1.45 and 1.47; width of 
base/width of head 1.53 and 1.51 ; sides rounded almost to base 
then briefly sinuate just before basal angles. Elytra: width 
elytra/width prothorax 1.29 and 1.28. Measurements: length 
3.0-3.3 ; width 1.2-1.3 mm. 

Types. Holotype £ (M.C.Z. No. 30,171) and 2 ( S 9 ) para- 
types all from Sansapor, Vogelkop, Neth. N. G., Aug. 1944 
(Darlington). 

Measured specimens. The $ holotype and 9 paratype. 

Notes. See key to species of serra group. 



408 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Tachys tenuiserra n. sp. 

Description. Generally similar to serra but smaller and more 
slender (fig. 43) ; color, faint iridescence, and microsculpture 
similar. Head .64 and .66 width prothorax; details as de- 
scribed for serra. Prothorax more narrowly subcordate and with 
relatively narrower base; width/length 1.37 and 1.35; base/apex 
1.29 and 1.24; width of base/width of head 1.31 and 1.26; sides 
arcuate anteriorly, broadly and strongly sinuate before acute 
posterior angles; other details as described for serra. Elytra: 
width elytra/ width por thorax 1.40 and 1.49; other details as in 
serra. Inner wings fully developed. Lower surface, legs, and 
secondary sexual characters as in serra. Measurements: length 
2.5-2.7 ; width 1.0-1.1 mm. 

Types. Holotype $ (M.C.Z. No. 30,172) and 28 paratypes all 
from Dobodura, Papua, Mar.-July 1944 (Darlington). 

Measured specimens. The $ holotype and 1 9 paratype. 

Notes. Under the microscope, serra and the present species are 
perfectly distinct, but I did not distinguish them in the field 
and cannot say whether they occur in exactly the same habitats. 

Tachys serrula n. sp. 

Description. Form of serra group (or convex fasciatus group) ; 
rufous, head (slightly) and elytral discs (more obviously) usu- 
ally darker, appendages testaceous; microsculpture light and 
somewhat transverse on head, not distinct on pronotum and 
elytra, although slight opalescent iridescence suggests unresolved 
microsculpture. Head small, .61 and .63 width prothorax ; eyes 
rather small, somewhat variable in size and convexity; genae 
oblique or convex in profile ; antennae normal, segments 2 and 
3 subequal, median segments about 2X long as wide ; frontal 
grooves irregularly subparallel ; mentum toothed, with 2 con- 
spicuous pores at base. Prothorax transverse-subcordate ; width/- 
length 1.46 and 1.46 ; moderately narrowed in front and behind ; 
base/apex 1.26 and 1.30; width of base/width of head 1.39 and 
1.40; sides arcuate in about anterior %, sinuate posteriorly; 
apex broadly emarginate, base sub truncate (modified as usual in 
group) ; anterior angles rounded, posterior angles c. right, 
sharp, not carinate ; lateral margins narrow, scarcely wider 
basally ; no distinct baso-lateral foveae ; disc with anterior trans- 
verse line obsolete, median line distinct, basal sulcus deep, irreg- 
ular or slightly punctulate but without special median foveae. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 409 

Elytra rather broad; width elytra/width prothorax 1.34 and 
1.39; humeri prominent but rounded; basal margin ending 
opposite or inside base of 4th stria; margin conspicuously den- 
tate at and behind humeri; striation entire (except most striae 
abbreviated anteriorly and posteriorly as usual) ; striae 1 to 7 
lightly impressed or represented by rows of moderate punctures ; 
stria 8 deep and entire ; apical stride as in serra; intervals 
nearly flat; anterior dorsal puncture on 4th stria at or behind 
anterior 14, posterior puncture inside apical striole behind hook. 
Inner wings fully developed in all specimens. Lower surface 
with slight pubescence on last ventral segment. Legs normal. 
Secondary sexual characters as in serra. Measurements: length 
2.0-2.4; width 0.8-1.0 mm. 

Types. Holotype $ (Hungarian National Mus.) and 150 
paratypes (some in M.C.Z., Type No. 30,173) from Madang 
("Friedrich-Wilh.-hafen' ? ), N-E. N. G., 1901 (Biro). These 
specimens were probably taken by sifting, but there is no indi- 
cation of the habitat. 

Other material. N-E. N. G. : 48, Stephansort, Astrolabe Bay, 
1898 (Biro) ; 12, Erima, Astrolabe Bay, 1896 (Biro) ; 4, Hanse- 
man, Astrolabe Bay, 1901 (Biro). Papua: 1, Karema, Brown 
R., Mar. 8-11, 1955 (E. O. Wilson, M.C.Z.), taken in lowland 
rainforest. Also 7, Cape Gloucester, New Britain, Jan. -Feb. 1944 
(Darlington). 

Measured specimens. The $ holotype and 1 5 paratype 
from Madang. 

Notes. See key to species of serra group. 

Tachys serrula inales n. subsp. 

Description. Similar to typical serrula but differing in pro- 
portions and in atrophy of wings. Head .62 and .65 width pro- 
thorax. Prothorax: width/length 1.41 and 1.37; base/apex 1.23 
and 1.22; width of base/width of head 1.33 and 1.31; sides 
arcuate anteriorly and sinuate posteriorly about as in typical 
serrula. Elytra: width elytra/width prothorax 1.33 and 1.40. 
Inner wings reduced in all specimens, c. y 2 as long as elytra. 
Measurements: length 1.9-2.3; width 0.8-0.9 mm. 

Types. Holotype $ (M.C.Z. No. 30,174) and 10 paratypes 
all from Dobodura, Papua, Mar.-July 1944 (Darlington). 

Measured specimens. The 6 holotype and 1 9 paratype. 

Notes. I have no record whether this third, smallest species 
of the serra group occurs in exactly the same habitat as the 
other species at Dobodura. 



410 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Tachys fasciatus Group 

The following are noteworthy characters of the fasciatus group 
of Tachys in New Guinea (the group is more diverse in some 
other parts of the world). Form somewhat variable but usually 
moderately broad and depressed ; color variable ; reticulate 
microsculpture variable on front of head (present or absent in 
different species), usually indistinct on discs of pronotum and 
elytra, but surface usually faintly iridescent or silky. Head: 
mentum 2-foveate, usually toothed (tooth individually variable) ; 
antennae with segment 2 usually slightly longer than 3 ; frontal 
grooves moderately impressed, subparallel between eyes, sinuous 
anteriorly. Prothorax usually transversely subcordate; apex sub- 
truncate or broadly emarginate (sometimes vaguely lobed at 
middle), base usually subtruncate or with weak lobe at middle 
but more strongly lobed in sublobatus; posterior angles vari- 
able, not carinate ; anterior transverse impression obsolete, 
middle line finely impressed, basal transverse sulcus deep but 
sometimes interrupted at middle, usually crenate, with or with- 
out a conspicuous fovea at middle. Elytra with humeri prom- 
inent but rounded (slightly narrowed in species with reduced 
wings) ; margins ending inwardly opposite bases of 4th striae, 
not serrate (margins are serrate in some Oriental species of 
group) ; striation variable, usually not entire; 8th stria widely 
interrupted at middle, the posterior part (behind the principal 
interruption) entire or nearly so in the larger species, frag- 
mentary in the smaller ones ; apical striole well developed, about 
midway between suture and lateral margin, more or less hooked 
at tip (anterior end) ; anterior dorsal puncture usually on or 
near 4th stria (near 3rd stria in privus and on 6th interval in 
sericcus, but latter may not occur in New Guinea), posterior 
puncture usually inside apical striole behind (not attached to) 
its hooked tip (fig. 40). Inner wings usually fully developed, but 
reduced in last 2 species. Lower surface impunctate ; prosternum 
usually slightly longitudinally impressed; mesosternum nar- 
rowly margined anteriorly. Secondary sexual characters: $ with 
2 segments each front tarsus dilated, squamulose below ; $ with 
1, 5 2 setae each side last ventral segment, the 2nd (inner) 
setae in 9 distant from the margin ; and 9 usually with a 
little short, sparse, very inconspicuous pubescence on last ven- 
tral segment. 

The fasciatus group of Tachys is nearly world-wide in dis- 
tribution, within the geographical limits of the genus. Tachys 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 411 

fasciatus itself occurs throughout most of the warmer part of 
the Old World. Apparent representative forms of Tachys mas- 
tersi occur in eastern Australia, New Guinea, the Moluccas, and 
the Philippines. The 11 other species of the group in New 
Guinea are endemic or extend only to the Moluccas or other 
islands near New Guinea, so far as is known. 

Key to Species of Tachys of fasciatus Group of New Guinea 

1. Anterior dorsal puncture of elytron not on or near 4th stria 2 

— Anterior dorsal puncture of elytron on or near 4th stria 3 

2. Anterior dorsal puncture of elytron on 6th interval (p. 412) (sericeus) 
— • Anterior dorsal puncture of elytron near (outside of) 3rd stria 

(p. 413) privus 

3. Basal transverse sulcus of pronotum with a distinct fovea at middle; 

and eyes large or moderate, genae (in profile from above) forming 
right or nearly right angles with neck; and elytra fasciate or macu- 
late, not unicolorous (except in teneral specimens) 4 

— Not as above in one or more ways: basal sulcus of pronotum without 

distinct median fovea (except individually in master si subsp.) ; eyes 
moderate or small, in latter case forming very obtuse angles with 
neck ; elytra often unicolorous, but sometimes fasciate or maculate. 9 

4. At least 6 dorsal striae indicated (outer ones sometimes very faint) 

on each elytron; front usually with reticulate microsculpture of 
entire meshes plainly visible at 50X (but faint in apex) 5 

— Less than 6 dorsal striae indicated on each elytron; front with reticu- 

late microsculpture faint or absent 7 

5. Larger, 3.1-3.3 mm.; elytra dark in c. basal % with relatively small 

post-humeral spots (sometimes almost lacking) and large apical area 
pale; (3rd and 4th elytral striae almost connected at anterior dorsal 
puncture) (p. 414) apex 

— Smaller, 2.2-2.8 mm.; elytra fasciate or 4-maculate with anterior and 

posterior pale areas e. equal 6 

6. Prothorax pale; elytral striae slightly more impressed (p. 414) 

fasciatus 

— Prothorax dark (as dark or darker than head) ; elytral striae usually 

slightly less impressed (p. 416) fumax 

7. Base of prothorax with broad but distinct lobe set off by strong sinua- 

tions (fig. 44) sublolatus 

(7a) Base of prothorax slightly narrower than width of head; ely- 
tron usually with only the sutural stria well impressed; elytra 

usually fasciate (p. 418) (sublobatus s. s.) 

(7b) Base of prothorax c. equal width of head; each elytron with 

2 striae well impressed; elytra 2-maculate (dark, each elytron 

with a pale subapieal spot) (p. 420) (subsp. suffusus) 

— Base of prothorax not abnormally lobed 8 



412 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 

8. Form and size of fasciatus, length 2.4-2.9 mm.; pronotum pale (usu- 

ally paler than head) (p. 417) sibling 

— ■ More slender and larger, 3.0-3.2 mm.; pronotum dark (or at least not 
paler than head) (p. 417) beatus 

9. Elytra broader, more convex, and with wider margins than usual in 

fasciatus group (fig. 45) (mastersi) 

(9a) Basal area of pronotum (behind posterior transverse sulcus) 
usually not rugulose; color dark castaneous, often (not always) 

2 — maculate with pale (p. 421) (subsp. pinguis) 

(9b) Basal area of pronotum longitudinally rugulose; color more 
rufous, not maculate (p. 420) (subsp. exul) 

— Elytra normal for fasciatus group in shape and convexity 10 

10. Eyes moderate; inner wings fully developed 11 

— Eyes small, the genae forming very obtuse angles with sides of neck; 

wings often (not always) reduced 12 

11. Male with 2 segments each front tarsus widely dilated, 2nd segment 

much wider than long masculus 

(11a) Elytra 4-maculate (p. 422) (masculus s. s.) 

(lib) Color nearly uniform brown or castaneous (p. 423) 

(subsp. filius) 

— Male with 2 segments each front tarsus moderately dilated, 2nd seg- 

ment c. wide as long (p. 424) flavax 

12. Inner wings fully developed; elytra fasciate (p. 424) luscus 

— ■ Inner wings usually reduced (c. % or % long as elytra) ; if wings 

fully developed (individually in avius), elytra not fasciate 13 

13. Sides of prothorax broadly and strongly sinuate; elyt'-a fasciate or 

4-maculate (p. 425) ambulatus 

— Sides of prothorax briefly sinuate; unicolorous (p. 426) avius 

(Tachys sericeus Motschulsky) 

Motschulsky 1851, Bull. Soc. Nat. Moscou 24, Part 2, No. 4, p. 507. 
Andrewes 1935, Fauna British India, Coleop., Carabidae 2, pp. 214, 223 

(see for additional references and synonymy). 
Louwerens 1953, Verhandlungen Naturforschenden Gesellschaft Basel 64, 
p. 305. 

Description (recognition characters only). A large Tachys 
(3.25-4.0 mm. long) of the fasciatus group; reddish or reddish- 
castaneous with head slightly darker and elytral disc nearly 
black; anterior dorsal puncture of elytron on 6th interval about 
V4 from base. 

Type. From "Ind. or."; said by Andrewes to be in Moscow 
University Mus. 

Occurrence in New Guinea. Doubtful. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 413 

Notes. The only specimen of sericeus that I have seen that 
purports to be from New Guinea is in the British Mus. labeled 
"Dorey, Wallace." I have already given reasons (p. 331) for 
doubting that Wallace's "Dorey" specimens really came from 
New Guinea. Otherwise the known range of sericeus is from 
northeastern India and Burma to Sumatra, Borneo, and Sumba 
in the Lesser Sunda Is. The species' closest allies are Oriental. 

Tachys privus n. sp. 

Description. With characters of fasciatus group as here 
defined. Form average for group ; reddish yellow, a transverse 
post-median elytral fascia slightly darker; shining, faintly 
iridescent or silky; head without distinct reticulate microsculp- 
ture on middle of front but traces of it laterally. Head .73 
width prothorax, eyes moderate (smaller and less prominent 
than in fasciatus), genae forming slightly obtuse (not quite 
right) angles with neck; antennae rather short, middle segments 
about 2X long as wide. Prothorax transversely subcordate ; 
width/length 1.51 ; base/apex 1.12 ; base/head 1.08 ; sides arcu- 
ate anteriorly, broadly and rather strongly sinuate posteriorly ; 
base and apex subtruncate ; posterior angles right and well de- 
fined ; basal transverse sulcus subcrenulate, with conspicuous 
longitudinal pore at middle. Elytra about 14 wider than pro- 
thorax (E/P 1.23), moderately elongate; 6 discal striae indi- 
cated on each elytron, inner ones moderately impressed, outer 
ones faint ; anterior dorsal puncture outside 3rd stria about % 
from base. Inner wings fully developed. Lower surface, legs, 
and secondary sexual characters (of $) normal; last ventral 
segment not pubescent. Measurements: length c. 2.8; width c. 
0.9 mm. 

Type. Holotype 9 (M.C.Z. No. 30,175) from Dobodura, 
Papua, Mar.- July 1944 (Darlington) ; unique. 

Measured specimen. The type. 

Notes. No other New Guinean species of the fasciatus group 
as here defined has the anterior dorsal puncture of the elytron 
so near the 3rd stria. In Andrewes' (1925) revision, this species 
would run to near zonatus Putzeys (of Celebes), but privus 
differs from this in a number of ways : sides of prothorax 
strongly sinuate (not sinuate in zonatus), basal sulcus with a 
conspicuous pore at middle (absent in zonatus), etc. 



414 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

Tachys APEX n. sp. 

Description. With characters of fasciatus group as here de- 
fined. Form normal for group ; head dark, prothorax pale, 
elytra dark in c. basal % with (usually) elongate posthumeral 
marks and (always) a broad apical area pale, appendages pale; 
moderately shining, slightly iridescent, microsculpture of head 
faint at middle, more distinct at sides. Head .79 and .79 width 
prothorax ; eyes large and prominent ; genae short, forming right 
angles with sides of neck ; antennae with median segments c. 
3X long as wide. Prothorax transversely subcordate ; width/- 
length 1.61 and 1.56; base/apex 1.12 and 1.06; base/head 1.01 
and .96 ; sides arcuate anteriorly, converging posteriorly to brief 
but rather strong basal sinuations; base and apex subtruncate; 
posterior angles c. right, well formed; basal sulcus deep, finely 
crenulate at sides, interrupted at middle by three elongate pores 
or sulci. Elytra about % wider than prothorax (E/P 1.31 and 
1.36), rather elongate; inner striae well impressed, outer ones 
including 6th and 7th faint; anterior dorsal puncture about % 
from base between 3rd and 4th striae but nearer line of 4th 
(both 3rd and 4th striae bent toward and attached to the punc- 
ture on both elytra of all specimens). Inner wings fully devel- 
oped. Lower surface, legs, and secondary sexual characters nor- 
mal ; $ with 2 segments each front tarsus moderately dilated ; 
last ventral segment not pubescent. Measurements: length c. 3.1- 
3.3 ; width 1.3-1.4 mm. 

Types. Holotype <5 (M.C.Z. No. 30,176) and 4 paratypes all 
from Nadzab, N-E. N. G., July 1944 (Darlington). 

Measured specimens. The $ holotype and 1 9 paratype. 

Notes. The large size and general appearance of the present 
new species suggest Tachys sericcus Motschulsky or venustus 
Andrewes of the Orient, but the position of the anterior dorsal 
puncture of the elytron indicates that the new species is in fact 
more related to fasciatus and its immediate allies. 

Tachys fasciatus (Motschulsky) 

Trechus fasciatus Motschulsky 1851, Bull. Soc. Nat. Moseou 24, Part 2, 

No. 4, p. 506. 
Tachys triangularis Nietner 1858, Ann. Mag. Nat. Hist. (3) 2, p. 422 

(B emMdium) . 
Sloane 1921, Proc. Linn. Soc. New South Wales 46, pp. 196 (elytra), 200 

(key), 207. 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 415 

Andrewes 1925, Ann. Mus. Civ. Genova (Genoa) 51, pp. 342 (key), 350, 

figs. 3, 14, 10, 22, 30, 34, 35, 38. 
Ta&hys fasciatus Auct. incl. Csiki 1928, Coleop. Cat., Carabidae, Harpa- 

linae 1, p. 178 (see for synonymy and additional references). 
Andrewes 1935, Fauna British India etc., Coleop., Carabidae 2, pp. 213 
(key), 217. 

Description. None required here. See key above and notes 
below. Proportions : head/prothorax .72, .73, .76 ; prothoracic 
width/length 1.58, 1.60, 1.61, base/apex 1.26, 1.25, 1.16; pro- 
thoracic base/width of head 1.13, 1.14, 1.06; width elytra/pro- 
thorax c. 1.32, 1.38, 1.42. Measurements: length c. 2.2-2.7; 
width c. 0.8-1.1 mm. 

Types. Of fasciatus, from "Ind. or.," in Moscow University 
Zool. Mus. ; of triangularis, from Ceylon, in Berlin University 
Zool. Mus. ; both seen by Andrewes. 

Occurrence in New Guinea. Papua.- 5, Milne Bay, Dec. 1943 
(Darlington) ; 38, Dobodura, Mar.- July 1944 (Darlington) ; 15, 
Oro Bay, Dec. 1943- Jan. 1944 (Darlington) ; 8, Port Moresby, 
Oct. 1944 (Darlington). N-E. N. G. : 6, Lae, Oct. 1944 (Dar- 
lington) ; 12, Nadzab, July 1944 (Darlington) ; 19, Chimbu 
Valley, Bismarck Range, 5,000-7,000 ft., Oct. 1944 (Darling- 
ton) ; 1, Finschhafen, May 12, 1944 (E. S. Ross, California 
Acad.); 6, Aitape, Aug. 1944 (Darlington). Neth. N. G. : 12, 
Hollandia, July-Sept. 1944 (Darlington), and 5, same locality, 
Nov. 1944 and Feb. 12 and May 1945 (H. Hoogstraal, Chicago 
Mus.) ; 10, Hollandia area, W. Sentani, Cyclops Mts., 50-100 m. 
(c. 150-325 ft.), June 22-24, 1959 (J. L. Gressitt, Bishop Mus.) ; 
5, Maffin Bay, Aug. 1944 (Darlington) and 3, same locality, 
Sept. 1944 (E. S. Ross, California Acad.) ; 7, Sansapor (Vogel- 
kop), Aug. 1944 (Darlington). Probably occurs throughout New 
Guinea at Ioav altitudes and to some extent in the mountains. 

Measured specimens. One $ 9 from Dobodura and 1 $ from 
Sansapor (proportions listed in this order in each case). 

Notes. This species occurs in parts of Africa, through southern 
Asia north to Japan, and south and east through the islands to 
Australia (south at least to southern New South Wales), and, 
east of New Guinea, I have seen it from Cape Gloucester, New 
Britain (Darlington), and Bougainville in the Solomons (A. B. 
Gurney, U.S.N.M.), and it is recorded from New Caledonia. 
It is common in a variety of wet, muddy, and grassy places in 
open country and sometimes also in forest. 

The typical form of fasciatus, which is apparently the only 
form in tropical Asia, is testaceous or ferrugineous with head 



416 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

darker and (except in unpigmented specimens) a somewhat 
variable dark fascia across the middle of the elytra, and the 
apices of the elytra are often slightly darkened too. In New 
Guinea, however, individuals occur (together with typically 
colored ones) in which the dark area of the elytra is more or 
less extended, and in extreme cases the elytra are black with 4 
pale blotches. The prothorax is always pale. 

Tachys fumax n. sp. 

Description. With characters of fasciatus group as here de- 
fined. Form broad; head reddish brown, pronotum slightly 
darker especially at sides, elytra testaceous with median fascia 
and apices dark, the dark color more or less produced along 
suture and margins (but actual margins pale), appendages pale; 
rather shining, faintly iridescent ; head with distinct isodia- 
metric microsculpture. Head .73 and .75 width prothorax; eyes 
large and prominent (as in fasciatus), genae very short and 
forming right angles with sides of neck ; antennae with middle 
segments c. 3X long as wide. Prothorax transversely sub- 
cordate; width/length 1.60 and 1.61; base/apex 1.20 and 1.16; 
base/head 1.09 and 1.04 sides arcuate anteriorly, broadly but 
somewhat variably sinuate posteriorly ; base and apex subtrun- 
cate ; posterior angles c. right ; basal sulcus crenulate laterally, 
interrupted at middle by a conspicuous longitudinal pore. 
Elytra wide, E/P 1.41 and 1.39 ; inner discal striae well im- 
pressed, outer ones including 6th faint, 7th not distinctly indi- 
cated ; anterior dorsal puncture just inside 4th stria c. % from 
base. Inner wings fully developed. Lower surface, legs, and 
secondary sexual characters normal ; $ with 2 segments each 
front tarsus rather widely dilated; last ventral 9 with a little 
very fine, short pubescence scarcely visible at c. 100X. Meas- 
urements: length 2.6-2.8; width 1.1-1.2 mm. 

Types. Holotype $ (M.C.Z. No. 30,177) and 12 paratypes all 
from Sansapor, Neth. N. G., Aug. 1944 (Darlington). 

Other material. Three, southern lowlands of Morotai Is., 
Moluccas, Sept. 1944 (Darlington). 

Measured specimens. The $ holotype and 1 $ paratype. 

Notes. This new species is so close to Tachys fasciatus that I 
at first thought it might be a Mendelian color form of that 
species. The two occur together, without intergradation, both at 
Sansapor and on Morotai Is. However, the color difference seems 
to be re-enforced by a slightly lighter striation of fumax, and I 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 417 

am therefore treating the latter as a separate species, although 
I am not sure of its status. 

Tachys sibling n. sp. 

Description. With characters of fasciatus group as here de- 
fined. Form normal for group ; yellow, head slightly browner, 
elytra with a median fascia and apices brown; shining, faintly 
iridescent ; microsculpture of front light, visible but not form- 
ing distinct complete isodiametric meshes. Head .74 and .76 
width prothorax ; eyes large and prominent, genae forming right 
angles with sides of neck ; antennae with median segments about 
3X long as wide. Prothorax transverse-subcordate ; width/- 
length 1.55 and 1.55; base/apex 1.16 and 1.19; base/head 1.06. 
and 1.05; sides arcuate anteriorly, sinuate (variably) poster- 
iorly ; base and apex subtruncate ; posterior angles c. right ; basal 
sulcus finely crenulate, interrupted at middle by a conspicuous 
longitudinal pore. Elytra rather wide; E/P 1.35 and 1.34; 
inner diseal striae impressed and irregularly subcrenulate, outer 
striae to 5th lighter, 6th not or scarcely indicated ; anterior 
dorsal puncture inside 4th stria about % from base. Inner 
wings fully developed. Lower surface, legs, and secondary sexual 
characters normal; $ with 2 segments each front tarsus moder- 
ately dilated; $ with last ventral segment finely and incon- 
spicuously pubescent. Measurements: length 2.4-2.9; width c. 
0.9-1.2 mm. 

Types. Holotype 6 (M.C.Z. No. 30,178) and 6 paratypes 
from Dobodura, Papua, Mar.-July 1944 (Darlington) ; and 2 
additional paratypes from Oro Bay, near Dobodura, Dec. 1943- 
Jan. 1944 (Darlington). 

Other material. One specimen from Nadzab, N-E. N. G., July 
1944 (Darlington) may represent this species or may be a 
related one. 

Measured specimens. The $ holotype and 1 9 paratype from 
Dobodura. 

Notes. This species is very close indeed to fasciatus, from 
which it differs chiefly as indicated in the preceding key. 

Tachys beatus n. sp. 

Description. With characters of fasciatus group as here de- 
fined. Slightly larger and more slender than average for group ; 
head and pronotum dark, elytra strikingly bicolored, yellow 



418 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

with broad transverse median fascia and apices blackish, the 
dark color extended along suture and margins, but actual mar- 
gins pale, appendages yellow ; shining, slightly iridescent ; front 
without distinct microsculpture. Head .73 and .75 width pro- 
thorax; eyes large and prominent, genae forming right angles 
with sides of neck; antennae with middle segments c. 3X (or 
slightly less) long as wide. Prothorax subcordate ; width/length 
1.50 and 1.49; base/apex 1.11 and 1.04; base/head 1.02 and 
0.96; sides arcuate anteriorly, rather strongly sinuate before 
base; base and apex subtruncate; posterior angles acute (slightly 
more pointed than right) ; basal sulcus crenulate, interrupted at 
middle by a conspicuous elongate pore. Elytra rather elongate ; 
about % wider than prothorax (E/P 1.34 and 1.31); 2 inner 
discal striae well impressed on each elytron, and 2 or 3 addi- 
tional striae lightly impressed or faintly indicated; anterior 
dorsal puncture at basal y 3 just inside 4th stria. Inner wings 
fully developed. Lower surface, legs, and secondary sexual char- 
acters normal ; $ with 2 segments each front tarsus moderately 
dilated; last ventral segment not distinctly pubescent or with 
very slight traces of pubescence in 9 . Measurements: length 
3.0-3.2; width 1.1-1.2 mm. 

Types. Holotype S (M.C.Z. No. 30,179) and 6 paratypes 
from Aitape, N-E. N. G., Aug. 1944 (Darlington) ; 4 paratypes 
from Sansapor, Neth. N. G., Aug. 1944 (Darlington) ; 1 para- 
type, Bernhard Camp, Neth. N. G., 50 m. (c. 150 ft.) altitude, 
July-Nov. 1938 (J. Olthof, Louwerens Coll.). 

Other material. Three, Dobodura Papua, Mar.-July 1944 
(Darlington); 2 Nadzab, N-E. N. G., July 1944 (Darlington). 
These specimens differ slightly from the types in size or propor- 
tions or color. 

Measured specimens. The $ holotype and 1 9 paratype from 
Aitape. 

Notes. This rather large and conspicuously marked Tachys 
is distinguished from related forms in the preceding key to 
species of the fasciatus group. 



Tachys sublobatus n. sp. 

Description. With characters of fasciatus group as here de- 
fined. Form as figured (fig. 44) ; slightly more slender than 
usual ; color rather variable, from piceous with elytra 4-plagiate 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 419 

to reddish yellow with suture and median fascia (together form- 
ing a broad cross) and apices of elytra dark brown; appendages 
pale, antennae sometimes slightly darker; shining, usually not 
distinctly iridescent but sometimes slightly so ; front without 
distinct reticulate microsculpture. Head .74 and .77 width pro- 
thorax ; eyes large and prominent, genae forming right angles 
with sides of neck ; antennae with middle segments about 3X 
long as wide ; frontal grooves more strongly impressed than 
usual anteriorly ; mentum with or without a tooth, 2-foveate at 
base as usual in group. Prothorax subcordate ; width/length 1.48 
and 1.47 ; base/apex 1.10 and 1.03 ; base/head 0.97 and 0.93 ; sides 
arcuate anteriorly, strongly and broadly sinuate posteriorly; 
apex subtruncate or slightly and broadly emarginate, base 
broadly and slightly lobed, lobe nearly truncate at middle but 
ending in rather strong (but brief) sinuations at ends of basal 
sulcus; posterior angles right or (usually) acute; basal sulcus 
crenulate, interrupted at middle by a conspicuous elongate pore. 
Elytra rather elongate ; about % or more wider than prothorax 
(E/P 1.33 and 1.40) ; usually only one (sutural) stria well 
impressed but second stria sometimes slightly impressed and 
one or two additional striae faintly indicated (but this is close 
to being a 1-striate species). Anterior dorsal puncture about 
Y 3 from base just inside position of (obsolete) 4th stria. Inner 
wings fully developed. Lower surface, legs, and secondary sex- 
ual characters normal; S with 2 first segments each front 
tarsus moderately dilated; 9 with last ventral with a little, 
scarcely detectable, short pubescence. Measurements: length 
2.2-2.8; width 0.8-1.1 mm. 

Types. Holotype <J (M.C.Z. No. 30,180) and 19 paratypes 
from Nadzab, N-E. N. G., July 1944 (Darlington) ; 6 paratypes 
from Lae, N-E. N. G., Oct. 1944 (Darlington) ; and 2 paratypes 
from Dobodura, Papua, March-July 1944 (Darlington). 

Measured specimens. The $ holotype and 1 9 paratype from 
Nadzab. 

Notes. This species probably represents Tachys ductus Put- 
zeys of Amboina, but differs (judging from Andrewes' redes- 
cription of ductus, 1925, Ann. Mus. Civ. Genova [Genoa] 51, p. 
357) in lacking distinct reticulate microsculpture on the head 
and probably in other details. Moreover, the following new 
subspecies occurs between areas inhabited by cinctus and sub- 
lobatus and differs from both. 



420 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

T\CHYS SUBLOBATUS SUFFUSUS 11. SUbsp. 

Description. Generally similar to typical sublobatus but 
stouter and darker. Color somewhat irregular reddish castan- 
eous with elytra 2-maculate, i.e. each elytron with a rather large 
subapical yellow spot ; appendages pale, except basal segments 
of antennae irregularly darker. Head .75 and .75 width pro- 
thorax. Prothorax relatively broader than in typical suMobatus 
and with relatively broader base; width/length 1.57 and 1.57; 
base/apex 1.16 and 1.14; base/head 1.03 and 1.02. Elytra 1.38 
and 1.37 width prothorax, about as in sublobatus (or slightly 
broader) except 2nd stria well impressed for part of its length 
on disc (rarely so in typical sublobatus) . Measurements: length 
2.5-2.8 ; width 1.0-1.2 mm. 

Types. Holotype S (M.C.Z. No. 30,181) and 2 paratypes 
from Maffin Bay Neth. N. G., Aug. 1944 (Darlington), and 1 
additional paratype from same locality, Aug. 1944 (E. S. Ross, 
California Acad.). 

Measured specimens. The S holotype and 1 (M.C.Z.) para- 
type. 

Notes. Sufficiently compared with typical sublobatus in the 
preceding description. 

Tachys mastersi exul n. subsp. 

Description. With characters of fasciatus group as here de- 
fined. Head relatively smaller and elytra relatively wider and 
with wider margins than usual in group ; reddish brown, faintly 
iridescent or silky, front with distinct reticulate microsculpture. 
Head .69 and .71 width prothorax ; eyes moderately large and 
prominent (less so than in fasciatus), genae forming obtuse 
(not quite right) angles with sides of neck; antennae with mid- 
dle segments slightly more than 3X long as wide. Prothorax 
transversely subcordate ; width/length 1.59 and 1.58 ; base/apex 
1.32 and 1.30; base/head 1.22 and 1.19; sides arcuate for most 
of length, rather weakly (somewhat variably) sinuate toward 
base ; apex subtruncate or broadly emarginate, sometimes slightly 
lobed at middle ; base subtruncate, broadly and slightly lobed ; 
posterior angles usually slightly obtuse (nearly right), slightly 
blunted; basal transverse sulcus less impressed than usual but 
distinct and crenulate, usually with a distinct (but variable) 
longitudinal sulciform impression at middle ; area behind sulcus 
longitudinally rugulose. Elytra broader and with sides more 



DARLINGTON : CARABID BEETLES OF NEW GUINEA 421 

rounded than usual; E/P 1.48 and 1.50; margins unusually 
broad laterally ; 2 or 3 inner discal striae more or less impressed 
(1st most impressed) and 1 or 2 additional striae indicated es- 
pecially anteriorly on disc; anterior dorsal puncture about % 
from base on (position of) 4th stria. Inner wings fully de- 
veloped. Lower surface, legs, and secondary sexual characters 
normal ; $ with 2 segments each front tarsus rather widely dil- 
ated ; last ventral segment not distinctly pubescent in either sex. 
Measurements: length 2.6-3.0; width 1.2-1.3 mm. 

Ttjpes. Holotype $ (M.C.Z. No. 30,182) and 14 paratypes 
all from Sansapor (Vogelkop), Neth. N. G., Aug. 1944 (Darling- 
ton). 

Measured specimens. The £ holotype and 1 9 paratype. 

Notes. Tachys master si Sloane ranges (with some variation) 
in eastern Australia from part of the Cape York Peninsula to 
southern New South Wales. The present subspecies, although 
from the farthest end of New Guinea, resembles typical mastersi 
rather closely, differing from it in having slightly better defined 
basal prothoracic angles, slightly longer antennal segments, and 
in other minor ways. This species is represented in the main 
part of New Guinea by another, more distinct subspecies ; and 
there is another form in the Philippines (Leyte). I should add 
that I am not sure that all these forms really represent a single 
species. Their adequate study would require much more time 
than I can give them at present. But to treat them as subspecies 
now emphasizes their probable relationships in a useful fashion. 
All the forms occur among dead leave